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1.  Sensitivity to Interaural Time Differences in the Inferior Colliculus with Bilateral Cochlear Implants 
Bilateral cochlear implantation attempts to increase performance over a monaural prosthesis by harnessing the binaural processing of the auditory system. Although many bilaterally implanted human subjects discriminate interaural time differences (ITDs), a major cue for sound localization and signal detection in noise, their performance is typically poorer than that of normal-hearing listeners. We developed an animal model of bilateral cochlear implantation to study neural ITD sensitivity for trains of electric current pulses delivered via bilaterally implanted intracochlear electrodes. We found that a majority of single units in the inferior colliculus of acutely deafened, anesthetized cats are sensitive to ITD and that electric ITD tuning is as sharp as found for acoustic stimulation with broadband noise in normal-hearing animals. However, the sharpness and shape of ITD tuning often depended strongly on stimulus intensity; some neurons had dynamic ranges of ITD sensitivity as low as 1 dB. We also found that neural ITD sensitivity was best at pulse rates below 100 Hz and decreased with increasing pulse rate. This rate limitation parallels behavioral ITD discrimination in bilaterally implanted individuals. The sharp neural ITD sensitivity found with electric stimulation at the appropriate intensity is encouraging for the prospect of restoring the functional benefits of binaural hearing in bilaterally implanted human subjects and suggests that neural plasticity resulting from previous deafness and deprivation of binaural experience may play a role in the poor ITD discrimination with current bilateral implants.
doi:10.1523/JNEUROSCI.0052-07.2007
PMCID: PMC2041852  PMID: 17581961
binaural hearing; electric stimulation; neural prosthesis; cochlear implant; inferior colliculus; ITD
2.  Preservation of Spectrotemporal Tuning Between the Nucleus Laminaris and the Inferior Colliculus of the Barn Owl 
Journal of neurophysiology  2007;97(5):3544-3553.
Performing sound recognition is a task that requires an encoding of the time-varying spectral structure of the auditory stimulus. Similarly, computation of the interaural time difference (ITD) requires knowledge of the precise timing of the stimulus. Consistent with this, low-level nuclei of birds and mammals implicated in ITD processing encode the ongoing phase of a stimulus. However, the brain areas that follow the binaural convergence for the computation of ITD show a reduced capacity for phase locking. In addition, we have shown that in the barn owl there is a pooling of ITD-responsive neurons to improve the reliability of ITD coding. Here we demonstrate that despite two stages of convergence and an effective loss of phase information, the auditory system of the anesthetized barn owl displays a graceful transition to an envelope coding that preserves the spectrotemporal information throughout the ITD pathway to the neurons of the core of the central nucleus of the inferior colliculus.
doi:10.1152/jn.01162.2006
PMCID: PMC2532515  PMID: 17314241
3.  Multiplicative Auditory Spatial Receptive Fields Created by a Hierarchy of Population Codes 
PLoS ONE  2009;4(11):e8015.
A multiplicative combination of tuning to interaural time difference (ITD) and interaural level difference (ILD) contributes to the generation of spatially selective auditory neurons in the owl's midbrain. Previous analyses of multiplicative responses in the owl have not taken into consideration the frequency-dependence of ITD and ILD cues that occur under natural listening conditions. Here, we present a model for the responses of ITD- and ILD-sensitive neurons in the barn owl's inferior colliculus which satisfies constraints raised by experimental data on frequency convergence, multiplicative interaction of ITD and ILD, and response properties of afferent neurons. We propose that multiplication between ITD- and ILD-dependent signals occurs only within frequency channels and that frequency integration occurs using a linear-threshold mechanism. The model reproduces the experimentally observed nonlinear responses to ITD and ILD in the inferior colliculus, with greater accuracy than previous models. We show that linear-threshold frequency integration allows the system to represent multiple sound sources with natural sound localization cues, whereas multiplicative frequency integration does not. Nonlinear responses in the owl's inferior colliculus can thus be generated using a combination of cellular and network mechanisms, showing that multiple elements of previous theories can be combined in a single system.
doi:10.1371/journal.pone.0008015
PMCID: PMC2776990  PMID: 19956693
4.  Effects of reverberation on the directional sensitivity of auditory neurons across the tonotopic axis: Influences of ITD and ILD 
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 (ITD) are the only available directional cue, low-CF cells sensitive to ITD in the waveform fine time structure maintain better directional sensitivity in reverberation than high-CF cells sensitive to ITD in the envelope induced by cochlear filtering. On the other hand, 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.
doi:10.1523/JNEUROSCI.5517-09.2010
PMCID: PMC2896784  PMID: 20534831
directional sensitivity; interaural level difference; inferior colliculus; interaural time difference; reverberation; sound localization
5.  Behavioral Sensitivity to Interaural Time Differences in the Rabbit 
Hearing research  2007;235(1-2):134-142.
An important cue for sound localization and separation of signals from noise is the interaural time difference (ITD). Humans are able to localize sounds within 1–2° and can detect very small changes in the ITD (10–20 μs). In contrast, many animals localize sounds with less precision than humans. Rabbits, for example, have sound localization thresholds of ~22°. There is only limited information about behavioral ITD discrimination in animals with poor sound localization acuity that are typically used for the neural recordings. For this study, we measured behavioral discrimination of ITDs in the rabbit for a range of reference ITDs from 0 to ± 300 μs. The behavioral task was conditioned avoidance and the stimulus was band-limited noise (500–1500 Hz). Across animals, the average discrimination threshold was 50–60 μs for reference ITDs of 0 to ± 200 μs. There was no trend in the thresholds across this range of reference ITDs. For a reference ITD of ± 300 μs, which is near the limit of the physiological window defined by the head width in this species, the discrimination threshold increased to ~100 μs. The ITD discrimination in rabbits less acute than in cats, which have a similar head size. This result supports the suggestion that ITD discrimination, like sound localization (see Heffner, 1997, Acta Otolaryngol Suppl 532:46–53, 1997) is determined by factors other than head size.
doi:10.1016/j.heares.2007.11.003
PMCID: PMC2692955  PMID: 18093767
Sound localization; animal psychoacoustics; neural discrimination
6.  Sensitivity of Inferior Colliculus Neurons to Interaural Time Differences in the Envelope Versus the Fine Structure With Bilateral Cochlear Implants 
Journal of neurophysiology  2008;99(5):2390-2407.
Bilateral cochlear implantation seeks to improve hearing by taking advantage of the binaural processing of the central auditory system. Cochlear implants typically encode sound in each spectral channel by amplitude modulating (AM) a fixed-rate pulse train, thus interaural time differences (ITD) are only delivered in the envelope. We investigated the ITD sensitivity of inferior colliculus (IC) neurons with sinusoidally AM pulse trains. ITD was introduced independently to the AM and/or carrier pulses to measure the relative efficacy of envelope and fine structure for delivering ITD information. We found that many IC cells are sensitive to ITD in both the envelope (ITDenv) and fine structure (ITDfs) for appropriate modulation frequencies and carrier rates. ITDenv sensitivity was generally similar to that seen in normal-hearing animals with AM tones. ITDenv tuning generally improved with increasing modulation frequency up to the maximum modulation frequency that elicited a sustained response in a neuron (tested ≤Hz). ITDfs sensitivity was present in about half the neurons for 1,000 pulse/s (pps) carriers and was nonexistent at 5,000 pps. The neurons that were sensitive to ITDfs at 1,000 pps were those that showed the best ITD sensitivity to low-rate pulse trains. Overall, the best ITD sensitivity was found for ITD contained in the fine structure of a moderate rate AM pulse train (1,000 pps). These results suggest that the interaural timing of current pulses should be accurately controlled in a bilateral cochlear implant processing strategy that provides salient ITD cues.
doi:10.1152/jn.00751.2007
PMCID: PMC2570106  PMID: 18287556
7.  Detection of Large Interaural Delays and Its Implication for Models of Binaural Interaction  
The interaural time difference (ITD) is a major cue to sound localization along the horizontal plane. The maximum natural ITD occurs when a sound source is positioned opposite to one ear. We examined the ability of owls and humans to detect large ITDs in sounds presented through headphones. Stimuli consisted of either broad or narrow bands of Gaussian noise, 100 ms in duration. Using headphones allowed presentation of ITDs that are greater than the maximum natural ITD. Owls were able to discriminate a sound leading to the left ear from one leading to the right ear, for ITDs that are 5 times the maximum natural delay. Neural recordings from optic-tectum neurons, however, show that best ITDs are usually well within the natural range and are never as large as ITDs that are behaviorally discriminable. A model of binaural cross-correlation with short delay lines is shown to explain behavioral detection of large ITDs. The model uses curved trajectories of a cross-correlation pattern as the basis for detection. These trajectories represent side peaks of neural ITD-tuning curves and successfully predict localization reversals by both owls and human subjects.
doi:10.1007/s101620020006
PMCID: PMC3202365  PMID: 12083726
interaural; binaural; owl; ITD
8.  Optical Imaging of Interaural Time Difference Representation in Rat Auditory Cortex 
We used in vivo voltage-sensitive dye optical imaging to examine the cortical representation of interaural time difference (ITD), which is believed to be involved in sound source localization. We found that acoustic stimuli with dissimilar ITD activate various localized domains in the auditory cortex. The main loci of the activation pattern shift up to 1 mm during the first 40 ms of the response period. We suppose that some of the neurons in each pool are sensitive to the definite ITD and involved in the transduction of information about sound source localization, based on the ITD. This assumption gives a reasonable fit to the Jeffress model in which the neural network calculates the ITD to define the direction of the sound source. Such calculation forms the basis for the cortex's ability to detect the azimuth of the sound source.
doi:10.3389/neuro.16.002.2009
PMCID: PMC2654020  PMID: 19277218
auditory cortex; interaural time difference; optical imaging; voltage-sensitive dye
9.  Emergence of Multiplicative Auditory Responses in the Midbrain of the Barn Owl 
Journal of neurophysiology  2007;98(3):1181-1193.
Space-specific neurons in the barn owl’s auditory space map gain spatial selectivity through tuning to combinations of the interaural time difference (ITD) and interaural level difference (ILD). The combination of ITD and ILD in the subthreshold responses of space-specific neurons in the external nucleus of the inferior colliculus (ICx) is well described by a multiplication of ITD- and ILD-dependent components. It is unknown, however, how ITD and ILD are combined at the site of ITD and ILD convergence in the lateral shell of the central nucleus of the inferior colliculus (ICcl) and therefore whether ICx is the first site in the auditory pathway where multiplicative tuning to ITD-and ILD-dependent signals occurs. We used extracellular re-cording of single neurons to determine how ITD and ILD are combined in ICcl of the anesthetized barn owl (Tyto alba). A comparison of additive, multiplicative, and linear-threshold models of neural responses shows that ITD and ILD are combined nonlinearly in ICcl, but the interaction of ITD and ILD is not uniformly multiplicative over the sample. A subset (61%) of the neural responses is well described by the multiplicative model, indicating that ICcl is the first site where multiplicative tuning to ITD- and ILD-dependent signals occurs. ICx, however, is the first site where multiplicative tuning is observed consistently. A network model shows that a linear combination of ICcl responses to ITD–ILD pairs is sufficient to produce the multiplicative subthreshold responses to ITD and ILD seen in ICx.
doi:10.1152/jn.00370.2007
PMCID: PMC2532518  PMID: 17615132
10.  Phase Locking of Auditory-Nerve Fibers to the Envelopes of High-Frequency Sounds: Implications for Sound Localization 
Journal of neurophysiology  2006;96(5):2327-2341.
Although listeners are sensitive to interaural time differences (ITDs) in the envelope of high-frequency sounds, both ITD discrimination performance and the extent of lateralization are poorer for high-frequency sinusoidally amplitude-modulated (SAM) tones than for low-frequency pure tones. Psychophysical studies have shown that ITD discrimination at high frequencies can be improved by using novel transposed-tone stimuli, formed by modulating a high-frequency carrier by a half-wave–rectified sinusoid. Transposed tones are designed to produce the same temporal discharge patterns in high-characteristic frequency (CF) neurons as occur in low-CF neurons for pure-tone stimuli. To directly test this hypothesis, we compared responses of auditory-nerve fibers in anesthetized cats to pure tones, SAM tones, and transposed tones. Phase locking was characterized using both the synchronization index and autocorrelograms. With both measures, phase locking was better for transposed tones than for SAM tones, consistent with the rationale for using transposed tones. However, phase locking to transposed tones and that to pure tones were comparable only when all three conditions were met: stimulus levels near thresholds, low modulation frequencies (<250 Hz), and low spontaneous discharge rates. In particular, phase locking to both SAM tones and transposed tones substantially degraded with increasing stimulus level, while remaining more stable for pure tones. These results suggest caution in assuming a close similarity between temporal patterns of peripheral activity produced by transposed tones and pure tones in both psychophysical studies and neurophysiological studies of central neurons.
doi:10.1152/jn.00326.2006
PMCID: PMC2013745  PMID: 16807349
11.  Models of Brainstem Responses to Bilateral Electrical Stimulation 
A simple, biophysically specified cell model is used to predict responses of binaurally sensitive neurons to patterns of input spikes that represent stimulation by acoustic and electric waveforms. Specifically, the effects of changes in parameters of input spike trains on model responses to interaural time difference (ITD) were studied for low-frequency periodic stimuli, with or without amplitude modulation. Simulations were limited to purely excitatory, bilaterally driven cell models with basic ionic currents and multiple input fibers. Parameters explored include average firing rate, synchrony index, modulation frequency, and latency dispersion of the input trains as well as the excitatory conductance and time constant of individual synapses in the cell model. Results are compared to physiological recordings from the inferior colliculus (IC) and discussed in terms of ITD-discrimination abilities of listeners with cochlear implants. Several empirically observed aspects of ITD sensitivity were simulated without evoking complex neural processing. Specifically, our results show saturation effects in rate–ITD curves, the absence of sustained responses to high-rate unmodulated pulse trains, the renewal of sensitivity to ITD in high-rate trains when inputs are amplitude-modulated, and interactions between envelope and fine-structure delays for some modulation frequencies.
doi:10.1007/s10162-008-0141-z
PMCID: PMC2644392  PMID: 18941838
binaural hearing; auditory brainstem model; electric hearing; interaural time delay; cochlear implant
12.  Frequency-Invariant Representation of Interaural Time Differences in Mammals 
PLoS Computational Biology  2011;7(3):e1002013.
Interaural time differences (ITDs) are the major cue for localizing low-frequency sounds. The activity of neuronal populations in the brainstem encodes ITDs with an exquisite temporal acuity of about . The response of single neurons, however, also changes with other stimulus properties like the spectral composition of sound. The influence of stimulus frequency is very different across neurons and thus it is unclear how ITDs are encoded independently of stimulus frequency by populations of neurons. Here we fitted a statistical model to single-cell rate responses of the dorsal nucleus of the lateral lemniscus. The model was used to evaluate the impact of single-cell response characteristics on the frequency-invariant mutual information between rate response and ITD. We found a rough correspondence between the measured cell characteristics and those predicted by computing mutual information. Furthermore, we studied two readout mechanisms, a linear classifier and a two-channel rate difference decoder. The latter turned out to be better suited to decode the population patterns obtained from the fitted model.
Author Summary
Neuronal codes are usually studied by estimating how much information the brain activity carries about the stimulus. On a single cell level, the relevant features of neuronal activity such as the firing rate or spike timing are readily available. On a population level, where many neurons together encode a stimulus property, finding the most appropriate activity features is less obvious, particularly because the neurons respond with a huge cell-to-cell variability. Here, using the example of the neuronal representation of interaural time differences, we show that the quality of the population code strongly depends on the assumption — or the model — of the population readout. We argue that invariances are useful constraints to identify “good” population codes. Based on these ideas, we suggest that the representation of interaural time differences serves a two-channel code in which the difference between the summed activities of the neurons in the two hemispheres exhibits an invariant and linear dependence on interaural time difference.
doi:10.1371/journal.pcbi.1002013
PMCID: PMC3060160  PMID: 21445227
13.  Noise Reduction of Coincidence Detector Output by the Inferior Colliculus of the Barn Owl 
A recurring theme in theoretical work is that integration over populations of similarly tuned neurons can reduce neural noise. However, there are relatively few demonstrations of an explicit noise reduction mechanism in a neural network. Here we demonstrate that the brainstem of the barn owl includes a stage of processing apparently devoted to increasing the signal-to-noise ratio in the encoding of the interaural time difference (ITD), one of two primary binaural cues used to compute the position of a sound source in space. In the barn owl, the ITD is processed in a dedicated neural pathway that terminates at the core of the inferior colliculus (ICcc). The actual locus of the computation of the ITD is before ICcc in the nucleus laminaris (NL), and ICcc receives no inputs carrying information that did not originate in NL. Unlike in NL, the rate-ITD functions of ICcc neurons require as little as a single stimulus presentation per ITD to show coherent ITD tuning. ICcc neurons also displayed a greater dynamic range with a maximal difference in ITD response rates approximately double that seen in NL. These results indicate that ICcc neurons perform a computation functionally analogous to averaging across a population of similarly tuned NL neurons.
doi:10.1523/JNEUROSCI.0220-06.2006
PMCID: PMC2492673  PMID: 16738236
interaural time difference; sound localization; inferior colliculus; nucleus laminaris; barn owl; pooling
14.  Cross-Correlation in the Auditory Coincidence Detectors of Owls 
Interaural time difference (ITD) plays a central role in many auditory functions, most importantly in sound localization. The classic model for how ITD is computed was put forth by Jeffress (1948). One of the predictions of the Jeffress model is that the neurons that compute ITD should behave as cross-correlators. Whereas cross-correlation-like properties of the ITD-computing neurons have been reported, attempts to show that the shape of the ITD response function is determined by the spectral tuning of the neuron, a core prediction of cross-correlation, have been unsuccessful. Using reverse correlation analysis, we demonstrate in the barn owl that the relationship between the spectral tuning and the ITD response of the ITD-computing neurons is that predicted by cross-correlation. Moreover, we show that a model of coincidence detector responses derived from responses to binaurally uncorrelated noise is consistent with binaural interaction based on cross-correlation. These results are thus consistent with one of the key tenets of the Jeffress model. Our work sets forth both the methodology to answer whether cross-correlation describes coincidence detector responses and a demonstration that in the barn owl, the result is that expected by theory.
doi:10.1523/JNEUROSCI.1969-08.2008
PMCID: PMC2637928  PMID: 18685035
barn owl; interaural time difference; cross-correlation; coincidence detection; sound localization; nucleus laminaris
15.  Contributions of Intrinsic Neural and Stimulus Variance to Binaural Sensitivity 
The discrimination of a change in a stimulus is determined both by the magnitude of that change and by the variability in the neural response to the stimulus. When the stimulus is itself noisy, then the relative contributions of the neural (intrinsic) and stimulus induced variability becomes a critical question. We measured the contribution of intrinsic neural noise and interstimulus variability to the discrimination of interaural time differences (ITDs) and interaural correlation (IAC). We measured discharge rate versus characteristic frequency (CF) tone ITD functions, and CF-centered narrowband noise ITD and IAC functions in interleaved blocks in the same units in the inferior colliculus of urethane-anesthetized guinea pigs. Ten “frozen” tokens of noise were synthesized and the responses to each token were separately analyzed to allow the relative contributions of intrinsic and stimulus variability to be assessed. ITD and IAC discrimination thresholds were determined for a simulated two-interval forced-choice experiment, based on the firing rate distributions, using receiver operating characteristic analysis. On average, between stimulus variability contributed 19% (range, 1.5–30%) of the variance in noise ITD discrimination and 27% (range, 3–50%) in IAC discrimination. Noise ITD thresholds were slightly higher than tone ITD thresholds. Taking the mean of the thresholds for individual noise tokens gave a similar result to pooling across all noise tokens. This implies that although the stimulus induced variability is measurable, it is insignificant in relation to the intrinsic noise in ITD and IAC discrimination.
doi:10.1007/s10162-006-0054-7
PMCID: PMC2504630  PMID: 17053864
binaural; discrimination; guinea pig; inferior colliculus; interaural correlation; jnd; interaural time difference (ITD); intrinsic variability; stimulus variability
16.  Responses to Interaural Time Delay in Human Cortex 
Journal of Neurophysiology  2008;100(5):2712-2718.
Humans use differences in the timing of sounds at the two ears to determine the location of a sound source. Various models have been posited for the neural representation of these interaural time differences (ITDs). These models make opposing predictions about the lateralization of ITD processing in the human brain. The weighted-image model predicts that sounds leading in time at one ear activate maximally the opposite brain hemisphere for all values of ITD. In contrast, the π-limit model assumes that ITDs beyond half the period of the stimulus center frequency are not explicitly encoded in the brain and that such “long” ITDs activate maximally the side of the brain to which the sound is heard. A previous neuroimaging study revealed activity in the human inferior colliculus consistent with the π-limit. Here we show that cortical responses to sounds with ITDs within the π-limit are in line with the predictions of both models. However, contrary to the immediate predictions of both models, neural activation is bilateral for “long” ITDs, despite these being perceived as clearly lateralized. Furthermore, processing of long ITDs leads to higher activation in cortex than processing of short ITDs. These data show that coding of ITD in cortex is fundamentally different from coding of ITD in the brain stem. We discuss these results in the context of the two models.
doi:10.1152/jn.90210.2008
PMCID: PMC2585401  PMID: 18799604
17.  Responses of Auditory Nerve and Anteroventral Cochlear Nucleus Fibers to Broadband and Narrowband Noise: Implications for the Sensitivity to Interaural Delays 
The quality of temporal coding of sound waveforms in the monaural afferents that converge on binaural neurons in the brainstem limits the sensitivity to temporal differences at the two ears. The anteroventral cochlear nucleus (AVCN) houses the cells that project to the binaural nuclei, which are known to have enhanced temporal coding of low-frequency sounds relative to auditory nerve (AN) fibers. We applied a coincidence analysis within the framework of detection theory to investigate the extent to which AVCN processing affects interaural time delay (ITD) sensitivity. Using monaural spike trains to a 1-s broadband or narrowband noise token, we emulated the binaural task of ITD discrimination and calculated just noticeable differences (jnds). The ITD jnds derived from AVCN neurons were lower than those derived from AN fibers, showing that the enhanced temporal coding in the AVCN improves binaural sensitivity to ITDs. AVCN processing also increased the dynamic range of ITD sensitivity and changed the shape of the frequency dependence of ITD sensitivity. Bandwidth dependence of ITD jnds from AN as well as AVCN fibers agreed with psychophysical data. These findings demonstrate that monaural preprocessing in the AVCN improves the temporal code in a way that is beneficial for binaural processing and may be crucial in achieving the exquisite sensitivity to ITDs observed in binaural pathways.
doi:10.1007/s10162-011-0268-1
PMCID: PMC3123442  PMID: 21567250
coincidence detection; interaural time difference; discrimination; binaural; sound localization
18.  Responses of Auditory Nerve and Anteroventral Cochlear Nucleus Fibers to Broadband and Narrowband Noise: Implications for the Sensitivity to Interaural Delays 
The quality of temporal coding of sound waveforms in the monaural afferents that converge on binaural neurons in the brainstem limits the sensitivity to temporal differences at the two ears. The anteroventral cochlear nucleus (AVCN) houses the cells that project to the binaural nuclei, which are known to have enhanced temporal coding of low-frequency sounds relative to auditory nerve (AN) fibers. We applied a coincidence analysis within the framework of detection theory to investigate the extent to which AVCN processing affects interaural time delay (ITD) sensitivity. Using monaural spike trains to a 1-s broadband or narrowband noise token, we emulated the binaural task of ITD discrimination and calculated just noticeable differences (jnds). The ITD jnds derived from AVCN neurons were lower than those derived from AN fibers, showing that the enhanced temporal coding in the AVCN improves binaural sensitivity to ITDs. AVCN processing also increased the dynamic range of ITD sensitivity and changed the shape of the frequency dependence of ITD sensitivity. Bandwidth dependence of ITD jnds from AN as well as AVCN fibers agreed with psychophysical data. These findings demonstrate that monaural preprocessing in the AVCN improves the temporal code in a way that is beneficial for binaural processing and may be crucial in achieving the exquisite sensitivity to ITDs observed in binaural pathways.
doi:10.1007/s10162-011-0268-1
PMCID: PMC3123442  PMID: 21567250
coincidence detection; interaural time difference; discrimination; binaural; sound localization
19.  The representation of sound localization cues in the barn owl's inferior colliculus 
The barn owl is a well-known model system for studying auditory processing and sound localization. This article reviews the morphological and functional organization, as well as the role of the underlying microcircuits, of the barn owl's inferior colliculus (IC). We focus on the processing of frequency and interaural time (ITD) and level differences (ILD). We first summarize the morphology of the sub-nuclei belonging to the IC and their differentiation by antero- and retrograde labeling and by staining with various antibodies. We then focus on the response properties of neurons in the three major sub-nuclei of IC [core of the central nucleus of the IC (ICCc), lateral shell of the central nucleus of the IC (ICCls), and the external nucleus of the IC (ICX)]. ICCc projects to ICCls, which in turn sends its information to ICX. The responses of neurons in ICCc are sensitive to changes in ITD but not to changes in ILD. The distribution of ITD sensitivity with frequency in ICCc can only partly be explained by optimal coding. We continue with the tuning properties of ICCls neurons, the first station in the midbrain where the ITD and ILD pathways merge after they have split at the level of the cochlear nucleus. The ICCc and ICCls share similar ITD and frequency tuning. By contrast, ICCls shows sigmoidal ILD tuning which is absent in ICCc. Both ICCc and ICCls project to the forebrain, and ICCls also projects to ICX, where space-specific neurons are found. Space-specific neurons exhibit side peak suppression in ITD tuning, bell-shaped ILD tuning, and are broadly tuned to frequency. These neurons respond only to restricted positions of auditory space and form a map of two-dimensional auditory space. Finally, we briefly review major IC features, including multiplication-like computations, correlates of echo suppression, plasticity, and adaptation.
doi:10.3389/fncir.2012.00045
PMCID: PMC3394089  PMID: 22798945
sound localization; central nucleus of the inferior colliculus; auditory; plasticity; adaptation; interaural time difference; interaural level difference; frequency tuning
20.  Across-frequency combination of interaural time difference in bilateral cochlear implant listeners 
The current study examined how cochlear implant (CI) listeners combine temporally interleaved envelope-ITD information across two sites of stimulation. When two cochlear sites jointly transmit ITD information, one possibility is that CI listeners can extract the most reliable ITD cues available. As a result, ITD sensitivity would be sustained or enhanced compared to single-site stimulation. Alternatively, mutual interference across multiple sites of ITD stimulation could worsen dual-site performance compared to listening to the better of two electrode pairs. Two experiments used direct stimulation to examine how CI users can integrate ITDs across two pairs of electrodes. Experiment 1 tested ITD discrimination for two stimulation sites using 100-Hz sinusoidally modulated 1000-pps-carrier pulse trains. Experiment 2 used the same stimuli ramped with 100 ms windows, as a control condition with minimized onset cues. For all stimuli, performance improved monotonically with increasing modulation depth. Results show that when CI listeners are stimulated with electrode pairs at two cochlear sites, sensitivity to ITDs was similar to that seen when only the electrode pair with better sensitivity was activated. None of the listeners showed a decrement in performance from the worse electrode pair. This could be achieved either by listening to the better electrode pair or by truly integrating the information across cochlear sites.
doi:10.3389/fnsys.2014.00022
PMCID: PMC3949319  PMID: 24653681
cochlear implant; interaural time difference; envelope ITD; across-frequency integration; spatial sound
21.  Detection of Interaural Time Differences in the Alligator 
The auditory systems of birds and mammals use timing information from each ear to detect interaural time difference (ITD). To determine whether the Jeffress-type algorithms that underlie sensitivity to ITD in birds are an evolutionarily stable strategy, we recorded from the auditory nuclei of crocodilians, who are the sister group to the birds. In alligators, precisely timed spikes in the first-order nucleus magnocellularis (NM) encode the timing of sounds, and NM neurons project to neurons in the nucleus laminaris (NL) that detect interaural time differences. In vivo recordings from NL neurons show that the arrival time of phase-locked spikes differs between the ipsilateral and contralateral inputs. When this disparity is nullified by their best ITD, the neurons respond maximally. Thus NL neurons act as coincidence detectors. A biologically detailed model of NL with alligator parameters discriminated ITDs up to 1 kHz. The range of best ITDs represented in NL was much larger than in birds, however, and extended from 0 to 1000 μs contralateral, with a median ITD of 450 μs. Thus, crocodilians and birds employ similar algorithms for ITD detection, although crocodilians have larger heads.
doi:10.1523/JNEUROSCI.6154-08.2009
PMCID: PMC3170862  PMID: 19553438
22.  Neural coding of ITD with bilateral cochlear implants: Effects of congenital deafness 
Human bilateral cochlear implant users do poorly on tasks involving interaural time differences (ITD), a cue which provides important benefits to the normal hearing, especially in challenging acoustic environments. Yet the precision of neural ITD coding in acutely-deafened, bilaterally-implanted cats is essentially normal (Smith and Delgutte, J. Neurosci. 27:6740–6750). One explanation for this discrepancy is that the extended periods of binaural deprivation typically experienced by cochlear implant users degrades neural ITD sensitivity, either by impeding normal maturation of the neural circuitry or by altering it later in life. To test this hypothesis, we recorded from single units in inferior colliculus (IC) of two groups of bilaterally-implanted, anesthetized cats that contrast maximally in binaural experience: acutely-deafened cats, which had normal binaural hearing until experimentation, and congenitally deaf white cats, which received no auditory inputs until the experiment. Rate responses of only half as many neurons showed significant ITD sensitivity to low-rate pulse trains in congenitally deaf cats compared to acutely deafened cats. For neurons that were ITD sensitive, ITD tuning was broader and best ITDs were more variable in congenitally deaf cats, leading to poorer ITD coding within the naturally-occurring range. A signal detection model constrained by the observed physiology supports the idea that the degraded neural ITD coding resulting from deprivation of binaural experience contributes to poor ITD discrimination by human implantees.
doi:10.1523/JNEUROSCI.3213-10.2010
PMCID: PMC3025489  PMID: 20962228
binaural hearing; electric stimulation; congenital deafness; cochlear implant; inferior colliculus; ITD
23.  Trading of interaural differences in high-rate Gabor click trains 
Hearing research  2010;268(1-2):202-212.
In this study, combinations of interaraural time differences (ITD) and interaural level differences (ILD) were applied to trains of 4000 Hz Gabor clicks (Gaussian-filtered impulses) and presented to listeners over headphones. ITD / ILD equivalence functions, or “trading ratios” (TR) were estimated using two different procedures: a “closed-loop” procedure in which subjects adjusted (via head-turn) the ILD of a target click train to counteract the effects of an imposed ITD, and an “open-loop” procedure in which subjects indicated (also via head-turn) the lateral position of click trains containing independent combinations of ITD and ILD. For both tasks, TR values increasingly favored ILD over ITD as inter-click interval (ICI) decreased from 10 to 2 ms. Subsequent analysis confirmed that this change reflected a loss of sensitivity to envelope ITD at short ICI rather than a gain in sensitivity to ILD, consistent with prior studies demonstrating rate-limited processing of ongoing envelope ITD. Significant intersubject differences in the data included two subjects whose TR values obtained under both procedures were consistently lower (greater influence of ITD) than other subjects', and did not vary with ICI. Such differences suggest that multiple mechanisms of ITD/ILD combination may be utilized to varying degrees by individual listeners. By at least one of those mechanisms, ITD sensitivity (but not ILD sensitivity) is limited to low modulation rates.
doi:10.1016/j.heares.2010.06.002
PMCID: PMC2923247  PMID: 20547218
binaural hearing; time-intensity trading; onset dominance; rate limitation
24.  Neural and Behavioral Sensitivity to Interaural Time Differences Using Amplitude Modulated Tones with Mismatched Carrier Frequencies 
Bilateral cochlear implantation is intended to provide the advantages of binaural hearing, including sound localization and better speech recognition in noise. In most modern implants, temporal information is carried by the envelope of pulsatile stimulation, and thresholds to interaural time differences (ITDs) are generally high compared to those obtained in normal hearing observers. One factor thought to influence ITD sensitivity is the overlap of neural populations stimulated on each side. The present study investigated the effects of acoustically stimulating bilaterally mismatched neural populations in two related paradigms: rabbit neural recordings and human psychophysical testing. The neural coding of interaural envelope timing information was measured in recordings from neurons in the inferior colliculus of the unanesthetized rabbit. Binaural beat stimuli with a 1-Hz difference in modulation frequency were presented at the best modulation frequency and intensity as the carrier frequencies at each ear were varied. Some neurons encoded envelope ITDs with carrier frequency mismatches as great as several octaves. The synchronization strength was typically nonmonotonically related to intensity. Psychophysical data showed that human listeners could also make use of binaural envelope cues for carrier mismatches of up to 2–3 octaves. Thus, the physiological and psychophysical data were broadly consistent, and suggest that bilateral cochlear implants should provide information sufficient to detect envelope ITDs even in the face of bilateral mismatch in the neural populations responding to stimulation. However, the strongly nonmonotonic synchronization to envelope ITDs suggests that the limited dynamic range with electrical stimulation may be an important consideration for ITD encoding.
doi:10.1007/s10162-007-0088-5
PMCID: PMC2538436  PMID: 17657543
sound localization; binaural; inferior colliculus; psychophysics
25.  Behavioral Sensitivity to Broadband Binaural Localization Cues in the Ferret 
Although the ferret has become an important model species for studying both fundamental and clinical aspects of spatial hearing, previous behavioral work has focused on studies of sound localization and spatial release from masking in the free field. This makes it difficult to tease apart the role played by different spatial cues. In humans and other species, interaural time differences (ITDs) and interaural level differences (ILDs) play a critical role in sound localization in the azimuthal plane and also facilitate sound source separation in noisy environments. In this study, we used a range of broadband noise stimuli presented via customized earphones to measure ITD and ILD sensitivity in the ferret. Our behavioral data show that ferrets are extremely sensitive to changes in either binaural cue, with levels of performance approximating that found in humans. The measured thresholds were relatively stable despite extensive and prolonged (>16 weeks) testing on ITD and ILD tasks with broadband stimuli. For both cues, sensitivity was reduced at shorter durations. In addition, subtle effects of changing the stimulus envelope were observed on ITD, but not ILD, thresholds. Sensitivity to these cues also differed in other ways. Whereas ILD sensitivity was unaffected by changes in average binaural level or interaural correlation, the same manipulations produced much larger effects on ITD sensitivity, with thresholds declining when either of these parameters was reduced. The binaural sensitivity measured in this study can largely account for the ability of ferrets to localize broadband stimuli in the azimuthal plane. Our results are also broadly consistent with data from humans and confirm the ferret as an excellent experimental model for studying spatial hearing.
doi:10.1007/s10162-013-0390-3
PMCID: PMC3705081  PMID: 23615803
sound localization; spatial hearing; psychometric function; interaural time difference; interaural level difference; azimuth

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