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.

Passive eye displacement alters auditory spatial receptive fields of cat superior colliculus neurons.

The superior colliculus (SC) is thought to use a set of superimposed, topographically organized neural maps of visual, auditory, somatosensory and motor space to direct the eyes toward novel stimuli. Auditory spatial response fields (SRFs) of SC neurons may change when an animal moves its eyes, presumably to compensate for the resulting misalignment of visual and auditory sensory spatial reference frames, but the mechanisms responsible for these SRF changes remain unknown. Here we report that passive deviation of the eye in anesthetized, paralyzed animals can profoundly affect the auditory responsiveness of SC neurons, but seems insufficient by itself to provide adaptive shifts of auditory SRFs.

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.

Computerised infrared imaging system for studying thermal activation on the skull following somatic stimulation in small animals.

A computerised infrared imaging system has been developed to measure infrared radiation as a means of functionally mapping the cerebral cortex. In two species of small mammal, rat and gerbil, the authors localised the thermal changes at the skull overlying the somatic sensory cortex following somatic stimulation of the mystacial vibrissae. Though typically small in magnitude, a thermal response could be detected through the skull. To enhance detection sensitivity, a number of measures were taken to improve various aspects of data acquisition, stimulus delivery and control of experimental conditions. Regarding data analysis, a coordinate system based on skull landmarks was adopted to localise thermally-active regions for comparison across animals of the same species. To extract the region of weak temperature changes, a coarse-to-fine detection strategy was developed, which searched automatically for clusters of temporally- and spatially-correlated pixels above a data-driven threshold. Thus, the dynamic aspect of the thermal changes at any region of interest on the skull could be studied efficiently. The detection algorithm was tested against simulated responses in addition to empirical data obtained from animals. All of the above software was integrated in a user-friendly package.

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.

Thermal images of somatic sensory cortex obtained through the skull of rat and gerbil.

Infrared images of the skull surface were obtained in urethane-anesthetized rats and gerbils before, during and after mechanical stimulation of the face and mystacial vibrissae on one side. Areas of increased temperature on the skull, localized mainly over the face area of the primary somatosensory cortex contralateral to the side of stimulation, appeared within 4-5 s after the onset of stimulation. Rarely, such temperature change was recorded bilaterally. Temperatures did not remain high on the intact skull in rats, but fell to baseline within minutes after stimulus onset regardless of stimulus duration. In rats in which the skull had been thinned and in gerbils with intact skull, temperatures remained elevated during the course of stimulation. We were unable to resolve the activation of individual vibrissae.

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.

Sensitivity of auditory nerve fibers to spectral notches.

1. Listeners use direction-dependent spectral cues introduced by the torso, head, and pinnae to localize the source of a sound in space. Among the prominent direction-dependent spectral features in the free field-to-eardrum transfer function are narrow regions of low acoustic energy referred to as spectral notches. In this paper, we studied the sensitivity of single auditory nerve fibers in the barbiturate-anesthetized cat to broadband noise that had been filtered by a function whose shape approximated natural notches in the free field-to-eardrum transfer function. 2. Two experimental paradigms were employed. The first was the repeated presentation of a burst of broadband noise filtered by the simulated-notch function. Center frequency of the notch was held constant at or around the fiber characteristic frequency (CF). We refer to this as a "stationary" notch stimulus. The second paradigm was the repeated presentation of a broadband noise that was constructed from noise segments, each filtered by the simulated notch, whose CF was incremented and then decremented in a systematic way. We refer to this as a "moving" notch stimulus. Results from these two paradigms were compared with respect to notch detection. 3. Data were obtained from 161 single auditory nerve fibers having CFs ranging from 0.4 to 40 kHz. Most fibers studied had CFs > 5 kHz, and they detected the presence of the spectral notch in an intensity- and frequency-dependent manner. Each fiber responded vigorously to the presence of broadband noise. When the CF of the notch encroached on the response area of the fiber, there was a demonstrable reduction in discharge rate. The greatest reduction in discharge rate occurred when the notch was centered at the fiber's CF and when the level of the notch signal was some 15-55 dB above the fiber's noise threshold. There was close association in the frequency-intensity plane between the position of the most effective notch and the fiber's threshold tuning curve. 4. For high-spontaneous rate fibers, a moving-notch stimulus, but not a stationary one, reduced the discharge below the spontaneous rate at and in the immediate vicinity of the most effective notch frequency. This increases sensitivity to a spectral notch and suggests a mechanism by which localization ability is enhanced when there is relative motion between a sound source and the head.(ABSTRACT TRUNCATED AT 400 WORDS)

Virtual-space receptive fields of single auditory nerve fibers.

1. Sounds reaching the tympanic membranes are first modified by the acoustic properties of the torso, head, and external ear. For certain frequencies in the incident sound there results a complex, direction-dependent spatial distribution of sound pressure at the eardrum such that, within a sound field, localized areas of pressure maxima are flanked by areas of pressure minima. Listeners may use these spatial maxima and minima in localizing the source of a sound in space. The results presented describe how information about this spatial pressure pattern is transmitted from the cochlea to the central auditory system via single fibers of the auditory nerve. 2. Discharges of single fibers of the auditory nerve were studied in Nembutal-anesthetized cats [characteristic frequencies (CFs) ranged from 0.4 to 40 kHz]. Click stimuli were derived from sound-pressure waveforms that were generated by a loudspeaker placed at 1,800 locations around the cat's head and recorded at the tympanic membrane with miniature microphones. Recorded signals were converted to acoustic stimuli and delivered to the ear via a calibrated and sealed earphone. The full complement of signals is referred to as "virtual acoustic space," and the spatial distribution of discharges to this array of signals is referred to as a "virtual-space receptive field" (VSRF). 3. Fibers detect both pressure maxima and pressure minima in virtual acoustic space. Thus VSRFs take on complex shapes. 4. VSRFs of fibers of the same or similar CF having low spontaneous rates had the same overall pattern as those from high-spontaneous rate (HSR) fibers. For HSR fibers, the VSRF is obscured by the high background spike activity. 5. Comparison of the VSRF and isolevel contour maps of the stimulus derived at various frequencies revealed that auditory nerve fibers most accurately extract spectral information contained in the stimulus at a frequency close to or slightly higher than CF.

Encoding of amplitude-modulated tones by neurons of the inferior colliculus of the kitten.

Responses of single neurons of the central nucleus of the inferior colliculus (ICC) of kittens 4-43 days of age were studied using sinusoidally amplitude-modulated (AM) tones delivered monaurally or binaurally via sealed and calibrated earphones. The carrier frequency of the AM signal was set to the CF of the neuron. CFs ranged from 2-26 kHz. During the about first 2 weeks of postnatal life, ICC neurons responded to sound with periodic bursts of activity. In response to AM tones, discharges of ICC neurons at all ages studied were phase-locked to the envelope of the modulation waveform over a wide range of stimulus level and modulation depth. A linear relationship, independent of SPL, was found between the average phase of discharge on the modulation cycle and modulation frequency. The slope of the line represents a time delay, which was highly correlated with the first-spike latency to tone onset, and hence with the age of the animal. The mean effective phase of the discharge remained relatively constant with age. There was little systematic change in average phase of discharge with changing stimulus level or modulation depth, although the number of spikes evoked and the temporal pattern of the spikes within a modulation cycle could vary. The sensitivity function relating spike synchrony or spike count to modulation frequency was typically band-pass in nature. The most effective modulation frequency (MEMF) was, on average, 15 Hz, far below that reported for adult cat ICC cells. When AM tones were delivered binaurally, the discharge was a periodic function of the interaural phase difference of the stimulus envelopes. The results indicate that prior to the time the cochlea is able to respond to most environmental sounds, monaural and binaural circuits involving the ICC faithfully transmit information pertaining to amplitude-modulated signals in the rate and timing of their discharges. During the next several weeks, when neural thresholds fall to adult levels, ICC circuits are activated by amplitude modulated sounds at levels encountered in the normal acoustic environment even though they are restricted to modulation frequencies below those encoded by the adult.

Glutathione S-transferase isoenzymes in rat brain neurons and glia.

The glutathione S-transferases (GSTs) constitute a family of cytosolic isoenzymes and a structurally unrelated microsomal enzyme that is involved in the detoxication of electrophilic xenobiotics. These enzymes also participate in the intracellular binding and transport of a broad range of lipophilic compounds including bilirubin, and hormones such as the glucocorticoids and thyroid hormones. The present investigation demonstrates that GSTs are present in neurons of the brainstem, forebrain, and cerebellum. An isoenzyme-specific distribution of GSTs was found in cytoplasm, nuclei, and nucleoli. The regional and cellular distribution of cytosolic GSTs in the brain was studied by immunohistochemistry, spectrophotometric enzyme assay, and reverse-phase HPLC. Polyclonal antibody against microsomal GST was strongly reactive with Purkinje cells throughout the cerebellar cortex, and with neurons in the brainstem and hippocampus. Nuclei of Purkinje cells and of neurons in the brainstem, hippocampus, and cerebral cortex were immunopositive for alpha-class GST 1-1 (YaYa), whereas alpha-class GST 2-2 (YcYc) antibody was consistently immunoreactive with the nucleolus, but not with the nucleus or soma. All alpha-class GST antibodies studied were reactive, to various degrees, with astrocytes and choroid plexus; however, ependymal cells of the subventricular zones were immunonegative. alpha-class GST 8-8 (YkYk) immunoreactivity was specifically localized to endothelial cells and/or astrocytic end feet associated with blood vessels. Reverse-phase HPLC indicated that there were also substantial regional differences in the pattern of alpha-, mu-, and pi-class GST subunit expression. For example, the thalamus/hypothalamus had the highest GST activity and greatest concentration of total GST protein and mu-class GST subunit 6 (Yb3), whereas the brainstem had the greatest concentration of pi-class GST subunit (Yp). This regional variation in GST expression may be reflective of regional differences in cell populations. In cerebellar cortex, the concentration of mu-class GST subunit 4 (Yb2) was greatest in the flocculus and lowest in the vermis. This is of clinical interest because the pattern of expression of mu-class GST subunit 4 (Yb2) in the cerebellum coincides with the known regional susceptibility of this structure to degeneration after exposure to toxic or metabolic insults. The vermis is most susceptible to these insults, whereas the lateral lobes and flocculus are most resistant.(ABSTRACT TRUNCATED AT 400 WORDS)

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 to binaural intensity and phase difference cues in kitten inferior colliculus.

1. Responses of single neurons to monaural or binaural CF tones delivered through a closed and calibrated sound delivery system were studied in the central nucleus of the inferior colliculus (ICC) in ketamine and barbiturate-anesthetized kittens 4-105 days old. 2. Neurons from young kittens had elevated thresholds, some greater than 100 dB in the youngest kittens tested. Average thresholds in the ICC matched those previously measured in the auditory nerve (AN), cochlear nuclei (CN), and auditory cortex (CTX), suggesting that the drop in threshold as a function of age is primarily determined by development at the periphery. 3. Minimal first-spike latencies were relatively long in the youngest kittens, approaching adult values by the end of the third postnatal week. Latencies were distributed between values previously determined for the CN and auditory cortex and can be attributed to the centripetal development of the auditory system. 4. The range of frequencies that were effective in exciting ICC neurons was restricted in young kittens. Neurons having characteristic frequencies (CFs) greater than 7 kHz were not recorded before postnatal day 10. CF distribution matched that obtained in previous experiments from AN, CN, and CTX, reflecting the development of the cochlea. 5. Both monotonic and nonomonotonic spike count-versus-intensity functions were found in the youngest kittens. There was a tendency for monotonic functions from the youngest kittens to be steeper than those from older kittens. No age-related changes in the shapes of non-monotonic functions were found. 6. Sensitivity to interaural intensity difference (IID), tested by holding the intensity to the excitatory ear at a suprathreshold level and increasing the intensity of the stimulus to the inhibitory ear, was exhibited as early as 8 days after birth. The majority of the cells exhibiting sensitivity to IID (89.5%) were classified as EI cells, and almost all IID sensitive cells had CFs between 3 and 25 kHz. Within our sample the shapes of IID functions, as well as the operating range of the IID functions, closely resembled those obtained from the adult cat. Thresholds of excitation and of inhibition were highly correlated, suggesting that the ipsilateral and contralateral inputs to the ICC develop as a matched set. 7. Sensitivity to interaural phase difference (IPD), tested either by shifting the onset phase of a CF tone to one ear relative to the other or by presenting tones of slightly different frequency to the two ears, was present as early as 12 days after birth.(ABSTRACT TRUNCATED AT 400 WORDS)