Functional connection between posterior superior temporal gyrus and ventrolateral prefrontal cortex in human.

The connection between auditory fields of the temporal lobe and prefrontal cortex has been well characterized in nonhuman primates. Little is known of temporofrontal connectivity in humans, however, due largely to the fact that invasive experimental approaches used so successfully to trace anatomical pathways in laboratory animals cannot be used in humans. Instead, we used a functional tract-tracing method in 12 neurosurgical patients with multicontact electrode arrays chronically implanted over the left (n = 7) or right (n = 5) perisylvian temporal auditory cortex (area PLST) and the ventrolateral prefrontal cortex (VLPFC) of the inferior frontal gyrus (IFG) for diagnosis and treatment of medically intractable epilepsy. Area PLST was identified by the distribution of average auditory-evoked potentials obtained in response to simple and complex sounds. The same sounds evoked little if there is any activity in VLPFC. A single bipolar electrical pulse (0.2 ms, charge-balanced) applied between contacts within physiologically identified PLST resulted in polyphasic evoked potentials clustered in VLPFC, with greatest activation being in pars triangularis of the IFG. The average peak latency of the earliest negative deflection of the evoked potential on VLPFC was 13.48 ms (range: 9.0-18.5 ms), providing evidence for a rapidly conducting pathway between area PLST and VLPFC.

Auditory-visual processing represented in the human superior temporal gyrus.

In natural face-to-face communication, speech perception utilizes both auditory and visual information. We described previously an acoustically responsive area on the posterior lateral surface of the superior temporal gyrus (field PLST) that is distinguishable on physiological grounds from other auditory fields located within the superior temporal plane. Considering the empirical findings in humans and non-human primates of cortical locations responsive to heard sounds and/or seen sound-sources, we reasoned that area PLST would also contain neural signals reflecting audiovisual speech interactions. To test this hypothesis, event related potentials (ERPs) were recorded from area PLST using chronically implanted multi-contact subdural surface-recording electrodes in patient-subjects undergoing diagnosis and treatment of medically intractable epilepsy, and cortical ERP maps were acquired during five contrasting auditory, visual and bimodal speech conditions. Stimulus conditions included consonant-vowel (CV) syllable sounds alone, silent seen speech or CV sounds paired with a female face articulating matched or mismatched syllables. Data were analyzed using a MANOVA framework, with the results from planned comparisons used to construct cortical significance maps. Our findings indicate that evoked responses recorded from area PLST to auditory speech stimuli are influenced significantly by the addition of visual images of the moving lower face and lips, either articulating the audible syllable or carrying out a meaningless (gurning) motion. The area of cortex exhibiting this audiovisual influence was demonstrably greater in the speech-dominant hemisphere.

Auditory cortical spatial receptive fields.

Neurons in the primary auditory cortex (AI) of anesthetized cats were studied for their sensitivity to directions of transient sounds in virtual acoustic space under a variety of conditions. An effective transient sound evokes a single spike or short burst of spikes with a precisely timed onset. The aggregate of effective directions forms a spatial receptive field. Typically, spatial receptive fields are large, often occupying a quadrant or more of acoustic space. Within the receptive field onset latency varies systematically with direction thereby providing information about source direction. This receptive field structure is highly robust, remaining relatively stable under conditions of competing sounds. Maximum likelihood analysis suggests that psychophysical spatial acuity can be achieved with a relatively small ensemble of AI neurons with broad receptive fields having response gradients of latency. Using reverse correlation and white-noise analysis receptive fields were mapped in space and time. This analysis revealed that spatial receptive fields of AI neurons need not be static but may exhibit marked temporal dynamics. This suggests a sensitivity for direction and speed of moving sound sources.

Auditory space-time receptive field dynamics revealed by spherical white-noise analysis.

Numerous studies have investigated the spatial sensitivity of cat auditory cortical neurons, but possible dynamic properties of the spatial receptive fields have been largely ignored. Given the considerable amount of evidence that implicates the primary auditory field in the neural pathways responsible for the perception of sound source location, a logical extension to earlier observations of spectrotemporal receptive fields, which characterize the dynamics of frequency tuning, is a description that uses sound source direction, rather than sound frequency, to examine the evolution of spatial tuning over time. The object of this study was to describe auditory space-time receptive field dynamics using a new method based on cross-correlational techniques and white-noise analysis in spherical auditory space. This resulted in a characterization of auditory receptive fields in two spherical dimensions of space (azimuth and elevation) plus a third dimension of time. Further analysis has revealed that spatial receptive fields of neurons in auditory cortex, like those in the visual system, are not static but can exhibit marked temporal dynamics. This might result, for example, in a neuron becoming selective for the direction and speed of moving auditory sound sources. Our results show that approximately 14% of AI neurons evidence significant space-time interaction (inseparability).

[Studies of directional sensitivity of neurons in the cat primary auditory cortex]. / Issledovanie prostranstvennoi chuvstvitel'nosti neironov pervichnoi slukhovoi kory koshki.

A set of impulsive transient signals has been synthesized for earphone delivery whose waveform and amplitude spectra, measured at the eardrum, mimic those of sounds arriving from a free-field source. The complete stimulus set forms a "virtual acoustic space" (VAS) for the cat. VAS stimuli are delivered via calibrated earphones sealed into the external meatus in cats under barbiturate anesthesia. Neurons recorded extracellularly in primary (AI) auditory cortex exhibit sensitivity to the direction of sound in VAS. The aggregation of effective sound directions forms a virtual space receptive field (VSRF). At about 20 dB above minimal threshold, VSRFs recorded in otherwise quiet and anechoic space fall into categories based on spatial dimension and location. The size, shape and location of VSRFs remain stable over many hours of recording and are found to be shaped by excitatory and inhibitory interactions of activity arriving from the two ears. Within the VSRF response latency and strength vary systematically with stimulus direction. In an ensemble of such neurons these functional gradients provide information about stimulus direction, which closely accounts for a human listener's spatial acuity. Raising stimulus intensity, introducing continuous background noise or presenting a conditioning stimulus all influence the extent of the VSRF but leave intact the gradient structure of the field. These and other findings suggest that such functional gradients in VSRFs of ensembles of AI neurons are instrumental in coding sound direction and robust enough to overcome interference from competing environmental sounds.

Directional sensitivity of neurons in the primary auditory (AI) cortex of the cat to successive sounds ordered in time and space.

Two transient sounds, considered as a conditioner followed by a probe, were delivered successively from the same or different direction in virtual acoustic space (VAS) while recording from single neurons in primary auditory cortex (AI) of cats under general anesthesia. Typically, the response to the probe sound was progressively suppressed as the interval between the two sounds (ISI) was systematically reduced from 400 to 50 ms, and the sound-source directions were within the cell's virtual space receptive field (VSRF). Suppression of the cell's discharge could be accompanied by an increase in response latency. In some neurons, the joint response to two sounds delivered successively was summative or facilitative at ISIs below about 20 ms. These relationships held throughout the VSRF, including those directions on or near the cell's acoustic axis where sounds often elicit the strongest response. The strength of suppression varied systematically with the direction of the probe sound when the ISI was fixed and the conditioning sound arrived from the cell's acoustic axis. Consequently a VSRF defined by the response to the lagging probe sound was progressively reduced in size when ISIs were shortened from 400 to 50 ms. Although the presence of a previous sound reduced the size of the VSRF, for many of these VSRFs a systematic gradient of response latency was maintained. The maintenance of such a gradient may provide a mechanism by which directional acuity remains intact in an acoustic environment containing competing acoustic transients.

Auditory cortex on the human posterior superior temporal gyrus.

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.

Spatial receptive fields of primary auditory cortical neurons in quiet and in the presence of continuous background noise.

Spatial receptive fields of primary auditory (AI) neurons were studied by delivering, binaurally, synthesized virtual-space signals via earphones to cats under barbiturate anesthesia. Signals were broadband or narrowband transients presented in quiet anechoic space or in acoustic space filled with uncorrelated continuous broadband noise. In the absence of background noise, AI virtual space receptive fields (VSRFs) are typically large, representing a quadrant or more of acoustic space. Within the receptive field, onset latency and firing strength form functional gradients. We hypothesized earlier that functional gradients in the receptive field provide information about sound-source direction. Previous studies indicated that spatial gradients could remain relatively constant across changes in signal intensity. In the current experiments we tested the hypothesis that directional sensitivity to a transient signal, as reflected in the gradient structure of VSRFs of AI neurons, is also retained in the presence of a continuous background noise. When background noise was introduced three major affects on VSRFs were observed. 1) The size of the VSRF was reduced, accompanied by a reduction of firing strength and lengthening of response latency for signals at an acoustic axis and on-lines of constant azimuth and elevation passing through the acoustic axis. These effects were monotonically related to the intensity of the background noise over a noise intensity range of approximately 30 dB. 2) The noise intensity-dependent changes in VSRFs were mirrored by the changes that occurred when the signal intensity was changed in signal-alone conditions. Thus adding background noise was equivalent to a shift in the threshold of a directional signal, and this shift was seen across the spatial receptive field. 3) The spatial gradients of response strength and latency remained evident over the range of background noise intensity that reduced spike count and lengthened onset latency. Those gradients along the azimuth that spanned the frontal midline tended to remain constant in slope and position in the face of increasing intensity of background noise. These findings are consistent with our hypothesis that, under background noise conditions, information that underlies directional acuity and accuracy is retained within the spatial receptive fields of an ensemble of AI neurons.

Modeling of auditory spatial receptive fields with spherical approximation functions.

A spherical approximation technique is presented that affords a mathematical characterization of a virtual space receptive field (VSRF) based on first-spike latency in the auditory cortex of cat. Parameterizing directional sensitivity in this fashion is much akin to the use of difference-of-Gaussian (DOG) functions for modeling neural responses in visual cortex. Artificial neural networks and approximation techniques typically have been applied to problems conforming to a multidimensional Cartesian input space. The problem with using classical planar Gaussians is that radial symmetry and consistency on the plane actually translate into directionally dependent distortion on spherical surfaces. An alternative set of spherical basis functions, the von Mises basis function (VMBF), is used to eliminate spherical approximation distortion. Unlike the Fourier transform or spherical harmonic expansions, the VMBFs are nonorthogonal, and hence require some form of gradient-descent search for optimal estimation of parameters in the modeling of the VSRF. The optimization equations required to solve this problem are presented. Three descriptive classes of VSRF (contralateral, frontal, and ipsilateral) approximations are investigated, together with an examination of the residual error after parameter optimization. The use of the analytic receptive field model in computational models of population coding of sound direction is discussed, together with the importance of quantifying receptive field gradients. Because spatial hearing is by its very nature three dimensional or, more precisely, two dimensional (directional) on the sphere, we find that spatial receptive field models are best developed on the sphere.

The structure of spatial receptive fields of neurons in primary auditory cortex of the cat.

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.

Applications of least-squares FIR filters to virtual acoustic space.

A virtual acoustic space (VAS) employs the localization cues specified by the direction-dependent 'free-field to eardrum transfer function' (FETF) to synthesize sound-pressure waveforms present near the tympanum. The combination of a VAS and the earphone delivery of synthesized waveforms is useful to study parametrically the neural mechanisms of directional hearing. The VAS-earphone procedure requires accurate FETF estimation from free-field measurements and appropriate compensation for the undesirable spectral characteristics of the closed-field earphone sound delivery and measurement systems. Here we describe how specially designed finite-impulse-response (FIR) filters improve these two operations. The coefficients of an FIR filter are determined using a least-squares error criterion. The least-squares FIR filter is implemented entirely in the time domain and avoids the usual problems with division inherent in a frequency domain approach. The estimation of an FETF by a least-squares FIR filter is veracious since its impulse response can recover signals that were recorded near the eardrum in the free field with a very high fidelity. The correlation coefficient between recorded and recovered time waveforms typically exceeds 0.999. Similarly, least-squares FIR filters prove excellent in compensating closed-field sound systems since comparisons of waveforms delivered by a compensated earphone to their corresponding predistorted signals yield correlation coefficients that exceed 0.99 on average.

Simulation of free-field sound sources and its application to studies of cortical mechanisms of sound localization in the cat.

We synthesized a set of signals (clicks) for earphone delivery whose waveforms and amplitude spectra, measured at the eardrum, mimic those of sounds arriving from a free-field source. The complete stimulus set represents 1816 sound-source directions, which together surround the head to form a 'virtual acoustic space' for the cat. Virtual-space stimuli were delivered via calibrated earphones sealed into the external meatus in cats under barbiturate anesthesia. Neurons recorded in AI cortex exhibited sensitivity to the direction of sound in virtual acoustic space. The aggregation of effective sound directions formed a virtual space receptive field (VSRF). At 20 dB above minimal threshold, VSRFs fell into one of several categories based on spatial dimension and location. Most VSRFs were confined to either the contralateral (59%) or ipsilateral (10%) sound hemifield. Seven percent spanned the frontal quadrants and 16% were omnidirectional. Eight percent fit into no clear category and were termed 'complex'. The size, shape, and location of VSRFs remained stable over many hours of recording. The results are in essential agreement with free-field studies. VSRFs were found to be shaped by excitatory and inhibitory interactions of activity arriving from the two ears. Some cortical neurons were found to preserve the spectral information in the free-field sound which was generated by the acoustical properties of the head and pinna, filtered by the cochlea and transmitted by auditory nerve fibers.

Auditory cortical neurons are sensitive to static and continuously changing interaural phase cues.

1. The interaural-phase-difference (IPD) sensitivity of single neurons in the primary auditory (AI) cortex of the anesthetized cat was studied at stimulus frequencies ranging from 120 to 2,500 Hz. Best frequencies of the 43 AI cells sensitive to IPD ranged from 190 to 2,400 Hz. 2. A static IPD was produced when a pair of low-frequency tone bursts, differing from one another only in starting phase, were presented dichotically. The resulting IPD-sensitivity curves, which plot the number of discharges evoked by the binaural signal as a function of IPD, were deeply modulated circular functions. IPD functions were analyzed for their mean vector length (r) and mean interaural phase (phi). Phase sensitivity was relatively independent of best frequency (BF) but highly dependent on stimulus frequency. Regardless of BF or stimulus frequency within the excitatory response area the majority of cells fired maximally when the ipsilateral tone lagged the contralateral signal and fired least when this interaural-phase relationship was reversed. 3. Sensitivity to continuously changing IPD was studied by delivering to the two ears 3-s tones that differed slightly in frequency, resulting in a binaural beat. Approximately 26% of the cells that showed a sensitivity to static changes in IPD also showed a sensitivity to dynamically changing IPD created by this binaural tonal combination. The discharges were highly periodic and tightly synchronized to a particular phase of the binaural beat cycle. High synchrony can be attributed to the fact that cortical neurons typically respond to an excitatory stimulus with but a single spike that is often precisely timed to stimulus onset. A period histogram, binned on the binaural beat frequency (fb), produced an equivalent IPD-sensitivity function for dynamically changing interaural phase. For neurons sensitive to both static and continuously changing interaural phase there was good correspondence between their static (phi s) and dynamic (phi d) mean interaural phases. 4. All cells responding to a dynamically changing stimulus exhibited a linear relationship between mean interaural phase and beat frequency. Most cells responded equally well to binaural beats regardless of the initial direction of phase change. For a fixed duration stimulus, and at relatively low fb, the number of spikes evoked increased with increasing fb, reflecting the increasing number of effective stimulus cycles. At higher fb, AI neurons were unable to follow the rate at which the most effective phase repeated itself during the 3 s of stimulation.(ABSTRACT TRUNCATED AT 400 WORDS)

Sensitivity of auditory cortical neurons of kittens to monaural and binaural high frequency sound.

The experiments reported here describe the abilities of young auditory cortical neurons to encode information about tone bursts having frequencies above 2.5 kHz. The studies were carried out in anesthetized kittens ranging from 8 to 44 days of age. At all ages studied, stimulation of the contralateral ear was most effective in evoking spikes. Typically the response was confined to stimulus onset. Thresholds were comparatively high and response latencies were comparatively long in the youngest kittens studied. The time course of threshold development was very similar to that of the auditory nerve and cochlear nuclei indicating that most, if not all, age related thresholds and threshold changes at the cortical level are accounted for by mechanisms operating at the level of the cochlea and auditory nerve. Response latency shortened progressively over the first month of postnatal life and while the absolute change in response latency differed considerably from that of cells in the cochlear nuclei the proportional changes were very similar. These data indicate that the comparatively long response latency and latency changes recorded at the cortex are imposed by underdeveloped central auditory processes. Response areas of kitten cortical neurons resembled those of the adult. At all ages studied, binaural interactions were robust and similar in kind to those recorded in adult cats. We conclude that cortical neurons of kittens preserve the results of interactions occurring at lower brainstem levels and that the development of the circuits of which these neurons are a part develop as a functional unit.

Maps of auditory cortex in cats reared after unilateral cochlear ablation in the neonatal period.

The responses of many neurons recorded in the high best-frequency region of primary auditory cortical field, AI, of the normal adult cat depend upon intensity differences of the sounds arriving at the two ears. These binaural interactions are exhibited early in postnatal life, well before structural maturation of the auditory pathways from the ear to the cortex is complete. The aim of the present work was to study certain aspects of the functional development of the auditory cortex in adult cats unilaterally deaf from birth. In adult animals reared with a neonatal cochlear ablation, field AI ipsilateral to the non-operated ear showed a normal tonotopic map, which was derived from single neurons and neuron clusters driven securely by best-frequency tonal stimulation in virtually every electrode penetration. The acoustic thresholds at many recording sites were as low as those obtained in AI contralateral to the non-operated ear. These findings are in marked contrast to those from control experiments on normal adult cats where only about 65% of AI neurons were excited by a sound delivered to the ipsilateral ear and where thresholds to ipsilateral ear stimulation were significantly higher than contralateral thresholds. The spatial distribution of cortical neurons based on acoustic thresholds also appeared to be different in cats unilaterally deaf from birth when compared to control cats. Closely spaced electrode penetrations in AI ipsilateral to the non-operated ear suggested that neurons were separated into low-threshold regions and high-threshold regions. There was no evidence for this type of non-random segregation in control experiments.

Topography of binaural organization in primary auditory cortex of the cat: effects of changing interaural intensity.

Responses from neuron clusters were used to derive binaural and aural dominance maps within the 5- to 30-kHz frequency representation of the primary auditory cortical (AI) field in the barbiturate-anesthetized cat. Tone burst stimuli were presented dichotically using a calibrated and sealed acoustic delivery system to parametrically vary interaural intensity difference (IID). Neuron cluster responses were divided into three binaural interaction classes using audiovisual criteria: summation (56%), suppression (25%), and mixed (17%). Neurons in the summation and suppression classes demonstrated a single type of binaural interaction, regardless of intensity manipulations. Neurons in the mixed binaural class demonstrated summation responses when dichotic tonal intensities were near their threshold levels and the IID was small, but suppression responses when the IID was increased. The relative proportions of the three binaural interaction classes changed with distance along the dorsal-to-ventral isofrequency dimension. Nearly equal proportions of each class were observed at the ventral end of field AI, whereas quite different proportions of each class were seen at the dorsal extreme of the field. The average frequency of occurrence of the mixed binaural class increased nearly monotonically with increasing distance from the dorsal end of field AI. The majority of mapped AI loci exhibited a contralateral aural dominance (65%) with equidominance (25%), ipsilateral aural dominance (6%), and predominantly binaural (4%) classes accounting for the remainder. Average topographic distributions of aural dominance suggested that the ventral end of field AI consisted almost exclusively of the contralateral dominance class, whereas more equal proportions of the four classes were observed near the dorsal extreme of the field. The highest average proportions of ipsilateral aural dominance and predominantly binaural classes were found in the dorsal half of field AI. Single neurons, isolated at cortical loci assigned to the mixed binaural class during the mapping of neuron clusters, were shown to demonstrate both summation and suppression responses. Quantitative measurements relating either discharge rate or response latency to changes in the IID appeared to distinguish these cells from other single neurons studied. Typically, the probability of discharge was initially increased and subsequently decreased by progressive changes in IID that increased the intensity of the ipsilateral tone relative to the contralateral tone. The initial changes in IID characteristically shortened the latent period to the binaural response while subsequent increments in IID produced a more comp

Geometry and orientation of neuronal processes in cat primary auditory cortex (AI) related to characteristic-frequency maps.

Microelectrode mapping and horseradish peroxidase oxidase histochemistry were combined to study the relationship between the characteristic-frequency representation and the intrinsic connectivity of the primary auditory cortex in the cat. Small extracellular iontophoretic injections of horseradish peroxidase within the characteristic-frequency map resulted in labeling of neuronal processes that, in the tangential plane, radiated out asymmetrically from the injection site over distances of several millimeters. The heaviest concentration of labeled fibers was along an axis parallel with the orientation of the isofrequency line within which the injection had been made. Thus, primary field neurons that have the same or a similar characteristic frequency have the potential of being preferentially interconnected.

Auditory cortical field projections to the basal ganglia of the cat.

Projections to the basal ganglia from four auditory cortical fields in the cat were studied by combining microelectrode-mapping of the neurons' best frequencies with autoradiographic and histochemical tract-tracing techniques. Each auditory field is a source of projections to the homolateral basal ganglia. The distribution of labeling within the basal ganglia is related to the cortical field in which the injection site is located. The dorsal portion of the putamen and adjacent caudate nucleus are connected with cortical fields situated anteriorly and dorsally, while the ventral portion of the putamen and adjacent lateral amygdaloid nucleus are related to auditory fields situated posteriorly and ventrally. Injections of two different tracers into different best-frequency loci of one cortical field provided evidence that low best-frequency neurons project medially within the basal ganglia while high best-frequency neurons project more laterally. We concluded that there was a basic similarity among patterns of terminations in the basal ganglia from axons that originate in different auditory cortical fields. When the source of a projection was confined to a restricted portion of an auditory cortical field, labeling appeared as dense patches of silver grains separated from each other by areas of less dense labeling. Often, these patches were distributed within a sheet of tissue, elongated both dorsoventrally and anteroposteriorly. Loci having the same best-frequency representation, but situated in different auditory cortical fields, project upon overlapping but not coextensive portions of a single sheet of tissue. Thus the projections from geographically distant cortical loci possessing similar best-frequency representations are notably distinguished on a topographic basis. By comparison, two adjacent sheets of tissue were labeled when two injections were made into the low best-frequency and high best-frequency representations of the same auditory field. Double-injection, double-tracer experiments revealed that adjacent sheets of tissue received projections from different best-frequency loci. These observations suggested a degree of tonotopic organization to this projection system which was equipoise to the evidence obtained for a topographic organization.

Ipsilateral corticocortical projections related to binaural columns in cat primary auditory cortex.

Projections from the high-frequency representations of auditory cortical fields A and P were autoradiographically labeled using anterograde transport of a mixture of tritiated proline and leucine. A single injection of isotope into either cortical field resulted in multiple regions of dense labeling within ipsilateral AI. As the result of an injection of radioactive label into field A in one experiment, densely labeled regions were found to be systematically related to the binaural map throughout the high-frequency representation of AI. Contralateral dominant suppression responses were located in regions of dense labeling while summation responses were located in regions of less dense labeling. In contrast, it was previously found that callosal axon terminations were more densely concentrated in summation columns than in contralateral dominant suppression columns (Imig and Brugge, '78). Thus, the two classes of binaural columns differ with respect to the density of innervation they receive from these ipsilateral and contralateral populations of neurons. In other experiments in which isotope was injected into field A, a systematic relationship between density of ipsilateral labeling and binaural response class was only seen in a portion of AI; in other regions no relationship was evident. A simple interpretation of these data follows. Within a contiguous territory in field A is a population of neurons whose axons provide more dense innervation to contralateral dominant suppression columns than to summation columns in ipsilateral AI. Injections of radioactivity confined to this territory result in a systematic relationship between the density of labeling and the binaural map throughout AI. Outside this territory is a population of neurons whose axon terminations are not systematically related to the binaural map in AI. Isotope injections which engage both territories may result in a systematic relation between density of labeling and the binaural map in one portion of AI, while in another, no relationship may be evident. There is some indication that projections from field P may also be related to binaural columns in AI in the same manner as are the projections from field A.

Tonotopic organization in auditory cortex of the cat.

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.