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Unit study of monkey frontal cortex: active localization of auditory and of visual stimuli.

The influence of sound localization behavior on unit activity in the frontal cortex of awake rhesus monkeys was examined by comparing responses under three behavioral conditions: auditory localization, during which a response was required to the location of a sound (broad-band noise) source; auditory detect, during which a response was required to indicate the occurrence of the sound regardless of location; visual localization, during which no sounds were presented and a response was required to the location of a visual stimulus; and nonperform, presentation of auditory stimuli as in the first two conditions, but with the animal sitting passively. Extracellular microelectrode recordings were made in the periarcuate region and dorsal and ventral prefrontal areas near the principal sulcus. Four monkeys were used with a total of 498 cells studied. Of the total population, only five cells were found to have characteristics similar to those of auditory units in the primary auditory cortex and the surrounding belt area. More typically, units were found that had strong short-latency responses specific to the auditory and/or visual localization tasks. These units had no or weak responses when the same sound stimuli were presented in the auditory detect task or when a monkey received the sound stimuli in a nonperforming condition. Two regions were identified, one medial and/or posterior to the arcuate sulcus, in Brodmann's area 6; the second included parts of areas 8 and 9 within the genu of the arcuate sulcus. Units from these regions are referred to, respectively, as the postarcuate and the prearcuate populations. Both populations responded predominantly during active localization behavior. Sixty-two percent of the postarcuate population responded during auditory localization, 32% responded during auditory detect, and only 18% responded to acoustic stimuli presented in the nonperforming condition. In the prearcuate population percentages in these three conditions were 35, 25, and 12%, respectively. For visual localization, 54% in the postarcuate population responded, whereas 42% in the prearcuate responded. Spatial tuning of units during auditory localization was similar to that seen in units of the primary auditory cortex, with the greatest percentages of units responding to stimuli contralateral to the recording site. Similar tuning was observed for the visual localization task as well. Similarities in spatial tuning between the auditory and visual localization conditions were examined to assess the "bimodal" nature of the units.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals

Localization of tones by horses: use of binaural cues and the role of the superior olivary complex.

The ability of horses to use binaural time and intensity difference cues to localize sound was assessed in free-field localization tests by using pure tones. The animals were required to discriminate the locus of a single tone pip ranging in frequency from 250 Hz to 25 kHz emitted by loudspeakers located 30 degrees to the left and right of the animals' midline (60 degrees total separation). Three animals were tested with a two-choice procedure; 2 additional animals were tested with a conditioned avoidance procedure. All 5 animals were able to localize 250 Hz, 500 Hz, and 1 kHz but were completely unable to localize 2 kHz and above. Because the frequency of ambiguity for the binaural phase cue delta phi for horses in this test was calculated to be 1.5 kHz, these results indicate that horses can use binaural time differences in the form of delta phi but are unable to use binaural intensity differences. This finding was supported by an unconditioned orientation test involving 4 additional horses, which showed that horses correctly orient to a 500-Hz tone pip but not to an 8-kHz tone pip. Analysis of the superior olivary complex, the brain stem nucleus at which binaural interactions first take place, reveals that the lateral superior olive (LSO) is relatively small in the horse and lacks the laminar arrangement of bipolar cells characteristic of the LSO of most mammals that can use binaural delta I.

Animals

The auditory midbrain of a marsupial: the brush-tailed possum (Trichosurus vulpecula).

A microelectrode survey was made of the midbrain auditory nuclei of the brushtailed possum (Trichosurus vulpecula), a common Australian marsupial. Information was sought on the tuning characteristics of individual neurones, tonotopic organization and mechanisms of sound localization. It was felt that such information would be of use in future studies of the development and evolution of mammalian hearing. Twelve possums were anaesthetized with ketamine and chloralose-urethane, and recordings were made of extracellular unit discharges in the inferior colliculus during monaural and binaural tonal stimulation. The inferior colliculus of the possum consists of a central nucleus - a darkly stained, densely packed group of cells - flanked laterally by an external nucleus with a lower density of paler cells. Tonotopic organization was demonstrated by discretelytuned elements in the central nucleus, but was not observed in the external nucleus. In the latter region broad and irregular tuning was commonly seen. Most units in both divisions were influenced by binaural stimuli, with patterns of binaural interaction similar to those observed in the cat inferior colliculus. Cells influenced by changes in the interaural time and intensity difference were commonly observed, but only a subclass of these were suited in sensitivity for sound localization. In general, the midbrain auditory system of the possum was similar in unit discharge characteristics and organization to those of the eutherian mammals commonly studied.

Acoustic Stimulation

Role of spectral cues in median plane localization.

The role of spectral cues in the sound source to ear transfer function in median plane sound localization is investigated in this paper. At first, transfer functions were measured and analyzed. Then, these transfer functions were used in experiments where sounds from a source on the median plane were simulated and presented to subjects through headphones. In these simulation experiments, the transfer functions were smoothed by ARMA models with different degrees of simplification to investigate the role of microscopic and macroscopic patterns in the transfer functions for median plane localization. The results of the study are summarized as follows: (1) For front-rear judgment, information derived from microscopic peaks and dips in the low-frequency region (below 2 kHz) and the macroscopic patterns in the high-frequency region seems to be utilized; (2) for judgment of elevation angle, major cues exist in the high-frequency region above 5 kHz. The information in macroscopic patterns is utilized instead of that in small peaks and dips.

Adult

Rapid acquisition of an auditory localization discrimination by rats.

Acquisition of a sound localization discrimination by rats was investigated. Two loudspeakers were located outside an experimental enclosure containing two levers and a dipper feeder. In the same-side condition, responses on the lever nearest the sound-producing speaker were reinforced. Animals in this condition acquired the discrimination rapidly, generally within the first session. In the opposite-side condition, responses on the lever furthest from the sound-producing speaker were reinforced. Acquisition for animals in this condition began below the chance level (50% correct responses) and took on the order of 10 sessions to approach the final, high level. The course of acquisition in both cases appeared to depend upon an initial tendency of rats to respond on the lever nearest the source of sound in this situation. The rise-decay time of the 4-kHz tone burst signal clearly affected the performance level reached. It did not, however, affect the rate at which the discrimination was acquired.

Animals

Mismatch negativity to change in spatial location of an auditory stimulus.

Auditory stimulus blocks were presented to 12 reading subjects. Each block consisted of 2 types, standard (P = 90%) and deviant stimuli (P = 10%), delivered in a random order. The only difference between these stimuli was their spatial location of origin. The subject always heard the standards as coming straight in front and the deviants from an angle of either 10, 45, or 90 degrees to the right of the standards. The spatial locations were produced via earphones by introducing for low-frequency (600 Hz) tones an interaural phase difference and for high-frequency (3000 Hz) tones an interaural intensity difference. Standard and deviant stimuli were also delivered in more natural, free-field, conditions via differently positioned loudspeakers. The deviant tones elicited an event-related brain potential component called the mismatch negativity (MMN), followed by a P3a component. Thus changes in spatial location of an auditory stimulus produced by following either one of the two main principles of human sound localization elicited the MMN. Consequently, it was concluded that the spatial location of a sound source is coded in the hypothesized neuronal stimulus traces reflected by the MMN and, further, that a change in this location is automatically detected by the brain by means of the MMN generator process.

Adult

Auditory localization: effects of reflecting surfaces.

Based on anatomical and evolutionary conceptions of the human ear, an experiment was conducted in which forty-eight human subjects were asked to localize sounds (a human voice) emitted by one of twenty-seven stationary loudspeakers in an anechoic chamber. The position of the active loudspeaker varied with respect to azimuth, distance, and elevation in three steps each. The position of a single sound-reflecting surface (about 6 m2) was varied: on the floor, on the ceiling, to the left, and to the right. The accuracy of identifying the active loudspeaker for each position of the sound-reflecting surface was compared intraindividually with the absence of reflection. The results show an overall increase in correct localizations with a sound-reflecting surface on the floor. Especially the elevation of the sound source can be detected with greater precision. Additionally, the percentage of correct localizations decreased systematically with the presence of a sound-reflecting ceiling, while the presence of sound-reflecting walls did not systematically affect the localization performance. Judgments in the horizontal plane and those of distance were not systematically influenced by the presence of a sound-reflecting surface.

Adult

Azimuthal processing in the posterior auditory thalamus of cats.

The responses to free-field acoustic stimuli of 157 units in the auditory thalamus of anesthetized cats were studied in relation to the localization of pure tone stimuli in the azimuthal plane. Units were classified as 'directional' if their firing rates at sound levels in excess of 20 dB above threshold varied by more than 50% as a function of azimuth. Sixty-five % of the units in the nucleus of the brachium of the inferior colliculus and 30% in the ventral division of the medial geniculate body were found to be directional, suggesting different processing channels for sound localization between colliculus and cortex.

Acoustic Stimulation