1. Encoding temporal features of the acoustic waveform is an important attribute of the auditory system. Auditory nerve (AN) fibers synchronize or phase-lock to low-frequency tones and transmit this temporal information to cells in the anteroventral cochlear nucleus (AVCN). Phase-locking in the AVCN is usually reported to be similar to or weaker than in the AN. We studied phase-locking in axons of the trapezoid body (TB), which is the output tract of the AVCN, and found, to our surprise, that most TB axons exhibited enhanced synchronization compared with AN fibers. 2. Responses from axons in the TB of the cat were obtained with horseradish peroxidase (HRP)- or Neurobiotin-filled micropipettes or metal microelectrodes. A series of short tone bursts at increasing sound pressure level (SPL) was presented at the characteristic frequency (CF) of the fiber and phase-locking was quantified with the vector strength R at each SPL. For each fiber the maximum R value (Rmax) was then determined. 3. Low-frequency fibers in the TB showed very precise phase-locking: Rmax values could approach 0.99. For the majority of fibers (33/44, 75%) with CF < 700 Hz, Rmax was > or = 0.9 and therefore higher than is ever observed in the AN. We define such fibers as "high-sync." Most of these fibers also entrained to the stimulus, i.e., they fired a precisely timed action potential to almost every stimulus cycle. Some fibers showed perfect entrainment, with maximum discharge rates equaling the stimulus frequency. 4. To exclude the possibility that stimulus paradigms or acoustic and recording equipment were the source of this enhancement, we obtained additional data on low-frequency AN fibers using the same experimental protocol as in our TB experiments. These AN data agree well with published reports. 5. The morphological class of some of the cells studied was identified on the basis of anatomic features revealed by intra-axonal injection of HRP or Neurobiotin. Labeled low-CF axons (N = 7), which were all high-sync, originated from AVCN bushy cells: five were globular and two were spherical bushy cell axons. 6. Spontaneous rate of high-sync fibers covered a range from 0 to 176 spikes/s but were biased toward low values (mean 16 spikes/s). Responses to broadband clicks and sinusoidally amplitude-modulated signals provided additional evidence of improved timing properties.(ABSTRACT TRUNCATED AT 400 WORDS)
A phenomenological model of the auditory periphery in cats was previously developed by Zilany and colleagues [J. Acoust. Soc. Am. 126, 2390-2412 (2009)] to examine the detailed transformation of acoustic signals into the auditory-nerve representation. In this paper, a few issues arising from the responses of the previous version have been addressed. The parameters of the synapse model have been readjusted to better simulate reported physiological discharge rates at saturation for higher characteristic frequencies [Liberman, J. Acoust. Soc. Am. 63, 442-455 (1978)]. This modification also corrects the responses of higher-characteristic frequency (CF) model fibers to low-frequency tones that were erroneously much higher than the responses of low-CF model fibers in the previous version. In addition, an analytical method has been implemented to compute the mean discharge rate and variance from the model's synapse output that takes into account the effects of absolute refractoriness.
There is growing evidence that the dynamics of biological systems that appear to be exponential over short time courses are in some cases better described over the long-term by power-law dynamics. A model of rate adaptation at the synapse between inner hair cells and auditory-nerve ͑AN͒ fibers that includes both exponential and power-law dynamics is presented here. Exponentially adapting components with rapid and short-term time constants, which are mainly responsible for shaping onset responses, are followed by two parallel paths with power-law adaptation that provide slowly and rapidly adapting responses. The slowly adapting power-law component significantly improves predictions of the recovery of the AN response after stimulus offset. The faster power-law adaptation is necessary to account for the "additivity" of rate in response to stimuli with amplitude increments. The proposed model is capable of accurately predicting several sets of AN data, including amplitude-modulation transfer functions, long-term adaptation, forward masking, and adaptation to increments and decrements in the amplitude of an ongoing stimulus.
We made intraaxonal recordings from 30 individual globular bushy cell axons in the trapezoid body of the cat using HRP-filled glass microelectrodes. With subsequent HRP injection, we determined their axonal projection patterns. For cells with characteristic frequencies (CFs) above 3 kHz, short-tone peristimulus time histograms (PSTHs) at CF were typically primarylike at low tone intensities and primarylike with notch (PLN) or onset with low sustained activity (OL) at higher stimulus levels. Cells with CFs between 1 and 3 kHz showed the same response features with the spikes in the sustained region of the response phase-locked to the stimulus tone. Cells with CFs below 1 kHz showed phase-locked PSTHs with exceptionally high levels of synchrony compared to eighth nerve fibers with comparable CFs. This exceptional phase-locking was also noted when cells with CFs of 1-3 kHz were presented with tones below 1 kHz. Although the globular bushy cell axons were not completely filled from the soma of origin to terminal fields in the contralateral brainstem, a number of consistent anatomical features were distinguished in the population. All but one of the myelinated axons crossed the midline in the middle, large fiber component of the trapezoid body. Ipsilaterally, the axon always gave off from one to four collateral branches whose major targets were the posterior periolivary nucleus (PPO) and the lateral nucleus of the trapezoid body (LNTB). Minor termination sites for ipsilateral collateral branches were the dorsolateral periolivary nucleus (DLPO) and the lateral superior olive (LSO). Contralaterally the axon gave rise to one or two calyces of Held in the medial nucleus of the trapezoid body (MNTB). Three other major collateral branches arose from the contralateral axon and innervated a consistent set of areas. One headed caudally to innervate an area just ventromedial to the facial nucleus. Another followed the sixth nerve dorsally to innervate the dorsomedial periolivary nucleus (DMPO). A third collateral headed rostrally toward the ventral nucleus of the lateral lemniscus (VNLL), giving off occasional small sidebranches. Although each injected axon gave rise to a collateral that innervated the MNTB, it did not necessarily give rise to all three of the other collateral branches.
A phenomenological model was developed to describe responses of high-spontaneous-rate auditory-nerve (AN) fibers, including several nonlinear response properties. Level-dependent gain (compression), bandwidth, and phase properties were implemented with a control path that varied the gain and bandwidth of tuning in the signal-path filter. By making the bandwidth of the control path broad with respect to the signal path, the wide frequency range of two-tone suppression was included. By making the control-path filter level dependent and tuned to a frequency slightly higher than the signal-path filter, other properties of two-tone suppression were also included. These properties included the asymmetrical growth of suppression above and below the characteristic frequency and the frequency offset of the suppression tuning curve with respect to the excitatory tuning curve. The implementation of this model represents a relatively simple phenomenological description of a single mechanism that underlies several important nonlinear response properties of AN fibers. The model provides a tool for studying the roles of these nonlinearities in the encoding of simple and complex sounds in the responses of populations of AN fibers.
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