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Effects of GABA-mediated inhibition on direction-dependent frequency tuning in the frog inferior colliculus.

Earlier studies from our laboratory have shown that the frequency selectivity of neurons in the frog inferior colliculus is direction dependent. The goal of this study was to test the hypotheses that gamma-aminobutyric acid or GABA (but not glycine)-mediated synaptic inhibition was responsible for the direction-dependence in frequency tuning, and that GABA acted through creation of binaural inhibition. We performed single unit recordings and investigated the unit's free-field frequency tuning, and/or the unit's response to the interaural level differences (under dichotic stimulation), before and during local applications of antagonists specific to gamma-aminobutyric acid A and glycine receptors. Our results showed that application of bicuculline produced a broadening of free-field frequency tuning depending on sound direction, i.e., more pronounced at azimuths at which the unit exhibited narrower frequency tuning under the pre-drug condition, thereby typically abolishing direction dependence in tuning. Application of strychnine produced no change in frequency tuning. The results from dichotic stimulation further revealed that bicuculline typically elevated and/or flattened the unit's interaural-level-difference response function, indicating a reduction in the strength of binaural inhibition. Our study provides evidence that gamma-aminobutyric acid-mediated binaural inhibition is important for direction dependence in frequency tuning.

Acoustic Stimulation↗

Neural inhibition sharpens auditory spatial selectivity of bat inferior collicular neurons.

This study examines the role of neural inhibition in auditory spatial selectivity of inferior collicular neurons of the big brown bat, Eptesicus fuscus, using a two-tone inhibition paradigm. Two-tone inhibition decreases auditory spatial response areas but increases the slopes of directional sensitivity curves of inferior collicular neurons. Inferior collicular neurons have either directionally-selective or hemifield directional sensitivity curves. A directionally-selective curve always has a peak which is at least 50% larger than the minimum. A hemifield directional sensitivity curve rises from an ipsilateral angle by more than 50% and either reaches a plateau or declines by less than 50% over a range of contralateral angles. Two-tone inhibition does not change directionally-selective curves but changes most hemifield directional sensitivity curves into directionally-selective curves. Auditory spatial selectivity determined both with and without two-tone inhibition increases with increasing best-excitatory frequency. Sharpening of auditory spatial selectivity by two-tone inhibition is larger for neurons with smaller differences between excitatory and inhibitory best frequencies. The effect of two-tone inhibition on auditory spatial selectivity increases with increasing inhibitory tone intensity but decreases with increasing intertone interval. The implications of these findings in bat echolocation are discussed.

Animals↗

Auditory behaviour of a parasitoid fly (Emblemasoma auditrix, Sarcophagidae, Diptera).

Females of the parasitoid fly Emblemasoma auditrix find their host cicada (Okanagana rimosa) by its acoustic signals. In laboratory experiments, fly phonotaxis had a mean threshold of about 66 dB SPL when tested with the cicada calling song. Flies exhibited a frequency dependent phonotaxis when testing to song models with different carrier frequencies (pulses of 6 ms duration and a repetition rate of 80 pulses s(-1)). However, the phonotactic threshold was rather broadly tuned in the range from 5 kHz to 11 kHz. Phonotaxis was also dependent on the temporal parameters of the song models: repetition rates of 60 pulses s(-1) and 80 pulses s and pulse durations of 5-7 ms resulted in the highest percentages of phonotaxis performing animals coupled with the lowest threshold values. Thus, parasitoid phonotaxis is adapted especially to the temporal parameters of the calling song of the host. Choice experiments revealed a preference of a song model with 9 kHz carrier frequency (peak energy of the host song) compared with 5 kHz carrier frequency (electrophysiologically determined best hearing frequency). However, this preference changed with the relative sound pressure level of both signals. When presented simultaneously, E. auditrix preferred 5-kHz signals, if they were 5 dB SPL louder than the 9-kHz signal.

Animal Communication↗

Early and late patterns of stimulus-related activity in auditory cortex of trained animals.

Epidural electrocorticograms over the right auditory cortex (field AI) were measured using implanted 18-channel (3 x 6) electrode arrays in four animals (Mongolian gerbil) trained to discriminate between a rising and a falling frequency modulated tone (frequency range 2-4 kHz). Using a previously introduced classification procedure, transient patterns of cortical activity suitable to discriminate between the rising and the falling modulation were identified. Early (locked to stimulus onset) and late (emerging at variable times poststimulus) patterns could be differentiated. Deletion of increasing numbers of randomly selected electrodes was used to determine a critical density of recording channels required to capture the discriminative power of the early and late patterns. Statistical analysis of the classification revealed a sigmoid dependence of the discriminative power from the number of remaining electrodes with an inflection point at 12 electrodes. The analysis of the minima of the classification statistic revealed that in the early patterns discriminative information was focal on regions corresponding to the tonotopic representation of the stimuli, whereas in late patterns this information seemed to be distributed nonfocally across larger cortical regions. This analysis supports the previous notion of the coexistence of topographically organized activity states related to the physical stimulus features and nontopographically organized states determined largely by intrinsic factors (Ohl et al. 2001).

Acoustic Stimulation↗

Oscillating neurons in the cochlear nucleus: I. Experimental basis of a simulation paradigm.

Anatomical and physiological auditory data and pitch measurements are presented including some additional analysis. The data provide the basis for a new computer model of sustained chopper neurons in the ventral cochlear nucleus. New and old evidence indicating a preference for multiples of 0.4 ms in oscillations of chopper neurons in the cochlear nucleus of different species such as man, cats, and Guinea fowls, is summarized. Our hypothesis is that the time constant of 0.4 ms is due to the minimum synaptic delay of chopper neuron connections. Anatomical findings show that chopper neurons are indeed connected and can excite each other; a model of a circular network of neurons that are connected via synapses with a delay of 0.4 ms is thus plausible. Results concerning frequency tuning and dynamical properties of periodicity encoding of chopper neurons are reviewed. It is concluded that chopper neurons receive input both from auditory nerve fibres and onset neurons.

Acoustic Stimulation↗

Processing of irrelevant location information under dual-task conditions.

This study deals with the problem of whether the processing of irrelevant location information in Simon-like tasks is triggered exogenously or endogenously. In Experiment 1, the primary task required one to press, as fast as possible, a left-hand-side key or a right-hand-side key (R1) to the pitch of a tone that was presented binaurally (S1). The secondary task required identifying, without time constraints, a visual stimulus (S2) that appeared randomly to the left or right of screen center. Results showed that there was a correspondence (i.e., a cross-task Simon effect) between the location of R1 and the location of S2 when S2 was presented alone. The cross-task Simon effect became much smaller (and insignificant) when a noise stimulus was presented contralateral to S2. Experiment 2 was similar to Experiment 1, except that S2 appeared unpredictably in only one-third of the trials. Results of Experiment 2 closely replicated those of Experiment 1: the cross-task Simon effect was much greater when S2 was presented alone. Experiment 3 differed from Experiment 1 because S2 had to be processed in only one-third of the trials, in which its identity was to be reported. In the remaining two-thirds of the trials, participants could ignore S2. Results confirmed that the cross-task Simon effect was much greater when S2 was presented alone. In contrast, it did not matter whether S2 had to be processed or not. In conclusion, the present study supports the hypothesis that the task-irrelevant spatial code of the stimulus is formed automatically, likely through an exogenously triggered selection. The role of endogenously initiated selection, if any, is much less important.

Adult↗

Task-order coordination in dual-task performance and the lateral prefrontal cortex: an event-related fMRI study.

A crucial demand in dual tasks suffering from a capacity limited processing mechanism is task-order scheduling, i.e. the control of the order in which the two component tasks are processed by this limited processing mechanism. The present study aims to test whether the lateral prefrontal cortex (LPFC) is associated with this demand. For this, 15 participants performed a psychological refractory paradigm (PRP) type dual task in an event-related functional magnetic resonance (fMRI) experiment. In detail, two choice reaction tasks, a visual (response with right hand) and an auditory (response with left hand), were presented with a temporal offset of 200 ms, while the participants were required to respond to the tasks in the order of their presentation. Importantly, the presentation order of the tasks changed randomly. Based on previous evidence, we argue that trials in which the present task order changed as compared to the previous trial (different-order trials) impose higher demands on task coordination than same-order trials do. The analyses showed that cortical areas along the posterior part of the left inferior frontal sulcus as well as the right posterior middle frontal gyrus were more strongly activated in different-order than in same-order trials, thus supporting the conclusion that one function of the LPFC for dual-task performance is the temporal coordination of two tasks. Furthermore, it is discussed that the present findings favour the active scheduling over the passive queuing hypothesis of dual-task processing.

Adult↗

Emergent perceptual features in the benefit of consistent stimulus-response mappings on dual-task performance.

Duncan (1979) examined all combinations of compatible and incompatible stimulus-response mappings for two spatial three-choice tasks in the psychological refractory period paradigm. Performance was better when the mappings for the tasks were consistent than when they were not, even when both mappings were incompatible. He attributed the benefit for the consistent incompatible mapping to an emergent choice between mappings when they are inconsistent that slows performance. Consistent incompatible mappings also may benefit from emergent perceptual features. The present study examined the role of emergent perceptual and mapping-choice features in two experiments that used pairs of two-choice tasks. Results similar to Duncan's were obtained with visual stimuli mapped to keypresses at short (stimulus onset asynchrony) SOAs. However, the benefit of the consistent incompatible mapping condition over the inconsistent mapping conditions was eliminated at an SOA of 1,000 ms. Furthermore, this benefit was not evident when the stimuli were auditory for Task 1 and visual for Task 2. With two-choice tasks, the benefit for consistent mappings apparently is due primarily to an emergent perceptual feature.

Attention↗

Exploring the mental number line: evidence from a dual-task paradigm.

In a parity-judgment task smaller numbers are responded to faster with the left-hand key and vice versa for larger numbers (SNARC effect; Dehaene et al., in Journal of Experimental Psychology: General, 122, 371-396, 1993). We used the psychological refractory period paradigm involving a parity-judgment task and tone-discrimination task to address the question at which stage this effect arises. When the parity-judgment task is performed second, then we found equal SNARC effects for the short and the long SOA. According to the central bottleneck model, this indicates that the effect arises during the response-selection or execution stage. In Experiment 2 the parity-judgment task was performed first. The pattern of results indicates that the SNARC effect originates during the perceptual encoding or response-selection. Together, our results suggest that the SNARC effect originates while the response is selected.

Adult↗

Proactive interference in a two-tone pitch-comparison task without additional interfering tones.

Two-tone pitch-comparison tasks typically comprise several successive pairs of successive tones separated by silent intervals. The serial occurrence of such pairs has been associated with degraded task performance, but the nature of this association is not fully understood. Human adult participants were presented with successive pairs of successive tones. The latter, to-be-compared tone of a pair could differ from the former, to-be-remembered tone of 1046.5 Hz by no more than +/-15 Hz (25 cents). The direction of this difference was easier to identify when it was opposite to that of the preceding pair than when being the same. Merely responding accordingly (irrespectively of whether the response was correct or not) was found not to account for this finding. Our study demonstrates proactive interference in a two-tone pitch comparison task as the difficulty to remember when the first tone of the present pair occurred relative to the last tone of the immediately preceding pair.

Acoustic Stimulation↗

Perceiving musical tension in long chord sequences.

We attempted to predict perceived musical tension in longer chord sequences by hierarchic and sequential models based on Lerdahl and Jackendoff's and Lerdahl's cognitive theories and on Parncutt's sensory-psychoacoustical theory. Musicians and nonmusicians were asked to rate the perceived tension of chords which were drawn either from a piece composed for the study (Exp. 1) or from a Chopin Prelude (Exps. 2-4). In Exps. 3 and 4, several experimental manipulations were made to emphasize either the global or the local structure of the piece and to verify how these manipultions would affect the respective contribution of the models in the ratings. In all experiments, musical tension was only weakly influenced by global harmonic structure. Instead, it mainly seemed to be determined locally, by harmonic cadences. The hierarchic model of Lerdahl and Jackendoff provided the best fit to tension ratings, not because it accounted for global hierarchic effects, but because it captured the local effect of cadences. By reacting to these local structures, tension ratings fit quite well with a hierarchic model, even though the participants were relatively insensitive to the global structure of the pieces. As a main outcome, it is argued that musical events were perceived through a short perceptual window sliding from cadence to cadence along a sequence.

Adult↗

Low-frequency tone pips elicit exaggerated startle reflexes in C57BL/6J mice with hearing loss.

The strength of the acoustic startle reflex (ASR) as a function of age was studied in adult C57BL/6J and CBA/CaJ mice, because altered ASR levels are a potential behavioral consequence of the neural reorganization that accompanies the early-onset hearing loss of the C57BL, in contrast to the normal-hearing CBA. For C57BL mice at 14-36 weeks of age, compared with 7-week-old mice, high-frequency thresholds measured with the auditory brainstem response (ABR) were less sensitive by about 25-30 dB while the hearing loss at low frequencies was 10-15 dB, but by 60 weeks losses of 45-50 dB were present across the entire spectrum. Their ASR amplitudes for 16 kHz tone pips were highest at 7 weeks and then declined with age, but, for 4 kHz tones the ASR increased in strength at 18 weeks and beyond to levels above that of the younger mice. This hyperreactivity persisted even in 60-week-old mice. The ASR for 16 kHz stimuli was positively correlated with hearing sensitivity, but the ASR for 4 kHz stimuli was positively correlated with hearing loss for mice that were 18-36 weeks of age. Furthermore, ASR amplitudes for 4 kHz stimuli were positively correlated with the 16 kHz ASR in young C57BL mice but negatively correlated in older mice. There were no similar ASR or ABR changes in adult CBA mice through 19 weeks of age. Correlations between ASR and ABR scores were always weakly positive, and correlations between 4 kHz and 16 kHz ASR amplitudes were always strongly positive. The ASR data in older C57BL mice with hearing loss are consistent with reports describing their increased neural representation of low-frequency sounds and reinforce the value of this strain for studying the functional consequences that accompany age-related cochlear degeneration.

Acoustic Stimulation↗

Estimates of human cochlear tuning at low levels using forward and simultaneous masking.

Auditory filter shapes were derived from psychophysical measurements in eight normal-hearing listeners using a variant of the notched-noise method for brief signals in forward and simultaneous masking. Signal frequencies of 1, 2, 4, 6, and 8 kHz were tested. The signal level was fixed at 10 dB above absolute threshold in the forward-masking conditions and fixed at either 10 or 35 dB above absolute threshold in the simultaneous-masking conditions. The results show that filter equivalent rectangular bandwidths (ERBs) are substantially narrower in forward masking than has been found in previous studies using simultaneous masking. Furthermore, in contrast to earlier studies, the sharpness of tuning doubles over the range of frequencies tested, giving Q(ERB) values of about 10 and 20 at signal frequencies of 1 and 8 kHz, respectively. It is argued that the new estimates of auditory filter bandwidth provide a more accurate estimate of human cochlear tuning at low levels than earlier estimates using simultaneous masking at higher levels, and that they are therefore more suitable for comparison to cochlear tuning data from other species. The data may also prove helpful in defining the parameters for nonlinear models of human cochlear processing.

Acoustic Stimulation↗

The role of spectral and temporal cues in voice gender discrimination by normal-hearing listeners and cochlear implant users.

The present study investigated the relative importance of temporal and spectral cues in voice gender discrimination and vowel recognition by normal-hearing subjects listening to an acoustic simulation of cochlear implant speech processing and by cochlear implant users. In the simulation, the number of speech processing channels ranged from 4 to 32, thereby varying the spectral resolution; the cutoff frequencies of the channels' envelope filters ranged from 20 to 320 Hz, thereby manipulating the available temporal cues. For normal-hearing subjects, results showed that both voice gender discrimination and vowel recognition scores improved as the number of spectral channels was increased. When only 4 spectral channels were available, voice gender discrimination significantly improved as the envelope filter cutoff frequency was increased from 20 to 320 Hz. For all spectral conditions, increasing the amount of temporal information had no significant effect on vowel recognition. Both voice gender discrimination and vowel recognition scores were highly variable among implant users. The performance of cochlear implant listeners was similar to that of normal-hearing subjects listening to comparable speech processing (4-8 spectral channels). The results suggest that both spectral and temporal cues contribute to voice gender discrimination and that temporal cues are especially important for cochlear implant users to identify the voice gender when there is reduced spectral resolution.

Adult↗

Concurrent sound segregation in electric and acoustic hearing.

We investigated potential cues to sound segregation by cochlear implant (CI) and normal-hearing (NH) listeners. In each presentation interval of experiment 1a, CI listeners heard a mixture of four pulse trains applied concurrently to separate electrodes, preceded by a "probe" applied to a single electrode. In one of these two intervals, which the subject had to identify, the probe electrode was the same as a "target" electrode in the mixture. The pulse train on the target electrode had a higher level than the others in the mixture. Additionally, it could be presented either with a 200-ms onset delay, at a lower rate, or with an asynchrony produced by delaying each pulse by about 5 ms re those on the nontarget electrodes. Neither the rate difference nor the asynchrony aided performance over and above the level difference alone, but the onset delay produced a modest improvement. Experiment 1b showed that two subjects could perform the task using the onset delay alone, with no level difference. Experiment 2 used a method similar to that of experiment 1, but investigated the onset cue using NH listeners. In one condition, the mixture consisted of harmonics 5 to 40 of a 100-Hz fundamental, with the onset of either harmonics 13 to 17 or 26 to 30 delayed re the rest. Performance was modest in this condition, but could be improved markedly by using stimuli containing a spectral gap between the target and nontarget harmonics. The results suggest that (a) CI users are unlikely to use temporal pitch differences between adjacent channels to separate concurrent sounds, and that (b) they can use onset differences between channels, but the usefulness of this cue will be compromised by the spread of excitation along the nerve-fiber array. This deleterious effect of spread-of-excitation can also impair the use of onset cues by NH listeners.

Acoustic Stimulation↗

Tonotopic projections of the auditory nerve to the cochlear nucleus angularis in the barn owl.

The nucleus angularis (NA), one of the two cochlear nuclei of birds, plays an important role in the processing of sound intensity. To begin investigating the NA in detail in the barn owl, which is a popular animal model for neural mechanisms of sound localization, a frequency map for this nucleus is presented here. Focal injections of horseradish peroxidase or neurobiotin were placed either in the NA or in the cochlear nucleus magnocellularis, labeling small groups of auditory nerve fibers of known characteristic frequency (CF) from 0.25 to 9.6 kHz. The courses of their axonal branches were used to construct a composite average map of the tonotopic frequency representation in the nucleus angularis. Nucleus angularis in the barn owl, as seen in frontal sections, resembles a sheet of cells bent approximately into an S shape. The lowest frequencies were found represented at the ventromedial extreme. The representation of increasingly higher frequencies then followed the S shape, with the highest frequencies located at the ventrolateral tip. Auditory nerve fibers of a given CF always entered the nucleus angularis within a well-restricted area and then traveled along their isofrequency band within the NA while branching off terminals. The isofrequency bands were typically slanted from caudo-ventro-medial to rostro-dorso-lateral. The basic tonotopic organization is comparable to that found in other birds, the major differences being the large size and unusual shape of the barn owl's nucleus angularis.

Animals↗

Music-to-language transfer effect: may melodic ability improve learning of tonal languages by native nontonal speakers?

In tonal languages, as Mandarin Chinese and Thai, word meaning is partially determined by lexical tones. Previous studies suggest that lexical tones are processed by native listeners as linguistic information and not as pure tonal information. This study aims at verifying if, in nontonal languages speakers, the discrimination of lexical Mandarin tones varies in function of the melodic ability. Forty-six students with no previous experience of Mandarin or any other tonal language were presented with two short lists of spoken monosyllabic Mandarin words and invited to perform a same-different task trying to identify whether the variation were phonological or tonal. Main results show that subjects perform significantly better in identifying phonological variations rather than tonal ones and interestingly, the group with a high melodic ability (assessed by Wing subtest 3) shows a better performance exclusively in detecting tonal variations.

Adult↗