PubMed Health⌕ Search

Biomedical subjects

Philip X Joris

Publications and source records attributed to Philip X Joris.

13 recordsLinked to original sources

Panoramic measurements of the apex of the cochlea.

Our understanding of cochlear mechanics is impeded by the lack of truly panoramic data. Sensitive mechanical measurements cover only a narrow cochlear region, mostly in the base. The global spatiotemporal pattern of vibrations along the cochlea cannot be inferred from such local measurements but is often extrapolated beyond the measurement spot under the assumption of scaling invariance. Auditory nerve responses give an alternative window on the entire cochlea, but traditional techniques do not allow recovery of the effective vibration pattern. We developed a new analysis technique to measure cochlear amplitude and phase transfer of fibers with characteristic frequencies <5 kHz. Data from six cats yielded panoramic phase profiles along the apex of the cochlea for an approximately 5 octave range of stimulus frequencies. All profiles accumulated systematic phase lags from base to apex. Phase accumulation was not gradual but showed a two-segment character: a steep segment (slow propagation) around the characteristic position of the stimulus, and a shallow segment (fast propagation) basal to it. The transition between the segments occurred in a narrow region and was smooth. Wavelength near characteristic position decreased from approximately 3.5 to approximately 1 mm for frequencies from 200 to 4000 Hz, corresponding to phase velocities of approximately 0.5 to approximately 5 m/s. The accumulated phase lag between the eardrum and characteristic position varied from approximately 1 cycle at 200 Hz to approximately 2.5 cycle at 4 kHz, invalidating scaling invariance. The generic character of our analysis technique and its success in solving the difficult problem of reconstructing the effective sensory input from neural recordings suggest its wider application as a powerful alternative to customary system analysis techniques.

Acoustic Stimulation↗

Binaural and cochlear disparities.

Binaural auditory neurons exhibit "best delays" (BDs): They are maximally activated at certain acoustic delays between sounds at the two ears and thereby signal spatial sound location. BDs arise from delays internal to the auditory system, but their source is controversial. According to the classic Jeffress model, they reflect pure time delays generated by differences in axonal length between the inputs from the two ears to binaural neurons. However, a relationship has been reported between BDs and the frequency to which binaural neurons are most sensitive (the characteristic frequency), and this relationship is not predicted by the Jeffress model. An alternative hypothesis proposes that binaural neurons derive their input from slightly different places along the two cochleas, which induces BDs by virtue of the slowness of the cochlear traveling wave. To test this hypothesis, we performed a coincidence analysis on spiketrains of pairs of auditory nerve fibers originating from different cochlear locations. In effect, this analysis mimics the processing of phase-locked inputs from each ear by binaural neurons. We find that auditory nerve fibers that innervate different cochlear sites show a maximum number of coincidences when they are delayed relative to each other, and that the optimum delays decrease with characteristic frequency as in binaural neurons. These findings suggest that cochlear disparities make an important contribution to the internal delays observed in binaural neurons.

Animals↗

Correlation index: a new metric to quantify temporal coding.

The standard procedure to study temporal encoding of sound waveforms in the auditory system has been Fourier analysis of responses to periodic stimuli. We introduce a new metric--correlation index (CI)--which is based on a simple counting of spike coincidences. It can be used for responses to aperiodic stimuli and does not require knowledge of the stimulus. Moreover, the basic procedure of comparing spiketimes in spiketrains is more physiological than currently used methods for temporal analysis. The CI is the peak value of the normalized shuffled autocorrelogram (SAC), which provides a quantitative summary of temporal structure in the neural response to arbitrary stimuli. We illustrate the CI and SACs by comparing temporal coding in the auditory nerve and output fibers of the cochlear nucleus.

Acoustic Stimulation↗

Auditory midbrain and nerve responses to sinusoidal variations in interaural correlation.

The human sensitivity to interaural temporal differences in the acoustic waveforms to the two ears shows remarkable acuity but is also very sluggish. Fast changes in binaural parameters are not detectable, and this inability contrasts sharply with the excellent temporal resolution of the monaural auditory system. We studied the response of binaural neurons in the inferior colliculus of the cat to sinusoidal changes in the interaural correlation of broadband noise. Responses to the same waveforms were also obtained from auditory nerve fibers and further analyzed with a coincidence analysis. Overall, the auditory nerve and inferior colliculus showed a similar ability to code changes in interaural correlation. This ability extended to modulation frequencies an order of magnitude higher than the highest frequencies detected binaurally in humans. We conclude that binaural sluggishness is not caused by a lack of temporal encoding of fast binaural changes at the level of the midbrain. We hypothesize that there is no neural substrate at the level of the midbrain or higher to read out this temporal code and that this constitutes a low-pass "sluggishness" filter.

Acoustic Stimulation↗

Decorrelation sensitivity of auditory nerve and anteroventral cochlear nucleus fibers to broadband and narrowband noise.

Binaural neurons show remarkable sensitivity to temporal differences in the waveforms at the two ears. This ability obviously requires temporal coding of sound waveforms in the monaural afferents that converge on such binaural neurons. We introduce a new analysis to investigate how well responses of single monaural neurons support discrimination of decorrelations in waveforms. Spike trains from auditory nerve (AN) and anteroventral cochlear nucleus (AVCN) neurons of cats to many repetitions of a set of broadband and narrowband noise tokens were obtained. The normalized correlation between the noise tokens ranged from 0.99 to -1. A coincidence and signal detection analysis was used to perform a correlation discrimination task using the monaural spike trains. The correlation discrimination thresholds derived from AVCN neurons were lower than those derived from AN fibers and sometimes as low as human psychophysical just noticeable differences. Importantly, low detection thresholds required comparisons of spike trains at small internal delays. Bandwidth dependence of neural decorrelation thresholds agreed with psychophysical data when large internal delays contributed to the detection. We conclude that, in the context of correlation discrimination, coding by AVCN fibers is superior to that by AN fibers and that these discriminations require a distribution of internal or best delays in binaural processing that differs from the predictions from studies of discrimination in interaural time delays.

Acoustic Stimulation↗

Acoustic stria: anatomy of physiologically characterized cells and their axonal projection patterns.

The mammalian cochlear nucleus (CN) has been a model structure to study the relationship between physiological and morphological cell classes. Several issues remain, in particular with regard to the projection patterns and physiology of neurons that exit the CN dorsally via the dorsal (DAS), intermediate (IAS), and commissural stria. We studied these neurons physiologically and anatomically using the intra-axonal labeling method. Multipolar cells with onset chopper (O(C)) responses innervated the ipsilateral ventral and dorsal CN before exiting the CN via the commissural stria. Upon reaching the midline they turned caudally to innervate the opposite CN. No collaterals were seen innervating any olivary complex nuclei. Octopus cells typically showed onset responses with little or no sustained activity. The main axon used the IAS and followed one of two routes occasionally giving off olivary complex collaterals on their way to the contralateral ventral nucleus of the lateral lemniscus (VNLL). Here they can have elaborate terminal arbors that surround VNLL cells. Fusiform and giant cells have overlapping but not identical physiology. Fusiform but not giant cells typically show pauser or buildup responses. Axons of both cells exit via the DAS and take the same course to reach the contralateral IC without giving off any collaterals en route.

Acoustic Stimulation↗

Enhanced temporal response properties of anteroventral cochlear nucleus neurons to broadband noise.

Compared with auditory nerve (AN) fibers, trapezoid body (TB) fibers of the cat show enhanced synchronization to low-frequency tones. This phenomenon probably contributes to the high temporal resolution of binaural processing. We examined whether enhanced synchronization also occurs to sustained broadband noise. We recorded responses to a reference Gaussian noise and its polarity-inverted version in the TB of barbiturate-anesthetized cats. From these we constructed shuffled autocorrelograms (SACs) and quantified spike timing by measuring the amplitude and width of their central peak. Many TB fibers with low characteristic frequency (CF) showed SACs with higher and narrower central peaks than ever observed in the AN, indicating better consistency and precision of temporal coding. Larger peaks were also observed in TB fibers with high CF, but this was mostly caused by higher average firing rates, resulting in a larger number of coincident spikes across stimulus repetitions. The results document monaural preprocessing of the temporal information delivered to binaural nuclei in the olivary complex, which likely contributes to the high sensitivity to interaural time differences.

Acoustic Stimulation↗

Temporal damping in response to broadband noise. I. Inferior colliculus.

Many cells in the inferior colliculus (IC) are sensitive to interaural time differences (ITDs), in the form of an oscillatory dependency of average firing rate on ITD. We studied the degree of damping in such binaural responses, recording from neurons in the inferior colliculus of pentobarbital-anesthetized cats to binaural broadband noise and tones. Noise-delay functions and composite curves were characterized by computing the difference between responses to correlated and anticorrelated stimuli. We use a new metric, based on the envelope of this difference, to quantify damping. There is a clear relationship between damping and characteristic frequency (CF), but even neurons of the same CF can differ in their damping. For individual cells, damping can be stronger to tones or to noise; at the population level the two are positively correlated and are scarcely affected by SPL. The frequencies that dominate ITD sensitivity are near the CF in response to noise, but are often below CF in response to tones. These findings qualify conclusions from earlier reports but overall they support the conclusion that, at a population level, basic aspects of binaural responses to wideband noise are consistent with summed responses to pure tones.

Acoustic Stimulation↗

The speed of auditory low-side suppression.

The nonlinear cochlear phenomenon of two-tone suppression is known to be very fast, but precisely how fast is unknown. We studied the timing of low-side suppression in the auditory nerve of the cat using multitone complexes as auditory stimuli. An evalution of the group delays of the responses to these complexes allowed us to measure the timing of the responses with sub-millisecond accuracy for a large number of fibers with characteristic frequencies (CFs) between 2 and 40 kHz. In particular, we measured the delays with which the same below-CF tone complexes affected the response either as an excitor (when presented alone) or as a suppressor (when combined with a CF probe). For CFs <10 kHz, we found that the delay of suppression was larger than the delay of excitation by several hundred microseconds. The difference between the delay of suppression and that of excitation decreased with increasing CF, becoming negligible for CFs >15 kHz. The results are analyzed in terms of traveling-wave delays and a purported cochlear gain control. The data suggest that suppression originates from a gain-control mechanism with an integration time in the order of two cycles of CF.

Acoustic Stimulation↗

Temporal properties of responses to broadband noise in the auditory nerve.

Temporal information in the responses of auditory neurons to sustained sounds has been studied mostly with periodic stimuli, using measures that are based on Fourier analysis. Less information is available on temporal aspects of responses to nonperiodic wideband sounds. We recorded responses to a reference Gaussian noise and its polarity-inverted version in the auditory nerve of barbiturate-anesthetized cats and used shuffled autocorrelograms (SACs) to quantify spike timing. Two metrics were extracted from the central peak of autocorrelograms: the peak-height and the width at halfheight. Temporal information related to stimulus fine-structure was isolated from that to envelope by subtracting or adding responses to the reference and inverted noise. Peak-height and halfwidth generally behaved as expected from the existing body of data on phase-locking to pure tones and sinusoidally amplitude-modulated tones but showed some surprises as well. Compared with synchronization to low-frequency tones, SACs reveal large differences in temporal behavior between the different classes of nerve fibers (based on spontaneous rate) as well as a strong dependence on characteristic frequency (CF) throughout the phase-locking range. SACs also reveal a larger temporal consistency (i.e., tendency to discharge at the same point in time on repeated presentation of the same stimulus) in the responses to the stochastic noise stimulus than in the responses to periodic tones. Responses at high CFs reflect envelope phase-locking and are consistent with previous reports using sinusoidal AM. We conclude that the combined use of broadband noise and SAC analysis allow a more general characterization of temporal behavior than periodic stimuli and Fourier analysis.

Acoustic Stimulation↗

Cochlear phase and amplitude retrieved from the auditory nerve at arbitrary frequencies.

It is currently impossible to mechanically measure the overall vibration pattern of the intact mammalian cochlea because of its inaccessibility and vulnerability. At first sight, data from the auditory nerve are a poor substitute because of their limited temporal resolution. The nonlinear character of neural coding, however, causes low-frequency interactions among the components of multitone stimuli. We designed a novel stimulus for which these interactions take a particularly systematic form, and we recorded the response of the auditory nerve to this stimulus. A careful analysis of interactions in the data allowed us to reconstruct frequency transfer functions (both their amplitude and their phase) at multiple points spanning the entire length of the cochlea. The generic character of our stimuli and analysis suggests its wider use in nonlinear system analysis, particularly in those instances in which limitations in temporal resolution restrict the use of customary methods.

Acoustic Stimulation↗

Interaural time sensitivity dominated by cochlea-induced envelope patterns.

To localize sounds in space, humans heavily depend on minute interaural time differences (ITDs) generated by path-length differences to the two ears. Physiological studies of ITD sensitivity have mostly used deterministic, periodic sounds, in which either the waveform fine structure or a sinusoidal envelope is delayed interaurally. For natural broadband stimuli, however, auditory frequency selectivity causes individual channels to have their own envelopes; the temporal code in these channels is thus a mixture of fine structure and envelope. This study introduces a method to disentangle the contributions of fine structure and envelope in both binaural and monaural responses to broadband noise. In the inferior colliculus (IC) of the cat, a population of neurons was found in which envelope fluctuations dominate ITD sensitivity. This population extends over a surprisingly wide range of frequencies, including low frequencies for which fine-structure information is also available. A comparison with the auditory nerve suggests that an elaboration of envelope coding occurs between the nerve and the IC. These results suggest that internally generated envelopes play a more important role in binaural hearing than is commonly thought.

Acoustic Stimulation↗