Microelectrode mapping techniques were employed in the cat's auditory cortex to relate the best frequencies of a large population of neurons with their spatial loci. Based upon the best-frequency distribution, the auditory region was divided into four complete and orderly tonotopic representations and a surrounding belt of cortex in which the tonotopic organization was more complex. The four auditory fields occupy a crescent-shaped band of tissue which comprises portions of both the exposed gyral surfaces and sulcal banks of the ectosylvian cortex. The anterior auditory field (A) is situated most rostrally upon the anterior ectosylvian gyrus. It extends upon the ventral bank of the suprasylvian sulcus and upon the banks of the anterior ectosylvian sulcus. Adjoining field A caudally is the primary auditory field (AI), which extends across the middle ectosylvian gyrus and portions of both banks of the posterior ectosylvian sulcus. The representations of the highest best frequencies in fields A and AI are contiguous. Caudal and ventral to AI are located the posterior (P) and ventroposterior (VP) auditory fields. They lie mainly upon the caudal bank of the posterior ectosylvian sulcus but also extend upon the rostral bank and upon the posterior ectosylvian gyrus. The low best-frequency representations of fields AI and P are contiguous, whereas the low best-frequency representation of field VP lies near the ventral end of the posterior ectosylvian sulcus. Fields P and VP are joined along their middle and high best-frequency representations. Within each auditory field isofrequency lines defined by the spatial loci of neurons with similar best frequencies are oriented orthogonal to the low-to-high best-frequency gradients.

The human superior temporal cortex plays a critical role in hearing, speech, and language, yet its functional organization is poorly understood. Evoked potentials (EPs) to auditory click‐train stimulation presented binaurally were recorded chronically from penetrating electrodes implanted in Heschl's gyrus (HG), from pial‐surface electrodes placed on the lateral superior temporal gyrus (STG), or from both simultaneously, in awake humans undergoing surgery for medically intractable epilepsy. The distribution of averaged EPs was restricted to a relatively small area on the lateral surface of the posterior STG. In several cases, there were multiple foci of high amplitude EPs lying along this acoustically active portion of STG. EPs recorded simultaneously from HG and STG differed in their sensitivities to general anesthesia and to changes in rate of stimulus presentation. Results indicate that the acoustically active region on the STG is a separate auditory area, functionally distinct from the HG auditory field(s). We refer to this acoustically sensitive area of the STG as the posterior lateral superior temporal area (PLST). Electrical stimulation of HG resulted in short‐latency EPs in an area that overlaps PLST, indicating that PLST receives a corticocortical input, either directly or indirectly, from HG. These physiological findings are in accord with anatomic evidence in humans and in nonhuman primates that the superior temporal cortex contains multiple interconnected auditory areas. J. Comp. Neurol. 416:79–92, 2000. © 2000 Wiley‐Liss, Inc.

Speech comprehension relies on temporal cues contained in the speech envelope, and the auditory cortex has been implicated as playing a critical role in encoding this temporal information. We investigated auditory cortical responses to speech stimuli in subjects undergoing invasive electrophysiological monitoring for pharmacologically refractory epilepsy. Recordings were made from multicontact electrodes implanted in Heschl's gyrus (HG). Speech sentences, time compressed from 0.75 to 0.20 of natural speaking rate, elicited average evoked potentials (AEPs) and increases in event-related band power (ERBP) of cortical high-frequency (70 -250 Hz) activity. Cortex of posteromedial HG, the presumed core of human auditory cortex, represented the envelope of speech stimuli in the AEP and ERBP. Envelope following in ERBP, but not in AEP, was evident in both language-dominant and -nondominant hemispheres for relatively high degrees of compression where speech was not comprehensible. Compared to posteromedial HG, responses from anterolateral HG-an auditory belt field-exhibited longer latencies, lower amplitudes, and little or no time locking to the speech envelope. The ability of the core auditory cortex to follow the temporal speech envelope over a wide range of speaking rates leads us to conclude that such capacity in itself is not a limiting factor for speech comprehension.

The capacity of auditory cortex on Heschl's gyrus (HG) to encode repetitive transients was studied in human patients undergoing surgical evaluation for medically intractable epilepsy. Multicontact depth electrodes were chronically implanted in gray matter of HG. Bilaterally presented stimuli were click trains varying in rate from 4 to 200 Hz. Averaged evoked potentials (AEPs) and event-related band power (ERBP), computed from responses at each of 14 recording sites, identified two auditory fields. A core field, which occupies posteromedial HG, was characterized by a robust polyphasic AEP on which could be superimposed a frequency following response (FFR). The FFR was prominent at click rates below approximately 50 Hz, decreased rapidly as click rate was increased, but could reliably be detected at click rates as high as 200 Hz. These data are strikingly similar to those obtained by others in the monkey under essentially the same stimulus conditions, indicating that mechanisms underlying temporal processing in the auditory core may be highly conserved across primate species. ERBP, which reflects increases or decreases of both phase-locked and non-phase-locked power within given frequency bands, showed stimulus-related increases in gamma band frequencies as high as 250 Hz. The AEPs recorded in a belt field anterolateral to the core were typically of low amplitude, showing little or no evidence of short-latency waves or an FFR, even at the lowest click rates used. The non-phase-locked component of the response extracted from the ERBP showed a robust, long-latency response occurring here in response to the highest click rates in the series.

Transient broad-band stimuli that mimic in their spectrum and time waveform sounds arriving from a speaker in free space were delivered to the tympanic membranes of barbiturized cats via sealed and calibrated earphones. The full array of such signals constitutes a virtual acoustic space (VAS). The extra-cellular response to a single stimulus at each VAS direction, consisting of one or a few precisely time-locked spikes, was recorded from neurons in primary auditory cortex. Effective sound directions form a virtual space receptive field (VSRF). Near threshold, most VSRFs were confined to one quadrant of acoustic space and were located on or near the acoustic axis. Generally, VSRFs expanded monotonically with increases in stimulus intensity, with some occupying essentially all of the acoustic space. The VSRF was not homogeneous with respect to spike timing or firing strength. Typically, onset latency varied by as much as 4-5 msec across the VSRF. A substantial proportion of recorded cells exhibited a gradient of first-spike latency within the VSRF. Shortest latencies occupied a core of the VSRF, on or near the acoustic axis, with longer latency being represented progressively at directions more distant from the core. Remaining cells had VSRFs that exhibited no such gradient. The distribution of firing probability was mapped in those experiments in which multiple trials were carried out at each direction. For some cells there was a positive correlation between latency and firing probability.

Functional connections between auditory fields on Heschl's gyrus (HG) and the acoustically responsive posterior lateral superior temporal gyrus (field PLST) were studied using electrical stimulation and recording methods in patients undergoing diagnosis and treatment of intractable epilepsy. Averaged auditory (click-train) evoked potentials were recorded from multicontact subdural recording arrays chronically implanted over the lateral surface of the superior temporal gyrus (STG) and from modified depth electrodes inserted into HG. Biphasic electrical pulses (bipolar, constant current, 0.2 ms) were delivered to HG sites while recording from the electrode array over acoustically responsive STG cortex. Stimulation of sites along the mediolateral extent of HG resulted in complex waveforms distributed over posterolateral STG. These areas overlapped each other and field PLST. For any given HG stimulus site, the morphology of the electrically evoked waveform varied across the STG map. A characteristic waveform was recorded at the site of maximal amplitude of response to stimulation of mesial HG [presumed primary auditory field (AI)]. Latency measurements suggest that the earliest evoked wave resulted from activation of connections within the cortex. Waveforms changed with changes in rate of electrical HG stimulation or with shifts in the HG stimulus site. Data suggest widespread convergence and divergence of input from HG to posterior STG. Evidence is presented for a reciprocal functional projection, from posterolateral STG to HG. Results indicate that in humans there is a processing stream from AI on mesial HG to an associational auditory field (PLST) on the lateral surface of the superior temporal gyrus.

No abstract

Evidence regarding the functional subdivisions of human auditory cortex has been slow to converge on a definite model. In part, this reflects inadequacies of current understanding of how the cortex represents temporal information in acoustic signals. To address this, we investigated spatiotemporal properties of auditory responses in human posterolateral superior temporal (PLST) gyrus to acoustic click-train stimuli using intracranial recordings from neurosurgical patients. Subjects were patients undergoing chronic invasive monitoring for refractory epilepsy. The subjects listened passively to acoustic click-train stimuli of varying durations (160 or 1,000 ms) and rates (4-200 Hz), delivered diotically via insert earphones. Multicontact subdural grids placed over the perisylvian cortex recorded intracranial electrocorticographic responses from PLST and surrounding areas. Analyses focused on averaged evoked potentials (AEPs) and high gamma (70-150 Hz) event-related band power (ERBP). Responses to click trains featured prominent AEP waveforms and increases in ERBP. The magnitude of AEPs and ERBP typically increased with click rate. Superimposed on the AEPs were frequency-following responses (FFRs), most prominent at 50-Hz click rates but still detectable at stimulus rates up to 200 Hz. Loci with the largest high gamma responses on PLST were often different from those sites that exhibited the strongest FFRs. The data indicate that responses of non-core auditory cortex of PLST represent temporal stimulus features in multiple ways. These include an isomorphic representation of periodicity (as measured by the FFR), a representation based on increases in non-phase-locked activity (as measured by high gamma ERBP), and spatially distributed patterns of activity.