Frequency-dependent interaural delays in the medial superior olive: implications for interaural cochlear delays

2011 ◽  
Vol 106 (4) ◽  
pp. 1985-1999 ◽  
Author(s):  
Mitchell L. Day ◽  
Malcolm N. Semple
Keyword(s):  
Interaural Time Difference ◽  
Superior Olivary Complex ◽  
Medial Superior Olive ◽  
Biophysical Parameters ◽  
Tone Frequency ◽  
Frequency Dependent ◽  
Superior Olive ◽  
Interaural Phase ◽  
Cochlear Delays ◽  
Delay Component

Neurons in the medial superior olive (MSO) are tuned to the interaural time difference (ITD) of sound arriving at the two ears. MSO neurons evoke a strongest response at their best delay (BD), at which the internal delay between bilateral inputs to MSO matches the external ITD. We performed extracellular recordings in the superior olivary complex of the anesthetized gerbil and found a majority of single units localized to the MSO to exhibit BDs that shifted with tone frequency. The relation of best interaural phase difference to tone frequency revealed nonlinearities in some MSO units and others with linear relations with characteristic phase between 0.4 and 0.6 cycles. The latter is usually associated with the interaction of ipsilateral excitation and contralateral inhibition, as in the lateral superior olive, yet all MSO units exhibited evidence of bilateral excitation. Interaural cochlear delays and phase-locked contralateral inhibition are two mechanisms of internal delay that have been suggested to create frequency-dependent delays. Best interaural phase-frequency relations were compared with a cross-correlation model of MSO that incorporated interaural cochlear delays and an additional frequency-independent delay component. The model with interaural cochlear delay fit phase-frequency relations exhibiting frequency-dependent delays with precision. Another model of MSO incorporating inhibition based on realistic biophysical parameters could not reproduce observed frequency-dependent delays.

Envelope coding in the lateral superior olive. II. Characteristic delays and comparison with responses in the medial superior olive

1996 ◽  
Vol 76 (4) ◽  
pp. 2137-2156 ◽  
Author(s):  
P. X. Joris
Keyword(s):  
Modulation Frequency ◽  
Small Sample ◽  
Spike Rate ◽  
Common Mechanism ◽  
Half Cycle ◽  
Medial Superior Olive ◽  
Lateral Superior Olive ◽  
Superior Olive ◽  
Binaural Interaction ◽  
Interaural Phase

1. Spike rates of cells in the cat's lateral superior olive (LSO) depend on interaural level differences (ILDs) and envelope interaural time differences (ITDs) of amplitude-modulated tones presented to both ears. We previously proposed that these sensitivities arise from a common mechanism, which is the IE binaural interaction (Inhibited by the contralateral and Excited by the ipsilateral ear). As a further test of that proposal and to gain a better understanding of the importance of this ITD-sensitivity, responses to monaural and binaural modulation are compared here over a range of modulation frequencies. 2. At low modulation frequencies, LSO-IE cells respond maximally when the envelopes of the amplitude-modulated stimuli at the two ears are out-of-phase by a half-cycle. This phase difference changes in a systematic way, which varies from cell to cell, when modulation frequency is increased. Mean interaural phase, measured over a range of modulation frequencies, was subjected to a characteristic delay analysis. Two measures were extracted: characteristic delay, which reflects differences in conduction delay between ipsi- and contralateral pathways, and characteristic phase, which reflects their sign of interaction. Most characteristic delays were within the physiological range of ITDs. There was a small bias toward positive delays, indicating a longer conduction time for the contralateral pathway. Characteristic phases were tightly distributed approximately 0.5 cycles, consistent with the proposed IE mechanism for ITD-sensitivity. 3. Increases in the modulation frequency of binaural stimuli beyond approximately 300 Hz consistently caused a profound decrease in average spike rate, as well as a decrease in the modulation of spike rate by ITD. The upper limit of ITD-sensitivity was 800 Hz. Sensitivity to envelope ITDs therefore is limited to a much lower range of frequencies than sensitivity to ITDs in fine-structure, e.g., as found in the medial superior olive (MSO), which operates up to several kilo Hertz. 4. A small sample of high-frequency EE cells (excited by both ears) in MSO also was tested with binaural amplitude-modulated stimuli. MSO-EE cells showed weak envelope ITD-sensitivity over a limited range of modulation frequencies. Consistent with the EE interaction, characteristic phases clustered approximately 0 cycles. 5. Mean interaural phase was compared with the phase of responses to monaural modulation. The difference between the ipsilateral and contralateral phases correlated well with the phase measured binaurally, both for LSO and MSO cells. 6. Many features of LSO-IE responses were mimicked by the simplest possible computer model, consisting of subtraction and rectification of low-pass filtered envelope waveforms. Differences between model and physiological results are suggestive of a temporal limitation in the binaural interaction that creates the ITD-sensitivity. 7. These results provide additional evidence for LSO ITD-sensitivity paralleling human psychophysical results. The stimulus boundaries within which ITD-sensitivity occurs suggest that it has a limited role in free-field conditions. It is traditionally thought that, to contribute to the perceived change in spatial location of a sound source, the LSO needs to show a change in overall firing rate summed across cells. This is achieved with small ILDs, but requires large ITDs, because the latter cue is less potent in single cells and has varied effects across cells by virtue of differences in characteristic delay.

Interaural time sensitivity in medial superior olive of cat

1990 ◽  
Vol 64 (2) ◽  
pp. 465-488 ◽  
Author(s):  
T. C. Yin ◽  
J. C. Chan
Keyword(s):  
Broad Band ◽  
Low Frequency ◽  
Sound Field ◽  
Phase Changes ◽  
Medial Superior Olive ◽  
Superior Olive ◽  
Band Noise ◽  
Interaural Phase ◽  
Combining Data ◽  
Monaural Stimulation

1. We studied the sensitivity of cells in the medial superior olive (MSO) of the anesthetized cat to variations in interaural phase differences (IPDs) of low-frequency tones and in interaural time differences (ITDs) of tones and broad-band noise signals. Our sample consisted of 39 cells histologically localized to the MSO. 2. All but one of the cells had characteristic frequencies less than 3 kHz, and 79% were sensitive to ITDs and IPDs. More than one-half (56%) of the cells responded to monaural stimulation of either ear, and both the binaural and monaural responses were highly phase locked. All of the cells that were sensitive to IPDs and monaurally driven by either ear responded in accord with that predicted by the coincidence model of Jeffress, as judged by comparisons of the phases at which the monaural and binaural responses occurred. The optimal IPDs were tightly clustered between 0.0 and 0.2 cycles. Most cells exhibited facilitation of the response at favorable ITDs and inhibition at unfavorable ITDs compared with the monaural responses. 3. Cells in the MSO exhibited characteristic delay, as judged by a linear relationship between the mean interaural phase and stimulating frequency. Characteristic phases were clustered near 0 indicating the most cells responded maximally when the two input tones were in phase. With the use of the binaural beat stimulus we found no differential selectivity for either the direction or speed of interaural phase changes. 4. The cells were also sensitive to ITDs of broad-band noise signals. The ITD curve in response to broad-band noise was similar to that predicted by the composite curve, which was calculated by linearly summating the tonal responses over the frequencies in the response area of the cell. Most (93%) of the peaks of the composite curves were between 0 and +400 microseconds, corresponding to locations in the contralateral sound field. Moreover, computer cross correlations of the monaural spike trains were similar to the ITD curve generated binaurally for both correlated and uncorrelated noise signals to the two ears. Thus our data suggest that the cells in the MSO behave much like cross-correlators. 5. By combining data from different animals and lcoating each cell on a standard MSO, we found evidence for a spatial map of ITDs across the anterior-posterior (A-P) axis of the MSO.(ABSTRACT TRUNCATED AT 400 WORDS)

scholarly journals Physiological and anatomical investigation of the auditory brainstem in the Fat-tailed Dunnart (Sminthopsis crassicaudata)

2019 ◽  
Author(s):  
Andrew Garrett ◽  
Virginia Lannigan ◽  
Nathanael Yates ◽  
Jennifer Rodger ◽  
Wilhelmina Mulders
Keyword(s):  
Auditory System ◽  
Mammalian Species ◽  
Auditory Brainstem ◽  
Auditory Brainstem Responses ◽  
Superior Olivary Complex ◽  
Medial Superior Olive ◽  
Superior Olive ◽  
Medial Nucleus ◽  
Sminthopsis Crassicaudata ◽  
Nissl Stain

The fat-tailed Dunnart (Sminthopsis crassicaudata) is a small (10-20g) native marsupial endemic to the south west of Western Australia. Currently little is known about the auditory capabilities of the dunnart, and of marsupials in general. Consequently, this study sought to investigate several electrophysiological and anatomical properties of the dunnart auditory system. Auditory brainstem responses (ABR) were recorded to brief (5ms) tone pips at a range of frequencies (4-47.5 kHz) and intensities to determine auditory brainstem thresholds. The dunnart ABR displayed multiple distinct peaks at all test frequencies, similar to other mammalian species. ABR showed the dunnart is most sensitive to higher frequencies increasing up to 47.5 kHz. Morphological observations (Nissl stain) revealed that the auditory structures thought to contribute to the first peaks of the ABR were all distinguishable in the dunnart. Structures identified include the dorsal and ventral subdivisions of the cochlear nucleus, including a cochlear nerve root nucleus as well as several distinct nuclei in the superior olivary complex, such as the medial nucleus of the trapezoid body, lateral superior olive and medial superior olive. This study is the first to show functional and anatomical aspects of the lower part of the auditory system in the Fat-tailed Dunnart.

scholarly journals Auditory brainstem models: adapting cochlear nuclei improve spatial encoding by the medial superior olive in reverberation

2019 ◽  
Author(s):  
Andrew Brughera ◽  
Jason Mikiel-Hunter ◽  
Mathias Dietz ◽  
David McAlpine
Keyword(s):  
Spatial Information ◽  
Interaural Time Difference ◽  
Auditory Pathway ◽  
Auditory Brainstem ◽  
Coincidence Detection ◽  
Type Model ◽  
Medial Superior Olive ◽  
Sound Energy ◽  
Superior Olive ◽  
Cochlear Nuclei

AbstractListeners perceive sound-energy as originating from the direction of its source, even as direct sound is followed milliseconds later by reflected sound from multiple different directions. Early-arriving sound is emphasised in the ascending auditory pathway, including the medial superior olive (MSO) where binaural neurons encode the interaural time difference (ITD) cue for spatial location. Behaviourally, weighting of ITD conveyed during rising sound-energy is stronger at 600 Hz, a frequency with higher reverberant energy, than at 200 Hz where reverberant energy is lower. Here we computationally explore the combined effectiveness of adaptation before ITD-encoding, and excitatory binaural coincidence detection within MSO neurons, in emphasising ITD conveyed in early-arriving sound. With excitatory inputs from adapting model spherical bushy cells (SBCs) of the bilateral cochlear nuclei, a Hodgkin-Huxley-type model MSO neuron reproduces the frequency-dependent emphasis of rising vs. peak sound-energy in ITD-encoding. Maintaining the adaptation in model SBCs, and adjusting membrane speed in model MSO neurons, hemispheric populations of model SBCs and MSO neurons, with simplified membranes for computational efficiency, also reproduce the stronger weighting of ITD information conveyed during rising sound-energy at 600 Hz compared to 200 Hz. This hemispheric model further demonstrates a link between strong weighting of spatial information during rising sound-energy, and correct unambiguous lateralisation of reverberant speech.

Interaural time and intensity coding in superior olivary complex and inferior colliculus of the echolocating bat Molossus ater

1985 ◽  
Vol 53 (1) ◽  
pp. 89-109 ◽  
Author(s):  
G. Harnischfeger ◽  
G. Neuweiler ◽  
P. Schlegel
Keyword(s):  
Inferior Colliculus ◽  
Stimulus Frequency ◽  
Stimulus Onset ◽  
Time Difference ◽  
Superior Olivary Complex ◽  
Inhibitory Input ◽  
Medial Superior Olive ◽  
Transient Time ◽  
Interaural Time Differences ◽  
Superior Olive

Single-unit responses to tonal stimulation with interaural disparities were recorded in the nuclei of the superior olivary complex (SOC) and the central nucleus of the inferior colliculus (ICC) of the echolocating bat, Molossus ater. Seventy-six units were recorded from the ICC and 74 from the SOC; of the SOC units, 31 were histologically verified in the medial superior olive (MSO), 10 in the lateral superior olive (LSO), and 33 in unidentified areas of the SOC. Best frequencies (BFs) of the units ranged from 10.3 to 89.6 kHz, and Q10 dB values ranged from 2 to 70 dB. Most ICC neurons responded phasically to stimulus onset and were either inhibitory/excitatory [I/E; (53)] or excitatory/excitatory [E/E; (21)] units. In the MSO, 23 units responded tonically and 7 phasically on, 18 were E/E or E/OF (facilitatory for other input) units, and 11 were I/E neurons. All LSO neurons responded in a "chopper" fashion, and the binaural neurons were E/I units. In E/E units the excitatory response to binaural stimulation was frequently larger than the sum of the monaurally evoked responses. Many neurons with E/I or I/E inputs had very steep binaural impulse-count functions and were sensitive to small interaural intensity differences. Twenty-eight units (24%) responded with a change in firing rate of at least 20% to interaural time differences of +/- 500 microseconds. Within this sample, 11 units (8 from ICC, 2 from MSO, and 1 from SOC) were sensitive to interaural time differences of only +/- 50 microseconds. Of these 11 units, 10 were I/E units responding phasically only to stimulus onset and were also sensitive to intensity differences (delta I), being suppressed completely by the inhibitory input over a delta I range of 20 dB or less. Of 117 units tested in the ICC and SOC nuclei, 86 units (76%) were not sensitive to interaural time disparities within +/- 500 microseconds. Because the BFs of these units sensitive to interaural transient time differences (delta t) ranged between 18 and 90 kHz, responses were elicited by pure tones, and responses did not change periodically with the period equal to that of the stimulus frequency, we conclude that the neurons reacted to interaural differences of stimulus-onset time (transient time difference) but not to phase differences (ongoing time difference). Sensitivity to interaural time differences was also correlated with interaural intensity differences.(ABSTRACT TRUNCATED AT 400 WORDS)

Functional organization of lateral cell groups of cat superior olivary complex

1977 ◽  
Vol 40 (2) ◽  
pp. 296-318 ◽  
Author(s):  
C. Tsuchitani
Keyword(s):  
Superior Olivary Complex ◽  
Maximum Output ◽  
Medial Superior Olive ◽  
Superior Olive ◽  
Response Characteristics ◽  
Cell Groups ◽  
Discharge Patterns ◽  
Intensity Functions ◽  
Contralateral Stimulus ◽  
Stimulation Of

1. Single-unit discharges to auditory stimuli were recorded extracellularly from superior olivary complex (SOC) units located lateral to the medial superior olive. Stimuli consisted of monaurally or binaurally presented tone bursts. The response measures obtained were effective ear, nature of effect, stimulus-frequency representation, maximum output, latency of response, and temporal pattern of tone burst-elicited discharges. Electrolytic marks were made at the unit studied or at the end of the electrode tract and in the medial superior olive. Following each experiment the locations of the units studied were determined histologically. An atlas of the laterally located SOC cell groups was developed to permit classification of units on the basis of localization within cell groups. Units were also classified according to the effects of stimulating the two ears. 2. All SOC units located lateral to the medial superior olive were excited by stimulation of the ipsilateral ear. Stimulation of the contralateral ear either excited, inhibited, had no effect, or had a potentiating effect on the discharges elicited by stimulating the ipsilateral ear. 3. Most lateral superior olivary (LSO) units were inhibited by contralateral stimulation, were narrowly tuned, produced low to high levels of maximum output, had short latencies, and produced regular discharge patterns characterized by chopper PST histograms with narrow initial peaks. 4. Most units within the caudal margins of the LSO (pLSO) were not affected or were inhibited by a contralateral stimulus; many were broadly tuned and exhibited intensity functions with large dynamic range and low slope. These units also had long latencies and produced chopper PST histograms with wide initial peaks. 5. Most units located dorsal to the LSO (DPO and DLPO) were not affected by the contralateral stimulus, were narrowly tuned, produced moderate levels of maximum discharge, had long latencies, and produced chopper PST histograms with wide initial peaks. 6. Units located ventral to the LSO appeared to have response characteristics related to unit location. Most units below the ventral hilum of the LSO (VLPO) were inhibited by the contralateral stimulus and many were broadly tuned VLPO units produced wide or poorly defined narrow-chopper discharge patterns and intensity functions with high maximum output. Most units located ventral to the lateral loop of the LSO (LNTB) were not affected by the contralateral stimulus and had response characteristics that may be related to the rostrocaudal location of the unit. 7. The cell groups located dorsal and ventral to the LSO were tonotopically organized with low-frequency-sensitive units located laterally and high-frequency-sensitive units located medially. The units located along the caudal margins of the LSO had a tonotopic organization similar to that of the LSO.

Binaural Response Properties of Low-Frequency Neurons in the Gerbil Dorsal Nucleus of the Lateral Lemniscus

2006 ◽  
Vol 96 (3) ◽  
pp. 1425-1440 ◽  
Author(s):  
Ida Siveke ◽  
Michael Pecka ◽  
Armin H. Seidl ◽  
Sylvie Baudoux ◽  
Benedikt Grothe
Keyword(s):  
Low Frequency ◽  
Superior Olivary Complex ◽  
Suitable Model ◽  
Medial Superior Olive ◽  
Lateral Lemniscus ◽  
Auditory Midbrain ◽  
Response Properties ◽  
Superior Olive ◽  
Dorsal Nucleus ◽  
Anatomical Evidence

Differences in intensity and arrival time of sounds at the two ears, interaural intensity and time differences (IID, ITD), are the chief cues for sound localization. Both cues are initially processed in the superior olivary complex (SOC), which projects to the dorsal nucleus of the lateral lemniscus (DNLL) and the auditory midbrain. Here we present basic response properties of low-frequency (<2 kHz) DNLL neurons and their binaural sensitivity to ITDs and IIDs in the anesthetized gerbil. We found many neurons showing binaural properties similar to those reported for SOC neurons. IID-properties were similar to that of the contralateral lateral superior olive (LSO). A majority of cells had an ITD sensitivity resembling that of either the ipsilateral medial superior olive (MSO) or the contralateral LSO. A smaller number of cells displayed intermediate types of ITD sensitivity. In neurons with MSO-like response ITDs that evoked maximal discharges were mostly outside of the range of ITDs the gerbil naturally experiences. The maxima of the first derivative of their ITD-functions (steepest slope), however, were well within the physiological range of ITDs. This finding is consistent with the concept of a population rather than a place code for ITDs. Moreover, we describe several other binaural properties as well as physiological and anatomical evidence for a small but significant input from the contralateral MSO. The large number of ITD-sensitive low-frequency neurons implicates a substantial role for the DNLL in ITD processing and promotes this nucleus as a suitable model for further studies on ITD-coding.

scholarly journals Physiological and anatomical investigation of the auditory brainstem in the Fat-tailed Dunnart (Sminthopsis crassicaudata)

2019 ◽  
Author(s):  
Andrew Garrett ◽  
Virginia Lannigan ◽  
Nathanael Yates ◽  
Jennifer Rodger ◽  
Wilhelmina Mulders
Keyword(s):  
Auditory System ◽  
Mammalian Species ◽  
Auditory Brainstem ◽  
Auditory Brainstem Responses ◽  
Superior Olivary Complex ◽  
Medial Superior Olive ◽  
Superior Olive ◽  
Medial Nucleus ◽  
Sminthopsis Crassicaudata ◽  
Nissl Stain

The fat-tailed Dunnart (Sminthopsis crassicaudata) is a small (10-20g) native marsupial endemic to the south west of Western Australia. Currently little is known about the auditory capabilities of the dunnart, and of marsupials in general. Consequently, this study sought to investigate several electrophysiological and anatomical properties of the dunnart auditory system. Auditory brainstem responses (ABR) were recorded to brief (5ms) tone pips at a range of frequencies (4-47.5 kHz) and intensities to determine auditory brainstem thresholds. The dunnart ABR displayed multiple distinct peaks at all test frequencies, similar to other mammalian species. ABR showed the dunnart is most sensitive to higher frequencies increasing up to 47.5 kHz. Morphological observations (Nissl stain) revealed that the auditory structures thought to contribute to the first peaks of the ABR were all distinguishable in the dunnart. Structures identified include the dorsal and ventral subdivisions of the cochlear nucleus, including a cochlear nerve root nucleus as well as several distinct nuclei in the superior olivary complex, such as the medial nucleus of the trapezoid body, lateral superior olive and medial superior olive. This study is the first to show functional and anatomical aspects of the lower part of the auditory system in the Fat-tailed Dunnart.

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