Although the auditory cortex plays a necessary role in sound localization, physiological investigations in the cortex reveal inhomogeneous sampling of auditory space that is difficult to reconcile with localization behavior under the assumption of local spatial coding. Most neurons respond maximally to sounds located far to the left or right side, with few neurons tuned to the frontal midline. Paradoxically, psychophysical studies show optimal spatial acuity across the frontal midline. In this paper, we revisit the problem of inhomogeneous spatial sampling in three fields of cat auditory cortex. In each field, we confirm that neural responses tend to be greatest for lateral positions, but show the greatest modulation for near-midline source locations. Moreover, identification of source locations based on cortical responses shows sharp discrimination of left from right but relatively inaccurate discrimination of locations within each half of space. Motivated by these findings, we explore an opponent-process theory in which sound-source locations are represented by differences in the activity of two broadly tuned channels formed by contra- and ipsilaterally preferring neurons. Finally, we demonstrate a simple model, based on spike-count differences across cortical populations, that provides bias-free, level-invariant localization—and thus also a solution to the “binding problem” of associating spatial information with other nonspatial attributes of sounds.
We compared the spatial sensitivity of neural responses in three areas of cat auditory cortex: primary auditory cortex (A1), the posterior auditory field (PAF), and the dorsal zone (DZ). Stimuli were 80-ms pure tones or broadband noise bursts varying in free-field azimuth (in the horizontal plane) or elevation (in the vertical median plane), presented at levels 20-40 dB above units' thresholds. We recorded extracellular spike activity simultaneously from 16 to 32 sites in one or two areas of alpha-chloralose-anesthetized cats. We examined the dependence of spike counts and response latencies on stimulus location as well as the information transmission by neural spike patterns. Compared with units in A1, DZ units exhibited more complex frequency tuning, longer-latency responses, increased prevalence and degree of nonmonotonic rate-level functions, and weaker responses to noise than to tonal stimulation. DZ responses also showed sharper tuning for stimulus azimuth, stronger azimuthal modulation of first-spike latency, and enhanced spatial information transmission by spike patterns, compared with A1. Each of these findings was similar to differences observed between PAF and A1. Compared with PAF, DZ responses were of shorter overall latency, and more DZ units preferred stimulation from ipsilateral azimuths, but the majority of analyses suggest strong similarity between PAF and DZ responses. These results suggest that DZ and A1 are physiologically distinct cortical fields and that fields like PAF and DZ might constitute a "belt" region of auditory cortex exhibiting enhanced spatial sensitivity and temporal coding of stimulus features.
Normal listeners were tested for their temporal auditory gap detection thresholds using free-field presentation of white-noise stimuli delivered from the left (L) and right (R) poles of the interaural axis. The noise bursts serving as the leading and trailing markers for the silent period were presented in either the same (LL,RR) or different (LR,RL) auditory locations. The duration of the leading marker was a second independent variable. Gap thresholds for stimuli in which the markers had the same location were low, and usually were independent of the duration of the leading marker. Gap thresholds for the LR and RL conditions were longer. These gap thresholds were sensitive to the duration of the leading marker, and increased as the leading marker duration decreased. This finding is consistent with the hypothesis that a relative timing operation mediates gap detection when the markers activate different perceptual channels. The present data suggest that this timing process can operate on perceptual channels emerging from central nervous system processing.
We assessed the spatial-tuning properties of units in the cat's anterior auditory field (AAF) and compared them with those observed previously in the primary (A1) and posterior auditory fields (PAF). Multi-channel, silicon-substrate probes were used to record single-and multi-unit activity from the right hemispheres of α-chloralose-anesthetized cats. Spatial tuning was assessed using broadband noise bursts that varied in azimuth or elevation. Response latencies were slightly, though significantly shorter in AAF than A1, and considerably shorter in both of those fields than in PAF. Compared to PAF, spike counts and latencies were more poorly modulated by changes in stimulus location in AAF and A1, particularly at higher sound pressure levels. Moreover, units in AAF and A1 demonstrated poorer level tolerance than units in PAF with spike rates modulated as much by changes in stimulus intensity as changes in stimulus location. Finally, spike-pattern-recognition analyses indicated that units in AAF transmitted less spatial information, on average, than did units in PAF-an observation consistent with recent evidence that PAF is necessary for sound-localization behavior, whereas AAF is not.
Bilateral auditory cortex lesions in Japanese macaques result in an aphasia-like deficit in which the animals are unable to discriminate two forms of their coo vocalizations. To determine whether this deficit is sensory in nature, two monkeys with bilateral lesions were tested for their ability to discriminate frequency and frequency change. The results indicated that although the animals were able to discriminate between sounds of different frequencies, they were unable to determine whether a sound was changing in frequency. Because the animals' coo vocalizations differ primarily in the predominant direction of their frequency change and not in their absolute frequency content, the aphasia-like deficit of animals with bilateral auditory cortex lesions appears to be a sensory disorder.
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