Different areas of human non-primary auditory cortex are activated by sounds with spatial and nonspatial properties.

In humans, neuroimaging studies have identified the planum temporale to be particularly responsive to both spatial and nonspatial attributes of sound. However, a functional segregation of the planum temporale along these acoustic dimensions has not been firmly established. We evaluated this scheme in a factorial design using modulated sounds that generated a percept of motion (spatial) or frequency modulation (nonspatial). In addition, these sounds were presented in the context of a motion detection and a frequency-modulation detection task to investigate the cortical effects of directing attention to different perceptual attributes of the sound. Motion produced stronger activation in the medial part of the planum temporale and frequency-modulation produced stronger activation in the lateral part of the planum temporale, as well as an additional non-primary area lateral to Heschl's gyrus. These separate subregions are consistent with the notion of divergent processing streams for spatial and nonspatial auditory information. Activation in the superior parietal cortex, putatively involved in the spatial pathway, was dependent on the task of motion detection and not simply on the presence of acoustic cues for motion. This finding suggests that the listening task is an important determinant of how the processing stream is engaged.

Amplitude and frequency-modulated stimuli activate common regions of human auditory cortex.

Hall et al. (Hall et al., 2002, Cerebral Cortex 12:140-149) recently showed that pulsed frequency-modulated tones generate considerably higher activation than their unmodulated counterparts in non-primary auditory regions immediately posterior and lateral to Heschl's gyrus (HG). Here, we use fMRI to explore the type of modulation necessary to evoke such differential activation. Carrier signals were a single tone and a harmonic-complex tone, with a 300 Hz fundamental, that were modulated at a rate of 5 Hz either in frequency, or in amplitude, to create six stimulus conditions (unmodulated, FM, AM). Relative to the silent baseline, the modulated tones, in particular, activated widespread regions of the auditory cortex bilaterally along the supra-temporal plane. When compared with the unmodulated tones, both AM and FM tones generated significantly greater activation in lateral HG and the planum temporale, replicating the previous findings. These activation patterns were largely overlapping, indicating a common sensitivity to both AM and FM. Direct comparisons between AM and FM revealed a higher magnitude of activation in response to the variation in amplitude than in frequency, plus a small part of the posterolateral region in the right hemisphere whose response was specifically AM-, and not FM-, dependent. The dominant pattern of activation was that of co-localized activation by AM and FM, which is consistent with a common neural code for AM and FM within these brain regions.

The sound-level-dependent growth in the extent of fMRI activation in Heschl's gyrus is different for low- and high-frequency tones.

fMRI (functional magnetic resonance imaging) was used to investigate whether the growth in activation of the human auditory cortex, with increasing sound level, is discernibly different for high- and low-frequency tones. Ten volunteers were scanned whilst listening to sequences of low-frequency (0.30-kHz) tones at sound levels between 42 and 96 dB sound pressure level (SPL), and 10 whilst listening to high-frequency (4.75-kHz) tones at the same sound levels. Activation was measured in Heschl's gyrus (including primary auditory cortex) which has been shown to be most sensitive to changes in sound level. For the 0.30-kHz tone, the extent of activation was flat up to 66 dB and then showed a rapid growth which continued up to the highest level studied (96 dB SPL). In contrast, increasing the level of 4.75-kHz tones produced a steady growth in the extent of activation across the range of levels studied. These results are consistent with physiological evidence suggesting that recruitment of primary auditory cortical neurones may be different at high and low frequencies.

Relationships between human auditory cortical structure and function.

The human auditory cortex comprises multiple areas, largely distributed across the supratemporal plane, but the precise number and configuration of auditory areas and their functional significance have not yet been clearly established. In this paper, we discuss recent research concerning architectonic and functional organisation within the human auditory cortex, as well as architectonic and neurophysiological studies in non-human species, which can provide a broad conceptual framework for interpreting functional specialisation in humans. We review the pattern in human auditory cortex of the functional responses to various acoustic cues, such as frequency, pitch, sound level, temporal variation, motion and spatial location, and we discuss their correspondence to what is known about the organisation of the auditory cortex in other primates. There is some neuroimaging evidence of multiple tonotopically organised fields in humans and of functional specialisations of the fields in the processing of different sound features. It is thought that the primary area, on Heschl's gyrus, may have a larger involvement in processing basic sound features, such as frequency and level, and that posterior non-primary areas on the planum temporale may play a larger role in processing more spectrotemporally complex sounds. Ways in which current knowledge of auditory cortical organisation and different data analysis approaches may benefit future functional neuroimaging studies which seek to link auditory cortical structure and function are discussed.

Heschl's gyrus is more sensitive to tone level than non-primary auditory cortex.

Previous neuroimaging studies generally demonstrate a growth in the cortical response with an increase in sound level. However, the details of the shape and topographic location of such growth remain largely unknown. One limiting methodological factor has been the relatively sparse sampling of sound intensities. Additionally, most studies have either analysed the entire auditory cortex without differentiating primary and non-primary regions or have limited their analyses to Heschl's gyrus (HG). Here, we characterise the pattern of responses to a 300-Hz tone presented in 6-dB steps from 42 to 96 dB sound pressure level as a function of its sound level, within three anatomically defined auditory areas; the primary area, on HG, and two non-primary areas, consisting of a small area lateral to the axis of HG (the anterior lateral area, ALA) and the posterior part of auditory cortex (the planum temporale, PT). Extent and magnitude of auditory activation increased non-linearly with sound level. In HG, the extent and magnitude were more sensitive to increasing level than in ALA and PT. Thus, HG appears to have a larger involvement in sound-level processing than does ALA or PT.