Human auditory cortical mechanisms of sound lateralization: II. Interaural time differences at sound onset

Neuromagnetic responses were recorded over the right temporal cortex using a 24-channel gradiometer. Stimuli were binaural click trains, presented with six separate interaural time differences (ITDs). N100m to sound onset was larger and earlier for stimuli presented with left- than with right-leading ITDs. With stimulus lateralization taken into account, monaural and binaural stimuli evoked responses of roughly equal amplitude. In selective adaptation and oddball experiments, stimuli presented with different ITDs excited overlapping neuronal populations, but the amount of overlap decreased as the ITD between the stimuli increased. There were no systematic differences in the cortical source locations of the N100m as a function of ITD, however. Thus it appears that ITD-sensitive neurons in the human auditory cortex are not organized into a large-scale, orderly representation, which could be resolved by MEG.     

The measurement of the lateralization of narrow bands of noise using an acoustic pointing paradigm: the effect of sound-pressure level

The effects of varying interaural time delay (ITD) and interaural intensity difference (IID) were measured in normal-hearing subjects as a function of eleven frequencies and at sound-pressure levels (SPL) from 60 to 90 dB SPL and at 25-dB sensation level. Using an "acoustic" pointing paradigm, the IID of a 500-Hz narrow-band (100 Hz) noise (the "pointer") was varied by the subject to coincide with that of a "target" ITD stimulus. ITDs of 0, +/- 200, and +/- 400 microseconds were obtained through total waveform delays of narrow-band noise (NBN), including envelope and fine structure. The results of this experiment confirm the traditional view of binaural hearing for like stimuli: There is little perceived displacement away from 0 IID at frequencies of 1250 Hz and above. In the low frequencies, subjects required IIDs greater than the expected 10 dB to perceive a fully lateralized image, and they varied in the maximum value of IID that they required, regardless of frequency. Our subjects did not always perceive the intracranial locations of ITD targets symmetrically: When the signal was delayed to one ear, the resultant matching IID was often different in magnitude than for the same ITD target delayed to the opposite ear for the identical frequency. The results of two subjects suggested that people with asymmetric normal hearing have adapted to their asymmetry for lateralization tasks: The subjects were found to lateralize toward the ear with the greater SPL stimulus, regardless of the ear to which the signal was delayed, when signals of equal SL were presented, and toward the leading ear when signals of equal SPL were presented (unequal SL). Increasing the presentation levels above 60 dB SPL had an effect on the perception of high-frequency ITD targets: As the intensity level increased, the slopes of the IID versus ITD functions increased indicating better discrimination of ITD. This study is in agreement with other studies in providing strong evidence of individual differences in lateralization experiments. These individual differences might be attributable to differential sensitivity to ambiguous time stimulus cues, differential task sensitivity, age effects, threshold asymmetries, or criterion variability.     

Lateralization of bands of noise as a function of combinations of interaural intensive differences, interaural temporal differences, and bandwidth

Listeners indicated the intracranial position of bands of noise (from 50 to 400 Hz in width) for several combinations of interaural intensive differences (IID), and interaural temporal differences (ITD), and/or interaural phase differences (IPD). All ITD and IPD combinations produced an interaural delay of 1500 microseconds at the center frequency of the noise. The interaural phase spectra were constructed to produce several patterns of putative cross-correlation functions. Potency of IIDs depended greatly on particular combinations of bandwidth, ITD and IPD. For some combinations, changing the IID by only 3 dB resulted in large shifts in laterality (sometimes moving the image from near one ear to near the other). The complex interactions observed make the results incompatible with the traditional notion that IIDs simply act as weights or scalars. Rather, IIDs act in two distinct manners: (1) as independent scalar quantities and (2) by interacting with specific combinations of bandwidth and ITD/IPD, which is believed to reflect an action within the cross correlation surface.     

Pathophysiology of aquaporin-2 in water balance disorders

The binaural interaction component (BIC) of the 500-Hz human frequency-following response (FFR) was evaluated as a function of interaural intensity difference (IID) using a lateralization paradigm. The robust FFR interaction component (FFR-BIC) was shown to decrease systematically with increasing IID with no discernible FFR-BIC for IID values larger than about 20 dB. These findings are similar to that observed for the high-frequency auditory brainstem response interaction component (ABR-BIC). Thus, like the ABR-BIC, the FFR-BIC may be correlated with binaural fusion and the perceived location of the fused image of the sound. These results taken together suggest that the binaural neurons in the brainstem are able to utilize IID cues presented in both low-frequency and high-frequency sounds.     

Modeling of percutaneous drug transport in vitro using skin-imitating Carbosil membrane

Extracellular recordings were made with microelectrodes from single neurons in the rat's dorsal nucleus of the lateral lemniscus (DNLL) and response characteristics were determined for monaural and binaural acoustic stimulation. The vast majority of DNLL neurons were narrowly tuned to sound frequency and their temporal responses to contralateral tone pulses fell into one of three broad categories: onset (57%), sustained (21%) or onset-pause-sustained (22%). Most DNLL neurons fired multiple action potentials to a single click delivered to the contralateral ear. The majority (77%) of DNLL neurons showed a monotonic increase in the number of spikes elicited by contralateral tone pulses of increasing sound pressure level; the remaining cells were weakly non-monotonic. No obvious tonotopic pattern was found in the distribution of characteristic frequency of neurons in DNLL. Most DNLL neurons exhibited either excitatory/inhibitory (74%) or excitatory/excitatory (9%) binaural response patterns. The remaining cells (17%) were monaural and driven exclusively by stimulation of the contralateral ear. The binaural neurons in DNLL were sensitive to both interaural intensity and interaural time differences as determined by presentation of dichotic tone bursts and clicks respectively. The responses of DNLL neurons could be distinguished on the basis of monaural and binaural response characteristics from those in surrounding areas including the sagulum, paralemniscal zone and the intermediate nucleus of the lateral lemniscus.     

Interaural delay-dependent changes in the binaural difference potential in cat auditory brainstem response: implications about the origin of the binaural interaction component

Auditory brainstem responses (ABRs) evoked by dichotic clicks with 12 different interaural delays (ITDs) between 0 and 1500 microsecond(s) were recorded from the vertices of 10 cats under ketamine anesthesia. The so-called binaural difference potential (BDP), considered to be an indicator of binaural interaction (BI), was computed by subtracting the sum of the two monaural responses from the binaural one. The earliest and most prominent component of BDP was a negative deflection (DN1) at a latency between 4 and 4.8 ms. Like all the other components of BDP, DNI was also due to binaural reduction rather than enhancement of the corresponding ABR wave, P4 in this case. Furthermore, the way its latency increased as a function of ITD was also not compatible with what would be predicted by the delay-line coincidence detector models based on the excitatory-excitatory units in the medial superior olive (MSO). We therefore proposed an alternative hypothesis for the origin of this BI component based on the inhibitory-excitatory (IE) units in the lateral superior olive (LSO). The computational model designed closely simulated the ITD-dependent attenuation and latency shifts observed in DN1. It was therefore concluded that the origin of this BI component in the cat's vertex-ABR could be the lateral lemniscal output of the LSO, although the delay lines which have been shown to exist also in the mammalian brain may play an important role in encoding ITDs.     

Effects of ear plugging on single-unit azimuth sensitivity in cat primary auditory cortex. I. Evidence for monaural directional cues

1. Single-unit recordings were carried out in primary auditory cortex (AI) of barbiturate-anesthetized cats. Neurons, sensitive to sound direction in the horizontal plane (azimuth), were identified by their responses to noise bursts, presented in the free field, that varied in azimuth and sound pressure level (SPL). SPLs typically varied between 0 and 80 dB and were presented at each azimuth that was tested. Each azimuth-sensitive neuron responded well to some SPLs at certain azimuths and did not respond well to any SPL at other azimuths. This report describes AI neurons that were sensitive to the azimuth of monaurally presented noise bursts. 2. Unilateral ear plugging was used to test each azimuth-sensitive neuron's response to monaural stimulation. Ear plugs, produced by injecting a plastic ear mold compound into the concha and ear canal, attenuated sound reaching the tympanic membrane by 25-70 dB. Binaural interactions were inferred by comparing responses obtained under binaural (no plug) and monaural (ear plug) conditions. 3. Of the total sample of 131 azimuth-sensitive cells whose responses to ear plugging were studied, 27 were sensitive to the azimuth of monaurally presented noise bursts. We refer to these as monaural directional (MD) cells, and this report describes their properties. The remainder of the sample consisted of cells that either required binaural stimulation for azimuth sensitivity (63/131), because they were insensitive to azimuth under unilateral ear plug conditions or responded too unreliably to permit detailed conclusions regarding the effect of ear plugging (41/131). 4. Most (25/27) MD cells received either monaural input (MD-E0) or binaural excitatory/inhibitory input (MD-EI), as inferred from ear plugging. Two MD cells showed other characteristics. The contralateral ear was excitatory for 25/27 MD cells. 5. MD-E0 cells (22%, 6/27) were monaural. They were unaffected by unilateral ear plugging, showing that they received excitatory input from one ear, and that stimulation of the other ear was without apparent effect. On the other hand, some monaural cells in AI were insensitive to the azimuth of noise bursts, showing that sensitivity to monaural directional cues is not a property of all monaural cells in AI. 6. MD-EI cells (70%, 19/27) exhibited an increase in responsiveness on the side of the plugged ear, showing that they received excitatory drive from one ear and inhibitory drive from the other. MD-EI cells remained azimuth sensitive with the inhibitory ear plugged, showing that they were sensitive to monaural directional cues at the excitatory ear.(ABSTRACT TRUNCATED AT 400 WORDS)     

Neuronal coding of interaural transient envelope disparities

Onsets are salient and important transient (i.e. dynamic) features of acoustic signals, and evoke vigorous responses from most auditory neurons, but paradoxically these onset responses have most often been analysed with respect to steady-state stimulus features, e.g. the sound pressure level (SPL). In nearly all studies concerned with the coding of differences in SPL at the two ears (interaural level differences; ILDs), which provide a major cue for the azimuthal location of high frequency sound sources, interaural onset disparities were covaried with ILD, but the possibly confounding effects of this covariation on neuronal responses have been entirely neglected. Therefore, dichotic stimulus paradigms were designed here in which onset and steady-state features were varied independently. Responses were recorded from single neurons in the inferior colliculus of rats, anaesthetized with pentobarbital and xylazine. It is demonstrated that onset responses, or the onset response components of neurons with more complex temporal response patterns, are dependent on the binaural combination of dynamic envelope features associated with conventional ILD stimulus paradigms, but not on the binaural combination of steady-state SPLs reached after the onset. In contrast, late or sustained response components appear more sensitive to the binaural combination of steady-state SPLs. These data stress the general necessity for a separate analysis of onset and late response components, with respect to different stimulus features, and suggest a need for re-evaluation of existing studies on ILD coding. The sensitivity of onset responses to the binaural combination of envelope transients, rather than to steady-state ILD, is in line with their sensitivity to other interaural envelope disparities, created by stationary or moving sounds.     

Effects of ear of entry and perceived location of synchronous and asynchronous components on mistuning detection

Listeners were required to detect mistuning imposed on the center ("target") component of a 200-ms complex consisting of the first seven harmonics of a 500-Hz fundamental. In the standard interval of each 2IFC trial, all components were frequency modulated in-phase by a 5-Hz sinusoid. In the signal interval the frequency modulation of the target component was inverted in-phase, thereby introducing a mistuning proportional to the depth of FM. In a similar experiment, using monaural presentation, Carlyon [J. Acoust. Soc. Am. 95, 2622-2630 (1994)] reported a substantial elevation of thresholds in the presence of an unmodulated asynchronous interferer with frequency identical to the mean frequency of the target. This was attributed to the interferer, causing the target component to be perceptually segregated from the remainder of the complex, thereby impairing across-frequency comparisons. Experiment 1 of the present study showed that an interferer presented contralaterally for 200 ms before and 100 ms after the signal complex (no simultaneous presentation) also impaired performance, but to a lesser extent than an ipsilaterally presented one. Experiment 2 showed that an interferer which was presented dichotically with an interaural level difference (ILD) of 10 dB, so that it was perceived contralaterally, had the same (large) effect as if it were presented ipsilaterally. Experiment 3 showed that, in the absence of any interferer, performance was impaired when the nontarget components were presented contralaterally to the target component. However, performance was not impaired when the nontarget components were presented dichotically with an ILD of 20 dB, so that they were perceived contralaterally to the target component. It is concluded that the level of performance in the mistuning task is determined by whether the target is presented to the same ear as the rest of the complex, rather than by its perceived location.     

Lateralized auditory spatial perception and the contralaterality of cortical processing as studied with functional magnetic resonance imaging and magnetoencephalography

Functional magnetic resonance imaging (fMRI) and magnetoencephalography (MEG) were used to study the relationships between lateralized auditory perception in humans and the contralaterality of processing in auditory cortex. Subjects listened to rapidly presented streams of short FM-sweep tone bursts to detect infrequent, slightly deviant tone bursts. The stimulus streams consisted of either monaural stimuli to one ear or the other or binaural stimuli with brief interaural onset delays. The onset delay gives the binaural sounds a lateralized auditory perception and is thought to be a key component of how our brains localize sounds in space. For the monaural stimuli, fMRI revealed a clear contralaterality in auditory cortex, with a contralaterality index (contralateral activity divided by the sum of contralateral and ipsilateral activity) of 67%. In contrast, the fMRI activations from the laterally perceived binaural stimuli indicated little or no contralaterality (index of 51%). The MEG recordings from the same subjects performing the same task converged qualitatively with the fMRI data, confirming a clear monaural contralaterality, with no contralaterality for the laterally perceived binaurals. However, the MEG monaural contralaterality (55%) was less than the fMRI and decreased across the several hundred millisecond poststimulus time period, going from 57% in the M50 latency range (20-70 ms) to 53% in the M200 range (170-250 ms). These data sets provide both quantification of the degree of contralaterality in the auditory pathways and insight into the locus and mechanism of the lateralized perception of spatially lateralized sounds.     

Binaural inhibition is important in shaping the free-field frequency selectivity of single neurons in the inferior colliculus

1. We have shown previously that under free-field stimulation in the frontal field, frequency selectivity of the majority of inferior colliculus (IC) neurons became sharper when the loudspeaker was shifted to ipsilateral azimuths. These results indicated that binaural inhibition may be responsible for the direction-dependent sharpening of frequency selectivity. To test the above hypothesis directly, we investigated the frequency selectivity of IC neurons under several conditions: monaural stimulation using a semiclosed acoustical stimulation system, binaural stimulation dichotically also using a semiclosed system, free-field stimulation from different azimuths, and free-field stimulation when the ipsilateral ear was occluded monaurally (coated with a thick layer of petroleum jelly, which effectively attenuated acoustic input to this ear). 2. The binaural interaction pattern of 98 IC neurons of northern leopard frogs (Rana pipiens pipiens) were evaluated; of these neurons, there were 34 EE and 64 EO neurons. The majority of IC neurons (92 of 98) showed some degree of binaural inhibition (i.e., showing diminished response when the ipsilateral and contralateral ears were stimulated simultaneously) whether they were designated as EE or EO; these IC neurons thus were classified as EE-I or EO-I. Neurons were classified as exhibiting strong inhibition if the ILD function showed a pronounced response decrement, i.e., a decrease of > or = 50% of the response to monaural stimulation of the contralateral ear. Those neurons that showed smaller response decrements (decrease was > or = 25% but < 50%) were designated as showing weak inhibition. Most of these EE-I and EO-I neurons (n = 68) showed strong binaural inhibition. 3. In agreement with results from our earlier studies, frequency threshold curves (FTCs) of IC neurons were altered by sound azimuth. Independent of binaural interaction pattern, most IC neurons (59 of 98) showed a narrowing of the FTC as sound direction was changed from contralateral 90 deg (c90 degrees) to ipsilateral 90 deg (i90 degrees). IC neurons that exhibited the largest direction-dependent changes in frequency selectivity were typically those that displayed stronger binaural inhibition. Occlusion of the ipsilateral ear, which reduced the strength of binaural inhibition by this ear, abolished direction-dependent frequency selectivity. 4. FTCs of IC neurons that exhibited little to moderate direction-dependent effects on frequency selectivity were associated typically with neurons that displayed weak binaural inhibition. Associated with this weak binaural inhibition, central neural responses under monaural occlusion also displayed only small effects; the FTCs were only slightly broader than those derived in the intact condition, and as before, the experimental manipulation resulted in abolishment of direction-dependent frequency selectivity. 5. In contrast to most IC neurons, which showed direction-dependent narrowing of the FTC, about one-third (34 of 98) of IC neurons studied showed a broadening of the FTC when sound direction was shifted to ipsilateral azimuths. Interestingly, for 90% of these 34 neurons, monaural occlusion resulted in narrowing of the bandwidth at each azimuth instead of broadening of the FTC bandwidth. We have evidence to suggest that this direction-dependent broadening is actually a consequence of a truncation or loss of the tip of the FTC derived at c90 degrees, which results from strong binaural inhibition. 6. To compare the frequency threshold tuning in response to monaural stimulation of each ear with free-field FTCs, we measured FTCs for each of the 34 EE neurons to independent contralateral and ipsilateral stimulation. FTCs derived from ipsilateral monaural stimulation were significantly narrower than those resulting from contralateral monaural stimulation, independent of a neuron's direction-dependent changes in frequency selectivity.     

Neural responses to simple simulated echoes in the auditory brain stem of the unanesthetized rabbit

1. In most natural environments, sound waves from a single source will reach a listener through both direct and reflected paths. Sound traveling the direct path arrives first, and determines the perceived location of the source despite the presence of reflections from many different locations. This phenomenon is called the "law of the first wavefront" or "precedence effect." The time at which the reflection is first perceived as a separately localizable sound defines the end of the precedence window and is called "echo threshold." The precedence effect represents an important property of the auditory system, the neural basis for which has only recently begun to be examined. Here we report the responses of single neurons in the inferior colliculus (IC) and superior olivary complex (SOC) of the unanesthetized rabbit to a sound and its simulated reflection. 2. Stimuli were pairs of monaural or binaural clicks delivered through earphones. The leading click, or conditioner, simulated a direct sound, and the lagging click, or probe, simulated a reflection. Interaural time differences (ITDs) were introduced in the binaural conditioners and probes to adjust their simulated locations. The probe was always set at the neuron's best ITD, whereas the conditioner was set at the neuron's best ITD or its worst ITD. To measure the time course of the effects of the conditioner on the probe, we examined the response to the probe as a function of the conditioner-probe interval (CPI). 3. When IC neurons were tested with conditioners and probes set at the neuron's best ITD, the response to the probe as a function of CPI had one of two forms: early-low or early-high. In early-low neurons the response to the probe was initially suppressed but recovered monotonically at longer CPIs. Early-high neurons showed a nonmonotonic recovery pattern. In these neurons the maximal suppression did not occur at the shortest CPIs, but rather after a period of less suppression. Beyond this point, recovery was similar to that of early-low neurons. The presence of early-high neurons meant that the overall population was never entirely suppressed, even at short CPIs. Taken as a whole. CPIs for 50% recovery of the response to the probe among neurons ranged from 1 to 64 ms with a median of approximately 6 ms. 4. The above results are consistent with the time course of the precedence effect for the following reasons. 1) The lack of complete suppression at any CPI is compatible with behavioral results that show the presence of a probe can be detected even at short CPIs when it is not separately localizable. 2) At a CPI corresponding to echo threshold for human listeners (approximately 4 ms CPI) there was a considerable response to the probe, consistent with it being heard as a separately localizable sound at this CPI. 3) Full recovery for all neurons required a period much longer than that associated with the precedence effect. This is consistent with the relatively long time required for conditioners and probes to be heard with equal loudness. 5. Conditioners with either the best ITD or worst ITD were used to determine the effect of ITD on the response to the probe. The relative amounts of suppression caused by the two ITDs varied among neurons. Some neurons were suppressed about equally by both types of conditioners, others were suppressed more by a conditioner with the best ITD, and still others by a conditioner with the worst ITD. Because the best ITD and worst ITD presumably activate different pathways, these results suggest that different neurons receive a different balance of inhibition from different sources. 6. The recovery functions of neurons not sensitive to ITDs were similar to those of ITD-sensitive, neurons. This suggests that the time course of suppression may be common among different IC populations. 7. We also studied neurons in the SOC. Although many showed binaural interactions, none were sensitive to ITDs. Thus the response of this population may not be     

Single olivocochlear neurons in the guinea pig. I. Binaural facilitation of responses to high-level noise

Single medial olivocochlear (MOC) neurons were recorded from the cochlea of the anesthetized guinea pig. We used tones and noise presented monaurally and binaurally and measured responses for sounds up to 105 dB sound pressure level (SPL). For monaural sound, MOC neuron firing rates were usually higher for noise bursts than tone bursts, a situation not observed for afferent fibers of the auditory nerve that were sampled in the same preparations. MOC neurons also differed from afferent fibers in having less saturation of response. Some MOC neurons had responses that continued to increase even at high sound levels. Differences between MOC and afferent responses suggest that there is convergence in the pathway to olivocochlear neurons, possibly a combination of inputs that are at the characteristic frequency (CF) with others that are off the CF. Opposite-ear noise almost always facilitated the responses of MOC neurons to sounds in the main ear, the ear that best drives the unit. This binaural facilitation depends on several characteristics that pertain to the main ear: it is higher in neurons having a contralateral main ear (contra units), it is higher at main-ear sound levels that are moderate (approximately 65 dB SPL), and it is higher in neurons with low discharge rates to main-ear stimuli. Facilitation also depends on parameters of the opposite-ear sound: facilitation increases with noise level in the opposite ear until saturating, is greater for continuous noise than noise bursts, and is usually greater for noise than for tones. Using optimal opposite-ear facilitators and high-level stimuli, the firing rates of olivocochlear neurons range up to 140 spikes/s, whereas for moderate-level monaural stimuli the rates are <80 spikes/s. At high sound levels, firing rates of olivocochlear neurons increase with CF, an increase that may compensate for the known lower effectiveness of olivocochlear synapses on outer hair cells responding to high frequencies. Overall, our results demonstrate a high MOC response for binaural noise and suggest a prominent role for the MOC system in environments containing binaural noise of high level.     

Single olivocochlear neurons in the guinea pig. II. Response plasticity due to noise conditioning

Previous studies have shown that daily, moderate-level sound exposure, or conditioning, can reduce injury from a subsequent high-level noise exposure. We tested the hypothesis that this conditioning produces an increased activity in the olivocochlear efferent reflex, a reflex known to provide protection to the cochlea. Guinea pigs were conditioned by a 10-day intermittent exposure to 2-4 kHz noise at 85 dB sound pressure level. This conditioning is known to reduce damage from a subsequent high-level exposure to the same noise band. Responses to monaural and binaural sound were recorded from single medial olivocochlear (MOC) efferent neurons, and data from conditioned animals were compared with those obtained from unexposed controls. MOC neurons were classified by their response to noise bursts in the ipsilateral or contralateral ears as ipsi units, contra units, or either-ear units. There were no significant differences in the distributions of these unit types between control and conditioned animals. There were also no differences in other responses to monaural stimuli, including the distribution of characteristic frequencies (CFs), the sharpness of tuning, or thresholds at the CF. For binaural sound at high levels, particularly relevant to sound-evoked activation of the MOC reflex during acoustic overstimulation, the firing rates of MOC neurons with CFs just above the conditioning band showed slight (but statistically significant) elevations relative to control animals. Frequency regions just above the conditioning band also demonstrated maximum conditioning-related protection; thus protection could be due, in part, to long-term changes in MOC discharge rates. For binaural sound at low levels, MOC firing rates in conditioned animals also were increased significantly relative to controls. Again, increases were largest for neurons with CFs just above the conditioning band. For equivalent monaural sound, rates were not significantly increased; thus, conditioning appears to increase binaural facilitation by opposite-ear sound. These data indicate that MOC neurons show long-term plasticity in acoustic responsiveness that is dependent on their acoustic history.     

Tracking of complex binaural images

Subjects were presented with an initially stationary binaural image formed by the fusion of two identical pulses, without interaural time difference (ITD), at the two ears. The image was then made to traverse the subject's auditory perceptual space by introducing ITD, varying linearly with time, under computer control. The direction of movement, i.e. towards the right or left ear, could be reversed by the listener, by pressing a button. Subjects were requested to keep the image central, by pressing the button when they judged that deviation from subjective centre had occurred. Experiments of this type can be considered as analogous to Békésy audiometry, where the subject automatically traces his threshold of hearing, in that here the listener traces out his auditory perceptual centre as it varies with time. Hence, equivalent analyses to those employed for Békésy audiometry are possible. Subsequent to the initial part of the experiment, an additional pulse was added to one channel, preceding the original pulse, to form a pulse pair. The monaural masking of the original pulse by this additional pulse thus acts to shift the pre-existing binaural image. The effect of varying the amplitude and onset time of the masking pulse, relative to the original pulse, on the Békésy-type trace was examined.     

Auditory spatial responses of young guinea pigs (Cavia porcellus) during and after ear blocking.

Tested 30 newborn guinea pigs to determine their ability to approach an auditory stimulus early in development. Observations of the behavior of 1–4 day old Ss in a circular 8-choice maze revealed a pronounced tendency to orient toward and approach a tape-recorded signal of guinea pig vocalizations. The occurrence of approach responses was reduced to chance in Ss tested with one ear occluded by wax ear plugs which attenuated but did not totally eliminate sound. The effect of monaural ear blocks was more severe than binaural blocks, which reflects the importance of binaural cues in the maintenance of approach responses to sound. In a 2nd study with 40 Ss the ability of older animals, 11–31 days of age, was examined. Directional approach responses to sound were also evident at this age, and ear plugs disrupted performance only under monaural conditions. Furthermore, in Ss raised from birth with monaural ear blocks but tested without ear plugs, there was a subsequent disruption of performance for at least 21 days. Results indicate the importance of binaural cues in the development of early auditory spatial reponses and suggest the need for appropriate binaural experience for subsequent localization of sounds. (52 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Some aspects of the lateralization of echoed sound in man. I. The classical interaural-delay based precedence effect

The precedence effect in two-click stimuli was investigated by measuring observers' sensitivity to interaural time delays (ITDs) as a function of interclick interval (ICI). A two-interval two-alternative forced-choice discrimination paradigm was used in two stimulus configurations: type I, a dichotic click with a given ITD preceded a diotic click; and type II, a dichotic click followed a diotic click. Threshold ITDs were measured in each configuration for a finely sampled distribution of ICIs that ranged from 0.1 to 25.6 ms. Performance was characterized by the "threshold elevation factor" (TEF) which normalized each of the observers' type I and type II ITD thresholds relative to their ITD threshold for a single dichotic click. The finer sampling of ICIs revealed two novel results: First, for two observers, sensitivity to ITD in the later arriving ITD (type II) oscillated in a consistent and systematic way with changes in ICI. Second, when the ICI reached 12.8 ms, ITD thresholds in the type I and type II configurations were equal but nearly a factor of 2 greater than for a single dichotic click. Some aspects of the data are consistent with the phenomenon of binaural adaptation.     

Psychophysical and speech perception studies: a case report on a binaural cochlear implant subject

Further improvements in speech perception for cochlear implant patients in quiet and in noise should be possible with speech processing strategies using binaural implants. For this reason, presented here is a series of initial psychophysical and speech perception studies on the authors' first binaural cochlear implant patient. For an approximate matching of the places of stimulation on the two sides, the patient usually reported a single percept when the two sides were simultaneously stimulated. Lateralization was strongly influenced by amplitude differences between the electrical stimuli on the two sides, but only weakly by interaural time delays. Speech testing, comparing monaural with binaural electrical stimulation, showed a binaural advantage particularly in noise.     

Early monaural occlusion alters the neural map of interaural level differences in the inferior colliculus of the barn owl

Monaural occlusion during early life causes adaptive changes in the tuning of units in the owl's optic tectum to interaural level differences (ILD) that tend to align the auditory with the visual map of space. We investigated whether these changes could be due to experience-dependent plasticity occurring in the auditory pathway prior to the optic tectum. Units were recorded in the external nucleus of the inferior colliculus (ICx), which is a major source of auditory input to the optic tectum. The tuning of ICx units to ILD was measured in normal barn owls and in barn owls raised with one ear occluded. ILD tuning at each recording site was measured with dichotic noise bursts, presented at a constant average binaural level, 20 dB above threshold. The best ILD at each site was defined as the midpoint of the range of ILD values which elicited more than 50% of the maximum response. A physiological map of ILD was found in the ICx of normal owls: best ILDs changed systematically from right-ear-greater to left-ear-greater as the electrode progressed from dorsal to ventral. Best ILDs ranged from 13 dB right-ear-greater to 15 dB left-ear-greater and progressed at an average rate of 12 dB/mm. The representations of ILD were similar on both sides of the brain. In the ICx of owls raised with one ear occluded, the map of ILD was shifted in the adaptive direction: ILD tuning was shifted towards values favoring the non-occluded ear (the direction that would restore a normal space map). The average magnitude of the shift was on the order of 8-10 dB in each of 4 owls. In one owl, the mean shift in ILD tuning was almost identical on both sides of the brain. In another owl, the mean shift was much larger on the side ipsilateral to the occlusion than on the contralateral side. In both cases, the mean shifts measured in each ICx were comparable to the mean shifts measured in the optic tectum on the same sides of the brain. Thus, the adjustments in ILD tuning that have been observed in the optic tectum in response to monaural occlusion are almost entirely due to adaptive mechanisms that operate at or before the level of the ICx.     

Development of the auditory orientation response in the albino rat (Rattus norvegicus).

The development of head orientation to auditory stimulation was examined in rat pups at Postnatal Days 8, 11, 14, 17, and 20. The animals were tested in a quiet environment with single bursts of 65 dB (SPL) broad-band noise. A reflexive head turn toward the sound was first seen on Postnatal Day 14 and subsequently on Days 17 and 20. This result demonstrates that the onset of directional auditory responses occurred between Day 11 and Day 14. The role of binaural cues in early sound orientation was examined in 17-day-old pups with monaural ligation of the external meatus. These animals were unable to localize a sound source and consistently turned toward the side of the unligated ear regardless of the position of the stimulus. Thus binaural cues were shown to be important for head orientation to sound in early development. In a separate study, head orientation to high and low frequency tone pips was examined. Directional responses were first seen on Day 12 for a 16-kHz tone and Day 14 for a 2-kHz tone. These results indicate an earlier onset for orientation to high frequency sounds in the rat. (PsycINFO Database Record (c) 2010 APA, all rights reserved)