Temporal encoding of phonetic features in auditory cortex.
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We studied the effects of sound presentation rate and attention on auditory supratemporal cortex (STC) activation in 12 healthy adults using functional magnetic resonance imaging (fMRI) at 3 T. The sounds (200 ms in duration) were presented at steady rates of 0.5, 1, 1.5, 2.5, or 4 Hz while subjects either had to focus their attention to the sounds or ignore the sounds and attend to visual stimuli presented with a mean rate of 1 Hz. Consistent with previous observations, we found that both increase in stimulation rate and attention to sounds enhanced activity in STC bilaterally. Further, we observed larger attention effects with higher stimulation rates. This interaction of attention and presentation rate has not been reported previously. In conclusion, our results show both rate-dependent and attention-related modulations of STC indicating that both factors should be controlled, or at least addressed, in fMRI studies of auditory processing.
The responses of neuronal clusters to amplitude-modulated tones were studied in five auditory cortical fields of the anesthetized cat: the primary auditory field (AI), second auditory field (AII), anterior auditory field (AAF), posterior auditory field (PAF) and the ventro-posterior auditory field (VPAF). Modulation transfer functions (MTFs) for amplitude-modulated tones were obtained at 172 cortical locations. MTFs were constructed by measuring firing rate (rate-MTFs) and response synchronization (synchronization-MTFs) to sinusoidal and rectangular waveform modulation of CF-tones. The MTFs were characterized by their 'best-modulation frequency' (BMF) and a measure of their quality of 'sharpness' (Q2dB). These characteristics were compared for the five fields. Rate and synchronization MTFs for sinusoidal and rectangular modulation produced similar estimates of BMF and Q2dB. Comparison of averaged BMFs between the cortical fields revealed relatively high BMFs in AAF (mean: 31.1 Hz for synchronization to sinusoidal AM) and moderately high BMFs in AI (14.2 Hz) whereas BMFs encountered in AII, VPAF and PAF were generally low (7.0, 5.2, and 6.8 Hz). The MTFs were relatively broadly tuned (low Q2dB) in AAF and sharper in a low modulation group containing AII, PAF and VPAF. The ventro-posterior field was the most sensitive to changes in the modulation waveform. We conclude that there are significant differences between auditory cortical fields with respect to their temporal response characteristics and that the assessment of these response characteristics reveals important aspects of the functional significance of auditory cortical fields for the coding and representation of complex sounds.
Functional Magnetic Resonance Imaging (fMRI) holds exciting potential as a research and clinical tool for exploring the human auditory system. This noninvasive technique allows the measurement of discrete changes in cerebral cortical blood flow in response to sensory stimuli, allowing determination of precise neuroanatomical locations of the underlying brain parenchymal activity. Application of fMRI in auditory research, however, has been limited. One problem is that fMRI utilizing echo-planar imaging technology (EPI) generates intense noise that could potentially affect the results of auditory experiments. Also, issues relating to the reliability of fMRI for listeners with normal hearing need to be resolved before this technique can be used to study listeners with hearing loss. This preliminary study examines the feasibility of using fMRI in auditory research by performing a simple set of experiments to test the reliability of scanning parameters that use a high resolution and high signal-to-noise ratio unlike that presently reported in the literature. We used consonant-vowel (CV) speech stimuli to investigate whether or not we could observe reproducible and consistent changes in cortical blood flow in listeners during a single scanning session, across more than one scanning session, and in more than one listener. In addition, we wanted to determine if there were differences between CV speech and nonspeech complex stimuli across listeners. Our study shows reproducibility within and across listeners for CV speech stimuli. Results were reproducible for CV speech stimuli within fMRI scanning sessions for 5 out of 9 listeners and were reproducible for 6 out of 8 listeners across fMRI scanning sessions. Results of nonspeech complex stimuli across listeners showed activity in 4 out of 9 individuals tested.
Hierarchical processing suggests that spectrally and temporally complex stimuli will evoke more activation than do simple stimuli, particularly in non-primary auditory fields. This hypothesis was tested using two tones, a single frequency tone and a harmonic tone, that were either static or frequency modulated to create four stimuli. We interpret the location of differences in activation by drawing comparisons between fMRI and human cytoarchitectonic data, reported in the same brain space. Harmonic tones produced more activation than single tones in right Heschl's gyrus (HG) and bilaterally in the lateral supratemporal plane (STP). Activation was also greater to frequency-modulated tones than to static tones in these areas, plus in left HG and bilaterally in an anterolateral part of the STP and the superior temporal sulcus. An elevated response magnitude to both frequency-modulated tones was found in the lateral portion of the primary area, and putatively in three surrounding non-primary regions on the lateral STP (one anterior and two posterior to HG). A focal site on the posterolateral STP showed an especially high response to the frequency-modulated harmonic tone. Our data highlight the involvement of both primary and lateral non-primary auditory regions.
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Histological methods and 2-deoxyglucose autoradiography were used in an attempt at finding distinguishing characteristics that would permit the clear definition of different auditory areas on the ectosylvian gyrus. This region was studied in both coronal and flattened tangential sections. In tangential sections a crescent-shaped region of high deoxyglucose uptake was identified. The centre of this crescent was in the position of the primary auditory area on the middle ectosylvian gyrus. The ventro-anterior arm of the crescent was on the surface of the anterior ectosylvian gyrus and the ventro-posterior arm on the posterior ectosylvian gyrus. All three parts of the crescent appear to have an auditory function, because ablating the inferior colliculus or inserting a contralateral earplug reduced their deoxyglucose uptake. This was shown by using two separately distinguishable forms of 2-deoxyglucose, incorporating the 18F and 14C isotopes. In addition, another area of high deoxyglucose activity was identified in the ventral wall of the suprasylvian sulcus, which seems to correspond to the anterior auditory field. These four areas with high deoxyglucose uptake also have high levels of succinate dehydrogenase activity and moderately high densities of myelinated fibres. Succinate dehydrogenase histochemistry provides a simple method for identifying auditory cortical areas and should be of use in future physiological studies. These results provide evidence that the ferret has four separate auditory areas with relatively high metabolic and functional activity.
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