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J C Middlebrooks

Publications and source records attributed to J C Middlebrooks.

29 records · Page 2Linked to original sources

Narrow-band sound localization related to external ear acoustics.

Human subjects localized brief 1/6-oct bandpassed noise bursts that were centered at 6, 8, 10, and 12 kHz. All testing was done under binaural conditions. The horizontal component of subjects' responses was accurate, comparable to that for broadband localization, but the vertical and front/back components exhibited systematic errors. Specifically, responses tended to cluster within restricted ranges that were specific for each center frequency. The directional transfer functions of the subjects' external ears were measured for 360 horizontal and vertical locations. The spectra of the sounds that were present in the subjects' ear canals, the "proximal stimulus" spectra, were computed by combining the spectra of the narrow-band sound sources with the directional transfer functions for particular stimulus locations. Subjects consistently localized sounds to regions within which the associated directional transfer function correlated most closely with the proximal stimulus spectrum. A quantitative model was constructed that successfully predicted subjects' responses based on interaural level difference and spectral cues. A test of the model, using techniques adapted from signal detection theory, indicated that subjects tend to use interaural level difference and spectral shape cues independently, limited only by a slight spatial correlation of the two cues. A testing procedure is described that provides a quantitative comparison of various predictive models of sound localization.

Acoustics↗

Sound localization by human listeners.

In keeping with our promise earlier in this review, we summarize here the process by which we believe spatial cues are used for localizing a sound source in a free-field listening situation. We believe it entails two parallel processes: 1. The azimuth of the source is determined using differences in interaural time or interaural intensity, whichever is present. Wightman and colleagues (1989) believe the low-frequency temporal information is dominant if both are present. 2. The elevation of the source is determined from spectral shape cues. The received sound spectrum, as modified by the pinna, is in effect compared with a stored set of directional transfer functions. These are actually the spectra of a nearly flat source heard at various elevations. The elevation that corresponds to the best-matching transfer function is selected as the locus of the sound. Pinnae are similar enough between people that certain general rules (e.g. Blauert's boosted bands or Butler's covert peaks) can describe this process. Head motion is probably not a critical part of the localization process, except in cases where time permits a very detailed assessment of location, in which case one tries to localize the source by turning the head toward the putative location. Sound localization is only moderately more precise when the listener points directly toward the source. The process is not analogous to localizing a visual source on the fovea of the retina. Thus, head motion provides only a moderate increase in localization accuracy. Finally, current evidence does not support the view that auditory motion perception is anything more than detection of changes in static location over time.

Attention↗

Directional dependence of interaural envelope delays.

Interaural envelope delays were measured in six human subjects as a function of the location of a movable sound source, bandpassed between 3 and 16 kHz. A total of 324 source locations were tested in horizontal and vertical increments of 10 degrees. A method is described for estimating the complex directional transfer function of the external ear, independent of the position of the recording microphone in the ear canal. To compute interaural envelope delays, directional transfer functions from the left and right ears were convolved with a critical-band filter, the envelopes were computed, and the envelopes were cross correlated. Interaural envelope delays, as well as interaural group delays, varied somewhat with the center frequency of the critical-band filter and with the vertical location of the sound source. Nevertheless, to a first approximation, envelope delays measured in the ear canals increased monotonically with increasing angle of incidence relative to the median plane, as they would for two microphones on the surface of a rigid sphere. The results are discussed in relation to the possible contribution of interaural envelope delays to sound localization behavior.

Attention↗

Two-dimensional sound localization by human listeners.

This study measured the ability of subjects to localize broadband sound sources that varied in both horizontal and vertical location. Brief (150 ms) sounds were presented in a free field, and subjects reported the apparent stimulus location by turning to face the sound source; head orientation was measured electromagnetically. Localization of continuous sounds also was tested to estimate errors in the motor act of orienting with the head. Localization performance was excellent for brief sounds presented in front of the subject. The smallest errors, averaged across subjects, were about 2 degrees and 3.5 degrees in the horizontal and vertical dimensions, respectively. The sizes of errors increased, for more peripheral stimulus locations, to maxima of about 20 degrees. Localization performance was better in the horizontal than in the vertical dimension for stimuli located on or near the frontal midline, but the opposite was true for most stimuli located further peripheral. Front/back confusions occurred in 6% of trials; the characteristics of those responses suggest that subjects derived horizontal localization information principally from interaural difference cues. The generally high level of performance obtained with the head orientation technique argues for its utility in continuing studies of sound localization.

Adult↗

Directional sensitivity of sound-pressure levels in the human ear canal.

Changes in sound pressures measured in the ear canal are reported for broadband sound sources positioned at various locations about the subject. These location-dependent pressures are one source of acoustical cues for sound localization by human listeners. Sound source locations were tested with horizontal and vertical resolution of 10 degrees. Sound levels were measured with miniature microphones placed inside the two ear canals. Although the measured amplitude spectra varied with the position of the microphone in the ear canal, it is shown that the directional sensitivity at any particular frequency of the broadband stimulus is independent of microphone position anywhere within the ear canal. At any given frequency, the distribution of sound pressures as a function of sound source location formed a characteristic spatial pattern comprising one or two discrete areas from which sound sources produced maximum levels in the ear canal. The locations of these discrete areas varied in horizontal and vertical location according to sound frequency. For example, around 8 kHz, two areas of maximum sensitivity typically were found that were located laterally and were separated from each other vertically, whereas, around 12 kHz, two such areas were found located on the horizontal plane and separated horizontally. The spatial patterns of sound levels were remarkably similar among different subjects, although some frequency scaling was required to accommodate for differences in the subjects' physical sizes. Interaural differences in sound-pressure level (ILDs) at frequencies below about 8 kHz tended to increase monotonically with increasing distance of the sound source from the frontal midline and tended to be relatively constant as a function of vertical source location. At higher frequencies, however, ILDs varied both with the horizontal and with the vertical location of the sound source. At some frequencies, asymmetries between the left and right ears in a given subject resulted in substantial ILDs even for midline sound sources. These results indicate the types of horizontal and vertical spatial information that are available from sound level cues over various ranges of frequency and, within a small subject population, indicate the nature of intersubject variability.

Acoustic Stimulation↗

Changes in external ear position modify the spatial tuning of auditory units in the cat's superior colliculus.

This study examines the influence of external ear position on the auditory spatial tuning of single units in the superior colliculus of the anesthetized cat. Unit responses to broad-band stimuli presented in a free sound field were measured with the external ears in a forward symmetrical position or with one or the other ear turned 40 degrees to the side; the ears are referred to as contra- or ipsilateral with respect to the side of the recording site. Changes in the position of either ear modified the spatial tuning of units. The region of space from which a stimulus was most effective in activating a unit is referred to as the unit's "best area". Whenever the contralateral ear was turned to the side, best areas shifted peripherally and somewhat upward, roughly in proportion to the magnitude of the change in ear position. A turn of the ipsilateral ear to the side had more variable effects, but best areas generally shifted frontally. Best areas located between approximately 10 and 40 degrees contralateral when the ears were forward were least affected by changes in ipsilateral ear position. Changes in ear position also modified the maximum response rates of many units. Units with best areas located within approximately 20 degrees of the frontal midline when the ears were forward exhibited a pronounced decrease in responsiveness when either ear was turned. Units with more peripheral best areas tended to show no change or a slight increase in responsiveness. The influence of ear position on the directionality of the external ears was determined by mapping the cochlear microphonic response to tones or one-third-octave bands of noise before and after turning the ear. When the ears were forward, maximum interaural intensity differences (IIDs) were produced by high-frequency sound sources (greater than or equal to 20 kHz) located 20-40 degrees from the frontal midline and by lower frequency sources located further peripherally. The influence of ear position on the locations from which maximum IIDs were produced was similar to the influence of ear position on unit best areas. Changes in ipsilateral ear position had different effects on high- and low-frequency IIDs that were comparable with the effects of changes in ear position on frontally and peripherally located best areas, respectively.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustics↗

Binaural mechanisms of spatial tuning in the cat's superior colliculus distinguished using monaural occlusion.

This study explores the mechanisms of auditory spatial tuning in the superior colliculus of the anesthetized cat by correlating spatial tuning within specific regions of space with particular types of binaural interaction. The auditory spatial tuning of units was measured using a movable, broad-band stimulus presented in a free sound field. The contribution of each ear to the response of a unit was identified by acutely plugging one or the other ear. Every unit became largely or entirely unresponsive when a foam-rubber earplug was placed in the ear contralateral to the recording site. Thus, every unit exhibited an excitatory or facilitatory influence from the contralateral ear. A plug placed in the ipsilateral ear had different effects on different units. For half of the units (16/32), an ipsilateral earplug produced increases in the sizes of the units' receptive fields and increases in the magnitudes of their responses to stimuli presented from most locations. Thus, these units exhibited inhibition from the ipsilateral ear. Another class of units (9/32) exhibited ipsilateral facilitation, in that an ipsilateral earplug caused decreases in the sizes of the units' receptive fields and prominent decreases in their response magnitudes. For the remaining units (7/32), an ipsilateral earplug resulted in decreases in the sizes of the units' receptive fields, but produced both decreases in the responses of units to stimuli presented in their best areas and increases in the responses to stimuli presented away from the best areas. Thus these units exhibited mixed facilitatory and inhibitory ipsilateral influences. The influence of an ipsilateral earplug on a unit's response tended to correlate with its spatial tuning. The region of space within which a sound source was most effective in activating a unit was its "best area". The best areas of units exhibiting ipsilateral inhibition were located furthest peripherally, those of units showing ipsilateral facilitation were located furthest frontally, and the best areas of units showing mixed ipsilateral influences were located in an intermediate area. The frequency tuning of units measured using a free-field tone source also tended to correlate with the locations of their best areas. Half of the units tested (27/54) responded to tones of the sound pressure levels (SPLs) that were used (up to 50 dB SPL).(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

A neural code for auditory space in the cat's superior colliculus.

We have examined the intermediate and deep layers of the cat's superior colliculus for evidence of a neural representation of auditory space. We measured the responses of single units to sounds presented in a free field. The results support the following generalizations. Most auditory units in the superior colliculus have sharply delimited receptive fields which form two discrete classes distinguished by their locations and sizes. The remaining units respond to sounds presented at any location. Each auditory unit responds maximally to sounds at a particular horizontal and vertical location within its receptive field, the unit's "best area." The best areas and receptive field borders of a unit are resistant to changes in the intensity of stimulus. The locations of best areas shift systematically as a function of unit position to form a continuous map of auditory space. The horizontal dimension of space is mapped rostrocaudally, and the vertical dimension is mapped mediolaterally. This map corresponds in orientation with the map of visual space. These data permit us to infer the distribution of unit activity elicited by a sound at any given location. Regardless of its location, a sound activates a substantial portion of the superior colliculus. Indeed, sounds at some locations activate nearly all of the auditory units. However the activated portion of the colliculus contains a restricted region of units which are excited to near their maximum firing rates. The position of this focus of greatest activity varies systematically according to the location of the sound source, thus mapping the location of the sound in space.

Acoustic Stimulation↗

Intrinsic organization of the cat's medial geniculate body identified by projections to binaural response-specific bands in the primary auditory cortex.

The area of the cat's primary auditory cortex (AI) within which high frequency sounds are represented can be subdivided using functional criteria. Within each subdivision, or "binaural interaction band," all recorded neurons display similar responses to binaural stimulation. The current study distinguishes the thalamic sources of input to these subdivisions of AI and characterizes the topography within the thalamic projection to each class of bands. The borders of binaural bands in AI were mapped using microelectrode recording with diotic tonal stimulation, then injections of one to three retrograde tracers were introduced into identified bands. Within the ventral division (V) of the medial geniculate body (the major thalamic source of input to AI), the neuronal populations that projected to different classes of binaural bands were strictly segregated from each other. This segregation of class-specific thalamic sources constitutes a laminar organization within an axis of V that is orthogonal to the previously described tonotopic organization. Excitatory/excitatory (EE) binaural neurons in AI were found to be segregated from excitatory/inhibitory (EI) neurons in alternating "bands." We consistently identified: (1) a ventral pair of rostrocaudally continuous EI and EE bands, (2) a middle area within which the pattern of binaural subdivisions was more variable and within which bands often were discontinuous rostrocaudally, and (3) a dorsal zone (DZ) within which the responses of neurons differed in binaural properties and in frequency specificity from the response patterns that were characteristic of neurons elsewhere in AI. Each EI band apparently derived input that converged from three thickened laminae of cells in V that were oriented approximately horizontally. The most ventral of these laminae encompassed the ovoidal part of V (Vo), suggesting that EI bands are the only recipients in AI of a projection from Vo. All of the EE bands and DZ derived their input from a single continuous structure which included the dorsal two-thirds of the rostral pole of V and a horizontal lamina interposed between the two dorsalmost EI-projecting laminae. Restricted portions of the complex EE-projecting structure in V projected preferentially to particular EE subdivisions of AI. The V-to-AI thalamocortical topography exhibited a high degree of convergence and divergence within the projections to each cortical binaural band and within the projections to each class of bands. These observations indicate that the high frequency representation in AI and its principal thalamic source of input, the ventral division of the medial geniculate body, may be thought of as assemblies of spatially discrete, functionally distinguishable subunits. The significance of this intrinsic organization is discussed in regard to the requirements for analysis of sound stimuli.

Animals↗

Binaural response-specific bands in primary auditory cortex (AI) of the cat: topographical organization orthogonal to isofrequency contours.

The spatial distribution of neurons with different binaural response properties has been studied within the three dimensions of the primary auditory cortex (AI) in the cat. Using dichotic stimulation, 92% of neurons encountered could be classified into either the excitatory/excitatory (EE) or excitatory/inhibitory (EI) interaction class. In nearly all of almost 800 penetrations introduced along radial axes, all neurons encountered along a given penetration were of the same binaural response class. Neurons of different binaural interaction classes were spatially segregated within the plane of the cortex. Electrode penetrations made parallel to isofrequency contours traversed the mediolateral extent of AI through the middle layers of the cortex. A sharp segregation of units by binaural response class was observed in these penetrations, i.e. sequences of neurons that were all of the EE class alternated with sequences of EI neurons. The regions of uniform response to binaural stimulation formed radially organized topographical subunits that were elongated along the rostrocaudal dimension of AI. These binaural interaction bands intersect the lines of re-representation of the cochlear sensory epithelium ('isofrequency contours') and, thus, create subdivisions of AI that each contain a representation of the entire audible frequency domain. The implications of these results for the concept of AI as a unitary element in auditory processing are discussed.

Acoustic Stimulation↗