Unanesthetized Rabbits Research Articles

Neural correlates of tonal loudness, intensity discrimination, and duration discrimination.

A long-standing quest in audition concerns understanding relations between behavioral measures and neural representations of changes in sound intensity. Here, we examined relations between aspects of intensity perception and central neural responses within the inferior colliculus of unanesthetized rabbits (by averaging the population's spike count/level functions). We found parallels between the population's neural output and: (1) how loudness grows with intensity; (2) how loudness grows with duration; (3) how discrimination of intensity improves with increasing sound level; (4) findings that intensity discrimination does not depend on duration; and (5) findings that duration discrimination is a constant fraction of base duration.

Effect of Reverberation on Neural Responses to Natural Speech in Rabbit Auditory Midbrain: No Evidence for a Neural Dereverberation Mechanism.

Reverberation is ubiquitous in everyday acoustic environments. It degrades both binaural cues and the envelope modulations of sounds and thus can impair speech perception. Still, both humans and animals can accurately perceive reverberant stimuli in most everyday settings. Previous neurophysiological and perceptual studies have suggested the existence of neural mechanisms that partially compensate for the effects of reverberation. However, these studies were limited by their use of either highly simplified stimuli or rudimentary reverberation simulations. To further characterize how reverberant stimuli are processed by the auditory system, we recorded single-unit (SU) and multiunit (MU) activity from the inferior colliculus (IC) of unanesthetized rabbits in response to natural speech utterances presented with no reverberation ("dry") and in various degrees of simulated reverberation (direct-to-reverberant energy ratios (DRRs) ranging from 9.4 to -8.2 dB). Linear stimulus reconstruction techniques (Mesgarani et al., 2009) were used to quantify the amount of speech information available in the responses of neural ensembles. We found that high-quality spectrogram reconstructions could be obtained for dry speech and in moderate reverberation from ensembles of 25 units. However, spectrogram reconstruction quality deteriorated in severe reverberation for both MUs and SUs such that the neural degradation paralleled the degradation in the stimulus spectrogram. Furthermore, spectrograms reconstructed from responses to reverberant stimuli resembled spectrograms of reverberant speech better than spectrograms of dry speech. Overall, the results provide no evidence for a dereverberation mechanism in neural responses from the rabbit IC when studied with linear reconstruction techniques.

Experimental retina's dystrophy and the curcumin recovery effect on the evoked activity of visual system structures

The rehabilitative curcumin effect on visual system function was investigated in studies conducted on awake, unanesthetized rabbits. Analysis of the results obtained showed that after curcumin administration, certain changes in the evoked activity of the retina, superior colliculus, lateral geniculate body, and primary visual cortex were observed in the model of retinal dystrophy. The neuronal apparatus of the glial layer of the retina damaged in moderate retinal dystrophy led to functional disintegration of the mechanisms of electrogenesis. The curcumin administration led to an increase in the amplitude parameters of evoked potentials in all structures. The most pronounced curcumin effect was observed in the potentials of the visual cortex evoked by light stimuli, where certain changes in the configuration of evoked responses were noted.

Robust Rate-Place Coding of Resolved Components in Harmonic and Inharmonic Complex Tones in Auditory Midbrain.

Harmonic complex tones (HCTs) commonly occurring in speech and music evoke a strong pitch at their fundamental frequency (F0), especially when they contain harmonics individually resolved by the cochlea. When all frequency components of an HCT are shifted by the same amount, the pitch of the resulting inharmonic tone (IHCT) can also shift, although the envelope repetition rate is unchanged. A rate-place code, whereby resolved harmonics are represented by local maxima in firing rates along the tonotopic axis, has been characterized in the auditory nerve and primary auditory cortex, but little is known about intermediate processing stages. We recorded single-neuron responses to HCT and IHCT with varying F0 and sound level in the inferior colliculus (IC) of unanesthetized rabbits of both sexes. Many neurons showed peaks in firing rate when a low-numbered harmonic aligned with the neuron's characteristic frequency, demonstrating "rate-place" coding. The IC rate-place code was most prevalent for F0 > 800 Hz, was only moderately dependent on sound level over a 40 dB range, and was not sensitive to stimulus harmonicity. A spectral receptive-field model incorporating broadband inhibition better predicted the neural responses than a purely excitatory model, suggesting an enhancement of the rate-place representation by inhibition. Some IC neurons showed facilitation in response to HCT relative to pure tones, similar to cortical "harmonic template neurons" (Feng and Wang, 2017), but to a lesser degree. Our findings shed light on the transformation of rate-place coding of resolved harmonics along the auditory pathway.SIGNIFICANCE STATEMENT Harmonic complex tones are ubiquitous in speech and music and produce strong pitch percepts when they contain frequency components that are individually resolved by the cochlea. Here, we characterize a "rate-place" code for resolved harmonics in the auditory midbrain that is more robust across sound levels than the peripheral rate-place code and insensitive to the harmonic relationships among frequency components. We use a computational model to show that inhibition may play an important role in shaping the rate-place code. Our study fills a major gap in understanding the transformations in neural representations of resolved harmonics along the auditory pathway.

Pitch of harmonic complex tones: rate and temporal coding of envelope repetition rate in inferior colliculus of unanesthetized rabbits.

Harmonic complex tones (HCTs) found in speech, music, and animal vocalizations evoke strong pitch percepts at their fundamental frequencies. The strongest pitches are produced by HCTs that contain harmonics resolved by cochlear frequency analysis, but HCTs containing solely unresolved harmonics also evoke a weaker pitch at their envelope repetition rate (ERR). In the auditory periphery, neurons phase lock to the stimulus envelope, but this temporal representation of ERR degrades and gives way to rate codes along the ascending auditory pathway. To assess the role of the inferior colliculus (IC) in such transformations, we recorded IC neuron responses to HCT and sinusoidally modulated broadband noise (SAMN) with varying ERR from unanesthetized rabbits. Different interharmonic phase relationships of HCT were used to manipulate the temporal envelope without changing the power spectrum. Many IC neurons demonstrated band-pass rate tuning to ERR between 60 and 1,600 Hz for HCT and between 40 and 500 Hz for SAMN. The tuning was not related to the pure-tone best frequency of neurons but was dependent on the shape of the stimulus envelope, indicating a temporal rather than spectral origin. A phenomenological model suggests that the tuning may arise from peripheral temporal response patterns via synaptic inhibition. We also characterized temporal coding to ERR. Some IC neurons could phase lock to the stimulus envelope up to 900 Hz for either HCT or SAMN, but phase locking was weaker with SAMN. Together, the rate code and the temporal code represent a wide range of ERR, providing strong cues for the pitch of unresolved harmonics.NEW & NOTEWORTHY Envelope repetition rate (ERR) provides crucial cues for pitch perception of frequency components that are not individually resolved by the cochlea, but the neural representation of ERR for stimuli containing many harmonics is poorly characterized. Here we show that the pitch of stimuli with unresolved harmonics is represented by both a rate code and a temporal code for ERR in auditory midbrain neurons and propose possible underlying neural mechanisms with a computational model.

Neural coding and perception of auditory motion direction based on interaural time differences.

While motion is important for parsing a complex auditory scene into perceptual objects, how it is encoded in the auditory system is unclear. Perceptual studies suggest that the ability to identify the direction of motion is limited by the duration of the moving sound, yet we can detect changes in interaural differences at even shorter durations. To understand the source of these distinct temporal limits, we recorded from single units in the inferior colliculus (IC) of unanesthetized rabbits in response to noise stimuli containing a brief segment with linearly time-varying interaural time difference ("ITD sweep") temporally embedded in interaurally uncorrelated noise. We also tested the ability of human listeners to either detect the ITD sweeps or identify the motion direction. Using a point-process model to separate the contributions of stimulus dependence and spiking history to single-neuron responses, we found that the neurons respond primarily by following the instantaneous ITD rather than exhibiting true direction selectivity. Furthermore, using an optimal classifier to decode the single-neuron responses, we found that neural threshold durations of ITD sweeps for both direction identification and detection overlapped with human threshold durations even though the average response of the neurons could track the instantaneous ITD beyond psychophysical limits. Our results suggest that the IC does not explicitly encode motion direction, but internal neural noise may limit the speed at which we can identify the direction of motion.NEW & NOTEWORTHY Recognizing motion and identifying an object's trajectory are important for parsing a complex auditory scene, but how we do so is unclear. We show that neurons in the auditory midbrain do not exhibit direction selectivity as found in the visual system but instead follow the trajectory of the motion in their temporal firing patterns. Our results suggest that the inherent variability in neural firings may limit our ability to identify motion direction at short durations.

Improved Neural Coding of ITD with Bilateral Cochlear Implants by Introducing Short Inter-pulse Intervals.

Bilateral cochlear implant (CI) users have poor perceptual sensitivity to interaural time differences (ITDs), which limits their ability to localize sounds and understand speech in noisy environments. This is especially true for high-rate (> 300pps) periodic pulse trains, which are used as carriers in CI processors. Here, we investigate a novel stimulation strategy in which extra pulses are added to high-rate periodic pulse trains to introduce short inter-pulse intervals (SIPIs). We hypothesized that SIPIs can improve neural ITD sensitivity similarly to the effect observed by randomly jittering IPIs (Hancock et al., J. Neurophysiol. 108:714-28, 2012). To test this hypothesis, we measured ITD sensitivity of single units in the inferior colliculus (IC) of unanesthetized rabbits with bilateral CIs. Introducing SIPIs into high-rate pulse trains significantly increased firing rates for ~ 60% of IC neurons, and the extra spikes tended to be synchronized to the SIPIs. The additional firings produced by SIPIs uncovered latent ITD sensitivity that was comparable to that observed with low-rate pulse trains. In some neurons, high spontaneous firing rates masked the ITD sensitivity introduced by SIPIs. ITD sensitivity in these neurons could be revealed by emphasizing stimulus-synchronized spikes with a coincidence detection analysis. Overall, these results with SIPIs are consistent with the effects observed previously with jittered pulse trains, with the added benefit of retaining control over the timing and number of SIPIs. A novel CI processing strategy could incorporate SIPIs by inserting them at selected times to high-rate pulse train carriers. Such a strategy could potentially improve ITD perception without degrading speech intelligibility and thereby improve outcomes for bilateral CI users.

Dual rate codes for the pitch of harmonic complex tones in the auditory midbrain

Harmonic complex tones contained in speech and music evoke a pitch at their fundamental frequency (F0). This pitch can be produced through either patterns of harmonics individually resolved by the cochlear frequency analysis or periodicity in the amplitude envelope resulting from beating between neighboring harmonics. To investigate how these two pitch cues are coded in the auditory midbrain, we recorded from single units in unanesthetized rabbits in response to harmonic complex tones with varying F0 so as to create conditions where harmonics were likely resolved or unresolved. For F0 above 600 Hz, some neurons showed local maxima in firing rate when a low-order (<6-8) harmonic of F0 coincided with the neuron’s best frequency (BF), thereby providing a rate-place code to resolved harmonics similar to that found in the auditory nerve. For F0 below 800 Hz, some neurons showed rate tuning to a particular F0 unrelated to the BF. Altering the phase relationships among the harmonics showed that this nontonotopic...

Neural coding of time-varying interaural time differences and time-varying amplitude in the inferior colliculus.

Binaural cues occurring in natural environments are frequently time varying, either from the motion of a sound source or through interactions between the cues produced by multiple sources. Yet, a broad understanding of how the auditory system processes dynamic binaural cues is still lacking. In the current study, we directly compared neural responses in the inferior colliculus (IC) of unanesthetized rabbits to broadband noise with time-varying interaural time differences (ITD) with responses to noise with sinusoidal amplitude modulation (SAM) over a wide range of modulation frequencies. On the basis of prior research, we hypothesized that the IC, one of the first stages to exhibit tuning of firing rate to modulation frequency, might use a common mechanism to encode time-varying information in general. Instead, we found weaker temporal coding for dynamic ITD compared with amplitude modulation and stronger effects of adaptation for amplitude modulation. The differences in temporal coding of dynamic ITD compared with SAM at the single-neuron level could be a neural correlate of "binaural sluggishness," the inability to perceive fluctuations in time-varying binaural cues at high modulation frequencies, for which a physiological explanation has so far remained elusive. At ITD-variation frequencies of 64 Hz and above, where a temporal code was less effective, noise with a dynamic ITD could still be distinguished from noise with a constant ITD through differences in average firing rate in many neurons, suggesting a frequency-dependent tradeoff between rate and temporal coding of time-varying binaural information.NEW & NOTEWORTHY Humans use time-varying binaural cues to parse auditory scenes comprising multiple sound sources and reverberation. However, the neural mechanisms for doing so are poorly understood. Our results demonstrate a potential neural correlate for the reduced detectability of fluctuations in time-varying binaural information at high speeds, as occurs in reverberation. The results also suggest that the neural mechanisms for processing time-varying binaural and monaural cues are largely distinct.

Neural Coding of Interaural Time Differences with Bilateral Cochlear Implants in Unanesthetized Rabbits.

Although bilateral cochlear implants (CIs) provide improvements in sound localization and speech perception in noise over unilateral CIs, bilateral CI users' sensitivity to interaural time differences (ITDs) is still poorer than normal. In particular, ITD sensitivity of most CI users degrades with increasing stimulation rate and is lacking at the high carrier pulse rates used in CI processors to deliver speech information. To gain a better understanding of the neural basis for this degradation, we characterized ITD tuning of single neurons in the inferior colliculus (IC) for pulse train stimuli in an unanesthetized rabbit model of bilateral CIs. Approximately 73% of IC neurons showed significant ITD sensitivity in their overall firing rates. On average, ITD sensitivity was best for pulse rates near 80-160 pulses per second (pps) and degraded for both lower and higher pulse rates. The degradation in ITD sensitivity at low pulse rates was caused by strong, unsynchronized background activity that masked stimulus-driven responses in many neurons. Selecting synchronized responses by temporal windowing revealed ITD sensitivity in these neurons. With temporal windowing, both the fraction of ITD-sensitive neurons and the degree of ITD sensitivity decreased monotonically with increasing pulse rate. To compare neural ITD sensitivity to human performance in ITD discrimination, neural just-noticeable differences (JNDs) in ITD were computed using signal detection theory. Using temporal windowing at lower pulse rates, and overall firing rate at higher pulse rates, neural ITD JNDs were within the range of perceptual JNDs in human CI users over a wide range of pulse rates. Many profoundly deaf people wearing cochlear implants (CIs) still face challenges in everyday situations, such as understanding conversations in noise. Even with CIs in both ears, they have difficulty making full use of subtle differences in the sounds reaching the two ears [interaural time difference (ITD)] to identify where the sound is coming from. This problem is especially acute at the high stimulation rates used in clinical CI processors. This study provides a better understanding of ITD processing with bilateral CIs and shows a parallel between human performance in ITD discrimination and neural responses in the auditory midbrain. The present study is the first report on binaural properties of auditory neurons with CIs in unanesthetized animals.

Neural population encoding and decoding of sound source location across sound level in the rabbit inferior colliculus.

At lower levels of sensory processing, the representation of a stimulus feature in the response of a neural population can vary in complex ways across different stimulus intensities, potentially changing the amount of feature-relevant information in the response. How higher-level neural circuits could implement feature decoding computations that compensate for these intensity-dependent variations remains unclear. Here we focused on neurons in the inferior colliculus (IC) of unanesthetized rabbits, whose firing rates are sensitive to both the azimuthal position of a sound source and its sound level. We found that the azimuth tuning curves of an IC neuron at different sound levels tend to be linear transformations of each other. These transformations could either increase or decrease the mutual information between source azimuth and spike count with increasing level for individual neurons, yet population azimuthal information remained constant across the absolute sound levels tested (35, 50, and 65 dB SPL), as inferred from the performance of a maximum-likelihood neural population decoder. We harnessed evidence of level-dependent linear transformations to reduce the number of free parameters in the creation of an accurate cross-level population decoder of azimuth. Interestingly, this decoder predicts monotonic azimuth tuning curves, broadly sensitive to contralateral azimuths, in neurons at higher levels in the auditory pathway.

Neural coding of sound envelope in reverberant environments.

Speech reception depends critically on temporal modulations in the amplitude envelope of the speech signal. Reverberation encountered in everyday environments can substantially attenuate these modulations. To assess the effect of reverberation on the neural coding of amplitude envelope, we recorded from single units in the inferior colliculus (IC) of unanesthetized rabbit using sinusoidally amplitude modulated (AM) broadband noise stimuli presented in simulated anechoic and reverberant environments. Although reverberation degraded both rate and temporal coding of AM in IC neurons, in most neurons, the degradation in temporal coding was smaller than the AM attenuation in the stimulus. This compensation could largely be accounted for by the compressive shape of the modulation input-output function (MIOF), which describes the nonlinear transformation of modulation depth from acoustic stimuli into neural responses. Additionally, in a subset of neurons, the temporal coding of AM was better for reverberant stimuli than for anechoic stimuli having the same modulation depth at the ear. Using hybrid anechoic stimuli that selectively possess certain properties of reverberant sounds, we show that this reverberant advantage is not caused by envelope distortion, static interaural decorrelation, or spectral coloration. Overall, our results suggest that the auditory system may possess dual mechanisms that make the coding of amplitude envelope relatively robust in reverberation: one general mechanism operating for all stimuli with small modulation depths, and another mechanism dependent on very specific properties of reverberant stimuli, possibly the periodic fluctuations in interaural correlation at the modulation frequency.

Azimuth and envelope coding in the inferior colliculus of the unanesthetized rabbit: effect of reverberation and distance.

Recognition and localization of a sound are the major functions of the auditory system. In real situations, the listener and different degrees of reverberation transform the signal between the source and the ears. The present study was designed to provide these transformations and examine their influence on neural responses. Using the virtual auditory space (VAS) method to create anechoic and moderately and highly reverberant environments, we found the following: 1) In reverberation, azimuth tuning was somewhat degraded with distance whereas the direction of azimuth tuning remained unchanged. These features remained unchanged in the anechoic condition. 2) In reverberation, azimuth tuning and envelope synchrony were degraded most for neurons with low best frequencies and least for neurons with high best frequencies. 3) More neurons showed envelope synchrony to binaural than to monaural stimulation in both anechoic and reverberant environments. 4) The percentage of envelope-coding neurons and their synchrony decreased in reverberation with distance, whereas it remained constant in the anechoic condition. 5) At far distances, for both binaural and monaural stimulation, the neural gain in reverberation could be as high as 30 dB and as much as 10 dB higher than those in the anechoic condition. 6) The majority of neurons were able to code both envelope and azimuth in all of the environments. This study provides a foundation for understanding the neural coding of azimuth and envelope synchrony at different distances in reverberant and anechoic environments. This is necessary to understand how the auditory system processes "where" and "what" information in real environments.

Dual sensitivity of inferior colliculus neurons to ITD in the envelopes of high-frequency sounds: experimental and modeling study

Human listeners are sensitive to interaural time differences (ITDs) in the envelopes of sounds, which can serve as a cue for sound localization. Many high-frequency neurons in the mammalian inferior colliculus (IC) are sensitive to envelope-ITDs of sinusoidally amplitude-modulated (SAM) sounds. Typically, envelope-ITD-sensitive IC neurons exhibit either peak-type sensitivity, discharging maximally at the same delay across frequencies, or trough-type sensitivity, discharging minimally at the same delay across frequencies, consistent with responses observed at the primary site of binaural interaction in the medial and lateral superior olives (MSO and LSO), respectively. However, some high-frequency IC neurons exhibit dual types of envelope-ITD sensitivity in their responses to SAM tones, that is, they exhibit peak-type sensitivity at some modulation frequencies and trough-type sensitivity at other frequencies. Here we show that high-frequency IC neurons in the unanesthetized rabbit can also exhibit dual types of envelope-ITD sensitivity in their responses to SAM noise. Such complex responses to SAM stimuli could be achieved by convergent inputs from MSO and LSO onto single IC neurons. We test this hypothesis by implementing a physiologically explicit, computational model of the binaural pathway. Specifically, we examined envelope-ITD sensitivity of a simple model IC neuron that receives convergent inputs from MSO and LSO model neurons. We show that dual envelope-ITD sensitivity emerges in the IC when convergent MSO and LSO inputs are differentially tuned for modulation frequency.

High Frequency Burst Firing of Granule Cells Ensures Transmission at the Parallel Fiber to Purkinje Cell Synapse at the Cost of Temporal Coding

Cerebellar granule cells (GrCs) convey information from mossy fibers (MFs) to Purkinje cells (PCs) via their parallel fibers (PFs). MF to GrC signaling allows transmission of frequencies up to 1 kHz and GrCs themselves can also fire bursts of action potentials with instantaneous frequencies up to 1 kHz. So far, in the scientific literature no evidence has been shown that these high-frequency bursts also exist in awake, behaving animals. More so, it remains to be shown whether such high-frequency bursts can transmit temporally coded information from MFs to PCs and/or whether these patterns of activity contribute to the spatiotemporal filtering properties of the GrC layer. Here, we show that, upon sensory stimulation in both un-anesthetized rabbits and mice, GrCs can show bursts that consist of tens of spikes at instantaneous frequencies over 800 Hz. In vitro recordings from individual GrC-PC pairs following high-frequency stimulation revealed an overall low initial release probability of ~0.17. Nevertheless, high-frequency burst activity induced a short-lived facilitation to ensure signaling within the first few spikes, which was rapidly followed by a reduction in transmitter release. The facilitation rate among individual GrC-PC pairs was heterogeneously distributed and could be classified as either “reluctant” or “responsive” according to their release characteristics. Despite the variety of efficacy at individual connections, grouped activity in GrCs resulted in a linear relationship between PC response and PF burst duration at frequencies up to 300 Hz allowing rate coding to persist at the network level. Together, these findings support the hypothesis that the cerebellar granular layer acts as a spatiotemporal filter between MF input and PC output (D’Angelo and De Zeeuw, 2009).

Neural encoding of sound source location in the presence of a concurrent, spatially separated source

In the presence of multiple, spatially separated sound sources, the binaural cues used for sound localization in the horizontal plane become distorted from the cues from each sound in isolation, yet localization in everyday multisource acoustic environments remains robust. We examined changes in the azimuth tuning functions of inferior colliculus (IC) neurons in unanesthetized rabbits to a target broadband noise when a concurrent broadband noise interferer was presented at different locations in virtual acoustic space. The presence of an interferer generally degraded sensitivity to target azimuth and distorted the shape of the tuning function, yet most neurons remained significantly sensitive to target azimuth and maintained tuning function shapes somewhat similar to those for the target alone. Using binaural cue manipulations in virtual acoustic space, we found that single-source tuning functions of neurons with high best frequencies (BFs) were primarily determined by interaural level differences (ILDs) or monaural level, with a small influence of interaural time differences (ITDs) in some neurons. However, with a centrally located interferer, the tuning functions of most high-BF neurons were strongly influenced by ITDs as well as ILDs. Model-based analysis showed that the shapes of these tuning functions were in part produced by decorrelation of the left and right cochlea-induced envelopes that occurs with source separation. The strong influence of ITD on the tuning functions of high-BF neurons poses a challenge to the "duplex theory" of sound localization and suggests that ITD may be important for localizing high-frequency sounds in multisource environments.

Extraocular muscle motor units characterized by spike-triggered averaging in alert monkey

Single-unit recording in macaque monkeys has been widely used to study extraocular motoneuron behavior during eye movements. However, primate extraocular motor units have only been studied using electrical stimulation in anesthetized animals. To study motor units in alert, behaving macaques, we combined chronic muscle force transducer (MFT) and single-unit extracellular motoneuron recordings. During steady fixation with low motoneuron firing rates, we used motoneuron spike-triggered averaging of MFT signals (STA-MFT) to extract individual motor unit twitches, thereby characterizing each motor unit in terms of twitch force and dynamics. It is then possible, as in conventional studies, to determine motoneuron activity during eye movements, but now with knowledge of underlying motor unit characteristics.We demonstrate the STA-MFT technique for medial rectus motor units. Recordings from 33 medial rectus motoneurons in three animals identified 20 motor units, which had peak twitch tensions of 0.5–5.25mg, initial twitch delays averaging 2.4ms, and time to peak contraction averaging 9.3ms. These twitch tensions are consistent with those reported in unanesthetized rabbits, and with estimates of the total number of medial rectus motoneurons and twitch tension generated by whole-nerve stimulation in monkey, but are substantially lower than those reported for lateral rectus motor units in anesthetized squirrel monkey. Motor units were recruited in order of twitch tension magnitude with stronger motor units reaching threshold further in the muscle's ON-direction, showing that, as in other skeletal muscles, medial rectus motor units are recruited according to the “size principle”.

Spatial tuning to sound-source azimuth in the inferior colliculus of unanesthetized rabbit.

Despite decades of research devoted to the study of inferior colliculus (IC) neurons' tuning to sound-source azimuth, there remain many unanswered questions because no previous study has examined azimuth tuning over a full range of 360° azimuths at a wide range of stimulus levels in an unanesthetized preparation. Furthermore, a comparison of azimuth tuning to binaural and contralateral ear stimulation over ranges of full azimuths and widely varying stimulus levels has not previously been reported. To fill this void, we have conducted a study of azimuth tuning in the IC of the unanesthetized rabbit over a 300° range of azimuths at stimulus levels of 10-50 dB above neural threshold to both binaural and contralateral ear stimulation using virtual auditory space stimuli. This study provides systematic evidence for neural coding of azimuth. We found the following: 1) level-tolerant azimuth tuning was observed in the top 35% regarding vector strength and in the top 15% regarding vector angle of IC neurons; 2) preserved azimuth tuning to binaural stimulation at high stimulus levels was created as a consequence of binaural facilitation in the contralateral sound field and binaural suppression in the ipsilateral sound field; 3) the direction of azimuth tuning to binaural stimulation was primarily in the contralateral sound field, and its center shifted laterally toward -90° with increasing stimulus level; 4) at 10 dB, azimuth tuning to binaural and contralateral stimulation was similar, indicating that it was mediated by monaural mechanisms; and 5) at higher stimulus levels, azimuth tuning to contralateral ear stimulation was severely degraded. These findings form a foundation for understanding neural mechanisms of localizing sound-source azimuth.

Pulse flows of populations of cortical neurons under microwave irradiation: Burst activity

The pulse flows of populations of sensomotor cortical neurons are studied in unanesthetized non-immobilized rabbits before, during, and after 1-min exposure to microwave electromagnetic irradiation (wavelength 37.5 cm, power density 0.2–0.3, 0.4, 0.5, and 40 mW/cm2) by analyzing the burst activity detected at time thresholds of 5, 10, and 20 ms. These exposures change the number of spike bursts. The changes in pulse flows are recorded both during irradiation and during the first minute after its cessation. The directions of shifts and their dynamics are determined by both the irradiation intensity and the types of bursts themselves.

Effects of Reverberation on the Directional Sensitivity of Auditory Neurons across the Tonotopic Axis: Influences of Interaural Time and Level Differences

In reverberant environments, acoustic reflections interfere with the direct sound arriving at a listener's ears, distorting the binaural cues for sound localization. We investigated the effects of reverberation on the directional sensitivity of single neurons in the inferior colliculus (IC) of unanesthetized rabbits. We find that reverberation degrades the directional sensitivity of single neurons, although the amount of degradation depends on the characteristic frequency (CF) and the type of binaural cues available. When interaural time differences (ITDs) are the only available directional cue, low-CF cells sensitive to ITDs in the waveform fine time structure maintain better directional sensitivity in reverberation than high-CF cells sensitive to ITDs in the envelope induced by cochlear filtering. Conversely, when both ITD and interaural level difference (ILD) cues are available, directional sensitivity in reverberation is comparable throughout the tonotopic axis of the IC. This result suggests that, at high frequencies, ILDs provide better directional information than envelope ITDs, emphasizing the importance of the ILD-processing pathway for sound localization in reverberation.