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Combination of binaural information across frequency bands.

Perceptual grouping of the frequency components from a source into a single auditory object is needed when localizing a complex sound in an environment where other sounds are also present. Two acoustic regularities that might allow for such grouping are a harmonic relation among the components and a commonality of their spatial positions. The utility of these cues was examined in a forced choice psychophysical task by measuring sensitivity to interaural differences of time (IDT) for low-frequency stimuli presented via earphones. In the first experiment, stimuli were composed of either one, two, or three frequencies. A signal detection analysis used to predict the effects of combining information across frequencies found summation to be optimal, regardless of the harmonicity of the complex. A second experiment presented two-frequency complexes in which one tone, the target, contained the IDT to be detected while the other, the distractor, was constant across all three intervals of the forced choice. For inharmonic complexes, performance for the target-distractor combinations was equivalent to that found for targets presented alone, suggesting segregation of the targets and distractors into separate auditory objects. However, for harmonic target-distractor combinations, performance was diminished. A signal detection analysis of these data supports the idea that for purposes of lateralization, the interaural information in the targets and distractors was combined into a variance-weighted value, even though it meant a lowering of performance. Thus it seems that for the grouping of complex acoustic stimuli in space, harmonic structure is more important than commonality of spatial position.

Attention↗

Audio-visual targeting reaction after unilateral lesions of the superior colliculus in cats.

Cats were trained to visually localize one of eight loudspeakers distributed in different spatial locations. The animals localized the activated sound source through a directed movement of ears, eyes and head towards it. The number of correct targeting reactions triggered by a tone stimulus from different spatial positions was strongly impaired in the operated animals in comparison to controls. The number of correct responses was higher when the tone stimuIus was presented from the side of the lesion. The role of the superior colliculus in the integration of audio-visual targeting reaction is discussed.

Animals↗

Representation of interaural time delay in the human auditory midbrain.

Interaural time difference (ITD) is a critical cue to sound-source localization. Traditional models assume that sounds leading at one ear, and perceived on that side, are processed in the opposite midbrain. Using functional magnetic resonance imaging we demonstrate that as the ITDs of sounds increase, midbrain activity can switch sides, even though perceived location remains on the same side. The data require a new model for human ITD processing.

Acoustic Stimulation↗

[Changes of evoked potentials caused by sound signals with different localization characteristics].

Characteristics of the mismatch negativity (MMN) were studied by presenting the subjects with four blocks of stimuli containing standard series of clicks (90%) simulating a stationery sound image located in the head midline, and one of three different deviant series of clicks (10%) simulating either a stationary sound image located near the left ear or a moving sound image which shifted from the head midline to the left ear or in the opposite direction. All the deviant stimuli elicited the MMN with the minimal peak amplitude and the greatest latency evoked by the deviant series of clicks simulating the sound image moving from the head midline to the left ear. These findings suggest that the MMN may be considered as a pre-perceptual physiological measure of the discrimination accuracy for the sound signals with various spatial locations.

Acoustic Stimulation↗

What and where in human audition: selective deficits following focal hemispheric lesions.

A sound that we hear in a natural setting allows us to identify the sound source and localize it in space. The two aspects can be disrupted independently as shown in a study of 15 patients with focal right-hemispheric lesions. Four patients were normal in sound recognition but severely impaired in sound localization, whereas three other patients had difficulties in recognizing sounds but localized them well. The lesions involved the inferior parietal and frontal cortices, and the superior temporal gyrus in patients with selective sound localization deficit; and the temporal pole and anterior part of the fusiform, inferior and middle temporal gyri in patients with selective recognition deficit. These results suggest separate cortical processing pathways for auditory recognition and localization.

Adult↗

Sound orientation behavior in cats. I. Localization of broadband noise.

Behavioral experiments measured the accuracy of the cat's voluntary head orientation responses to bursts of broadband noise at 104 locations in the frontal sound field. Cats were presented sound stimuli at randomly selected locations and received a food reward for releasing a lever when a light-emitting diode (LED) flashed at the same location. Head movements to the perceived location of the acoustic stimulus, and therefore expected location of the LED cue, were tracked by an electromagnetic sensor. Orientation responses to single sound bursts were directed to within 5 degrees of the azimuth and elevation of sounds within 15 degrees of the immediate frontal field and did not change for burst durations of 40, 100, and 200 ms. An increasing underestimation of target location was observed as the sound stimulus moved to more lateral azimuths and higher elevations. The "undershoot" phenomenon was reduced by allowing subjects to track paired stimulus bursts that repeated from the same location. These effects of sound location on the accuracy of orientation responses are predicted by the availability of mid-frequency spectral cues for sound localization.

Animals↗

Role of GABAergic inhibition in the coding of interaural time differences of low-frequency sounds in the inferior colliculus.

A major cue for the localization of sound in space is the interaural time difference (ITD). We examined the role of inhibition in the shaping of ITD responses in the inferior colliculus (IC) by iontophoretically ejecting gamma-aminobutyric acid (GABA) antagonists and GABA itself using a multibarrel pipette. The GABA antagonists block inhibition, whereas the applied GABA provides a constant level of inhibition. The effects on ITD responses were evaluated before, during and after the application of the drugs. If GABA-mediated inhibition is involved in shaping ITD tuning in IC neurons, then applying additional amounts of this inhibitory transmitter should alter ITD tuning. Indeed, for almost all neurons tested, applying GABA reduced the firing rate and consequently sharpened ITD tuning. Conversely, blocking GABA-mediated inhibition increased the activity of IC neurons, often reduced the signal-to-noise ratio and often broadened ITD tuning. Blocking GABA could also alter the shape of the ITD function and shift its peak suggesting that the role of inhibition is multifaceted. These effects indicate that GABAergic inhibition at the level of the IC is important for ITD coding.

Acoustic Stimulation↗

Maturation of glycinergic inhibition in the gerbil medial superior olive after hearing onset.

The neurones of the medial superior olive (MSO) are the most temporally sensitive neurones in the brain. They respond to the arrival time difference of sound at the two ears with a microsecond resolution; these interaural time differences are used to localize low-frequency sounds. In addition to the excitatory inputs from each ear, the MSO neurones also receive binaural glycinergic projections, which have a critical role in sound localization processing. Recently, it was shown that the glycinergic input to the MSO undergoes an experience-dependent structural reorganization after hearing onset. To explore the maturation of inhibition during the development of sound localization on a cellular level, glycinergic currents and potentials were measured in gerbil MSO principal cells from postnatal (P) day P12-P25 by whole-cell patch-clamp recordings. The synaptic glycinergic currents accelerated to rapid decay kinetics (approximately 2 ms) and rise times (approximately 0.4 ms) after hearing onset, reaching maturity around P17. Since the kinetics of miniature glycinergic currents did not change with age, it is likely that a higher degree of transmitter release synchrony is the underlying mechanism influencing the acceleration of the kinetics. During the same period, the synaptic glycinergic potentials accelerated four-fold, largely as a result of a prominent decrease in input resistance. In accordance with a reorganization of the glycinergic inputs, the evoked peak conductances decreased more than two-fold, together with a three-fold reduction in the frequency of miniature events after hearing onset. These age-dependent changes were absent in animals that had been reared in omni-directional noise, indicating that an experience-dependent pruning of synaptic inputs is important for the maturation of functional inhibition in the MSO. Taken together, these striking developmental adjustments of the glycinergic inhibition in the MSO most probably reflect an adaptation to improve the encoding of auditory cues with great temporal precision and fidelity during the maturation of sound localization behaviour.

Animals↗

Changes in evoked potentials during the action of sound signals with different localizing characteristics.

This report presents results of studies of the phenomenon of mismatch negativity (MMN) during exposure to four blocks of sound stimuli each containing identical standards creating an immobile sound image located along the midline of the head and one of a set of deviants, creating a sound image located either by the left ear or moving from the midline of the head towards the left ear or in the opposite direction. All deviants induced mismatch negativity; the minimal amplitude and longest latent period were seen in the mismatch negativity produced by the deviant modeling movement of the sound image from the midline of the head to the left ear. The question of the appearance of mismatch negativity as a criterion for the accurate discrimination of signals with different localizing characteristics is discussed.

Acoustic Stimulation↗