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Processing of novel sounds and frequency changes in the human auditory cortex: magnetoencephalographic recordings.

Whole-head magnetoencephalographic (MEG) responses to repeating standard tones and to infrequent slightly higher deviant tones and complex novel sounds were recorded together with event-related brain potentials (ERPs). Deviant tones and novel sounds elicited the mismatch negativity (MMN) component of the ERP and its MEG counterpart (MMNm) both when the auditory stimuli were attended to and when they were ignored. MMNm generators were located bilateral to the superior planes of the temporal lobes where preattentive auditory discrimination appears to occur. A subsequent positive P3a component was elicited by deviant tones and with a larger amplitude by novel sounds even when the sounds were to be ignored. Source localization for the MEG counterpart of P3a (P3am) suggested that the auditory cortex in the superior temporal plane is involved in the neural network of involuntary attention switching to changes in the acoustic environment.

Acoustic Stimulation↗

Spectral response patterns of auditory cortex neurons to harmonic complex tones in alert monkey (Macaca mulatta).

1. The auditory cortex in the superior temporal region of the alert rhesus monkey was explored for neuronal responses to pure and harmonic complex tones and noise. The monkeys had been previously trained to recognize the similarity between harmonic complex tones with and without fundamentals. Because this suggested that they could preceive the pitch of the lacking fundamental similarly to humans, we searched for neuronal responses relevant to this perception. 2. Combination-sensitive neurons that might explain pitch perception were not found in the surveyed cortical regions. Such neurons would exhibit similar responses to stimuli with similar periodicities but differing spectral compositions. The fact that no neuron with responses to a fundamental frequency responded also to a corresponding harmonic complex missing the fundamental indicates that cochlear distortion products at the fundamental may not have been responsible for missing fundamental-pitch perception in these monkeys. 3. Neuronal responses can be expressed as relatively simple filter functions. Neurons with excitatory response areas (tuning curves) displayed various inhibitory sidebands at lower and/or higher frequencies. Thus responses varied along a continuum of combined excitatory and inhibitory filter functions. 4. Five elementary response classes along this continuum are presented to illustrate the range of response patterns. 5. "Filter (F) neurons" had little or no inhibitory sidebands and responded well when any component of a complex tone entered its pure-tone receptive field. Bandwidths increased with intensity. Filter functions of these neurons were thus similar to cochlear nerve-fiber tuning curves. 6. "High-resolution filter (HRF) neurons" displayed narrow tuning curves with narrowband widths that displayed little growth with intensity. Such cells were able to resolve up to the lowest seven components of harmonic complex tones as distinct responses. They also responded well to wideband stimuli. 7. "Fundamental (F0) neurons" displayed similar tuning bandwidths for pure tones and corresponding fundamentals of harmonic complexes. This response pattern was due to lower harmonic complexes. This response pattern was due to lower inhibitory sidebands. Thus these cells cannot respond to missing fundamentals of harmonic complexes. Only physically present components in the pure-tone receptive field would excite such neurons. 8. Cells with no or very weak responses to pure tones or other narrowband stimuli responded well to harmonic complexes or wideband noise.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗

Temporally nonadjacent nonlinguistic sounds affect speech categorization.

Speech perception is an ecologically important example of the highly context-dependent nature of perception; adjacent speech, and even nonspeech, sounds influence how listeners categorize speech. Some theories emphasize linguistic or articulation-based processes in speech-elicited context effects and peripheral (cochlear) auditory perceptual interactions in non-speech-elicited context effects. The present studies challenge this division. Results of three experiments indicate that acoustic histories composed of sine-wave tones drawn from spectral distributions with different mean frequencies robustly affect speech categorization. These context effects were observed even when the acoustic context temporally adjacent to the speech stimulus was held constant and when more than a second of silence or multiple intervening sounds separated the nonlinguistic acoustic context and speech targets. These experiments indicate that speech categorization is sensitive to statistical distributions of spectral information, even if the distributions are composed of nonlinguistic elements. Acoustic context need be neither linguistic nor local to influence speech perception.

Adult↗

Functional organization of the cochlear nucleus of rufous horseshoe bats (Rhinolophus rouxi): frequencies and internal connections are arranged in slabs.

The functional organization of the cochlear nucleus (CN) was studied with physiological recording and anatomical tracing techniques. Recordings were made from single CN neurons to examine their temporal firing patterns to tone burst stimuli and their frequency tuning characteristics. Recording loci of individual neurons were carefully monitored in order to understand how the functional properties of a cell relate to its location within the CN. We found that tonal frequencies were systematically represented in each of the three CN divisions (anteroventral, AVCN; posteroventral, PVCN; dorsal, DCN). Eight temporal response patterns were observed in CN neurons when stimulated at units' best excitatory frequencies (BF). With a few exceptions, neurons in each CN division could generate all eight firing patterns with different distributions for the three division. A focal injection of horseradish peroxidase (HRP), at the end of the physiological study, to a group of neurons possessing a similar BF in one CN division resulted in anterograde labeling of nerve terminals in the other two divisions at precisely the areas where the same frequency band was processed in these divisions. Labeled terminals in each division were closely congregated in the form of a thin slab. The slab orientation was division specific whereas its location was frequency specific, which could be predicted on the basis of physiological data. HRP injections into the DCN also resulted in retrograde labeling of somata in the AVCN and PVCN. On the other hand, only DCN neurons were retrogradely labeled when HRP was injected into the AVCN or the PVCN. These data showed how the three CN divisions are internally connected. Furthermore, retrogradely labeled cells occupied the same slabs where we found anterogradely labeled nerve terminals. Additionally, in a group of bats, HRP was injected into various functionally (i.e., BF) identified regions of the central nucleus of the inferior coliculus (IC) to clarify the type and location of CN projecting neurons. Retrogradely labeled cells in individual CN divisions likewise were arranged in slabs whose locations in the CN nuclei depended on the BFs of neurons at the injection site in the IC. These results show that slabs represent units of functional organization (i.e., tonal frequency, local connection and central projection) in the CN.

Animals↗

Iso-frequency 2-DG contours in the inferior colliculus of the awake monkey.

Tone bursts produced bands of selective 2-[14C]-deoxyglucose labelling in the inferior colliculus (IC) of the awake monkey. Low tone frequencies produced labelling in dorsal regions and high tone frequencies produced labelling in ventral regions. The position of the bands coincided with the position of a single unit with a characteristic frequency, which was the same as the frequency producing the labelling. These findings indicate that the bands of labelling represent iso-frequency contours in IC. The iso-frequency contours extended across most of the nucleus and were oriented from dorsomedially to ventro-laterally at 20-30 degrees from the horizontal and became more vertical anteriorly. The width of the contours was as narrow as 200 micron, suggesting that the contours might represent 2 or 3 overlapping cellular laminae.

Animals↗

Perceptual scaling of synthesized musical timbres: common dimensions, specificities, and latent subject classes.

To study the perceptual structure of musical timbre and the effects of musical training, timbral dissimilarities of synthesized instrument sounds were rated by professional musicians, amateur musicians, and nonmusicians. The data were analyzed with an extended version of the multidimensional scaling algorithm CLASCAL (Winsberg & De Soete, 1993), which estimates the number of latent classes of subjects, the coordinates of each timbre on common Euclidean dimensions, a specificity value of unique attributes for each timbre, and a separate weight for each latent class on each of the common dimensions and the set of specificities. Five latent classes were found for a three-dimensional spatial model with specificities. Common dimensions were quantified psychophysically in terms of log-rise time, spectral centroid, and degree of spectral variation. The results further suggest that musical timbres possess specific attributes not accounted for by these shared perceptual dimensions. Weight patterns indicate that perceptual salience of dimensions and specificities varied across classes. A comparison of class structure with biographical factors associated with degree of musical training and activity was not clearly related to the class structure, though musicians gave more precise and coherent judgments than did non-musicians or amateurs. The model with latent classes and specificities gave a better fit to the data and made the acoustic correlates of the common dimensions more interpretable.

Adolescent↗

Location of rabbit auditory cortex and description of single unit activity.

The following data were obtained from microelectrode exploration of rabbit auditory cortex (AC). (1) Location of AC--Major portions of the rabbit's right hemisphere were stereotaxically mapped for auditory responsive and non-responsive regions. Auditory responsive regions were found to lie behind the rhinal sulcus, extending from its midpoint to its inferior border. (2) Frequency selectivity--Characteristic frequencies (CFs) ranged from 0.4 to 33 kHz. Sharp, multipeaked and broad tuning curves were described. (3) Response patterns--A variety of post-stimulus-time (PST) histogram shapes were obtained in response to tonal stimuli. The response of a given neuron often varied with changes in stimulus frequency or intensity. (4) Latency--Neural response latencies ranged from 10 to 100 msec following stimulus onset. (5) Spontaneous rate--In the absence of acoustic stimulation, spike discharges ranged from less than 1 spike/sec to 48 spikes/sec. Spontaneous rates of less than 5 spikes/sec were measured in 42% of the neurons. Rates in excess of 15 spikes/sec occurred in fewer than 18% of the cells examined. (6) Response variability--Stimulus-evoked activity was seen to vary over time. Changes consisted of an overall increase or attenuation of discharge rate, as well as modifications of selected portions of the response pattern (PST histogram shape). The characteristics of single unit activity in rabbit auditory cortex are consistent with findings reported in other species.

Animals↗

The effect of auditory stimulus rate on [14C]2-deoxyglucose uptake in rabbit inferior colliculus.

The relationship between auditory stimulus rate and level of [14C]2-deoxyglucose (2-DG) uptake was examined using the 2-DG technique developed by Sokoloff as an indication of glucose uptake and neural activity. Conscious restrained albino rabbits were given one monaural tone either once per second or once per minute for 45 min, or served as unstimulated controls. Autoradiographs of the inferior colliculi from these animals were compared densitometrically. It was found that the rabbits stimulated once per second had markedly elevated levels of 2-DG uptake compared to both other groups: the rabbits stimulated once per minute and to unstimulated control rabbits. In addition, the inferior colliculus autoradiographs of the rabbits stimulated with pure tones showed a distinctive banding pattern in central nucleus different from that seen in control rabbits.

Acoustic Stimulation↗

Auditory response properties of neurons in the anterior ectosylvian sulcus of the cat.

The auditory response properties of single neurons in the fundus and banks of the anterior ectosylvian sulcus (AES) were studied with simple dichotic stimuli (viz. noise- and tone-bursts) in cats anaesthetized with alpha-chloralose. Neurons within AES showed simple onset responses, were most commonly excited by stimulation of both ears, and showed either broad tuning or multiple high best frequencies. Some neurons were also tested for visual responsiveness and it was found that auditory cells and visual cells were intermingled within the sulcus. A small percentage of cells responded to both auditory and visual stimulation. Overall, the response properties of AES neurons differed from those of nearby auditory cortical fields. The region of AES studied appears to be outside the recently defined fourth somatosensory area (SIV), but overlaps para-SIV found deeper in the sulcus. It appears that deep within the sulcus and along most of its length there is a population of auditory, somatosensory and visual cells; to delineate this auditory population from the surrounding auditory cortical fields this region has been designated Field AES.

Animals↗

Mechanical and acoustical influences on spontaneous oto-acoustic emissions.

Spontaneous, tone-like emissions produced by normal ears and measured in the closed outer ear canal can be affected by mechanical and acoustical events. Such effects can be measured in steady-state conditions as well as for transient stimulation, and are seen in response to the stapedius reflex, to ear canal air pressure changes, and to the presentation of external tones. Frequency and level of the emissions follow certain characteristics which are described and discussed. The emissions seem to react with 2 ms delay and with an exponential rise and decay, the time constant of which is about 13 ms.

Acoustic Impedance Tests↗

Synchronized responses of primary auditory fibre-populations in Caiman crocodilus (L.) to single tones and clicks.

Measurements of the responses to tones and clicks were made from single primary auditory fibres of the caiman. The distribution of the amplitude and phase of the fundamental component of the response rate modulation over the best frequencies of the fibres is comparable to that reported in the cat, despite the fact that the basilar membrane in caiman is only 4.5 mm long. However, much higher intensities are needed in the caiman (75-85 dB SPL) than reported in the cat (20 dB SPL) to obtain systematic distributions of the phase of the responses, probably due to the larger scatter of the phase responses in the caiman. The slopes of the phase distributions are very similar to those in cat. Single unit phase responses as a function of stimulus frequency at 85 dB SPL can be approximated by one, or in fibres with low best frequency, two straight lines. At lower intensities the deviation of the phase-frequency responses from a straight line increases as the group delay at the best frequency becomes larger. The shortest latencies of click responses are obtained with rarefaction clicks. Group delay estimates obtained from the responses to clicks and from the straight line approximations of the phase-frequency responses are related in a way expected for linear filter systems and accurately predict the measured distributions of the phase of the responses over the neural best frequency. The obtained group delays and click latencies in the caiman are very similar to those reported by other workers in the cat, the squirrel monkey and the treefrog, despite large morphological and probably functional differences of their inner ears. The click latencies are also very similar to those in the pigeon. The results are consistent with the existence of a mechanical travelling wave reported previously on the basilar membrane of the caiman, but at the same stimulus level the phase characteristic of the present single unit responses is steeper and the wave length estimates from the neural population phase distributions are shorter than those observed directly in the motion of the basilar membrane. Since the neural responses are an indirect estimate of the basilar membrane motion it cannot be decided whether the difference between neural and mechanical data is due to deterioration of the basilar membrane responses during the direct measurements or whether the basilar membrane response is sharpened by additional tuning mechanisms.

Animals↗

Pre-attentive and attentive processing of temporal and frequency characteristics within long sounds.

Attention effects on the processing of deviations in the duration and the frequency dimension of a long sound were investigated in three conditions: (1) when auditory stimuli were ignored, (2) when they were attended and frequency dimension was task-relevant, and (3) when they were attended and duration dimension was task-relevant. The mismatch negativity (MMN) of the event-related potential (ERP) to infrequent shortenings of a sound (600 ms vs. 1000 ms) and to infrequent frequency modulations at one of nine possible intervals within the sound (change from 440 Hz to 480 Hz and back to 440 Hz, e.g. in the 600-650 ms interval) was measured. Duration MMN was slightly enhanced when directing attention towards the frequency dimension but notably enhanced when attention was focused on duration. The early phase of frequency-modulated MMN was of equal amplitude in all three conditions, and the late phase was equally enlarged in both attend conditions. Interestingly, MMN to frequency-modulated deviants decreased the later the deviation occurred within the sound; there was no indication for an MMN being present in Ignore condition when frequency modulations occurred 400 ms after sound onset or later. Thus, with increasing temporal distance between the onset of a sound and the onset of a deviation within the sound (e.g. frequency modulation or sound offset), MMN for frequency modulations and duration shortenings decreases. This suggests that the initial part of a sound ( approximately 300 ms) contributes more to the unitary sound representation underlying MMN than the later parts.

Acoustic Stimulation↗

Perception of flight information from EFIS displays.

A pilot's perception of variables presented on the Electronic Flight Instrument System, EFIS, was investigated. A stimulus response technique was used to determine the accuracy and speed of the perception process. By varying the exposure time of the stimuli, it is shown that the perception of a variable's magnitude is faster and more accurate than the perception of the first derivative or rate of that variable. Results of experiments on roll and pitch attitude perception, the influence of scale division, and the perception of the indicated airspeed, are shown.

Aerospace Medicine↗

Amygdalohippocampal involvement in tinnitus and auditory memory.

CONCLUSION: The preliminary results suggest that in chronic unilateral tinnitus the contralateral amygdalohippocampal complex does seem to be involved in tinnitus perception of pure tones. OBJECTIVES: Functional neuroimaging studies have revealed that the hippocampus and amygdala are involved in tinnitus perception. The amygdala and hippocampus are supplied by the anterior choroidal artery. Selective amobarbital injections in the anterior choroidal artery result in a non-functional amygdalohippocampal area for 10 min. The aim of this study was to assess the influence of this procedure on tinnitus perception. PATIENTS AND METHODS: Amobarbital (80 mg in total) was injected selectively in two sessions in the left and right anterior choroidal artery in six male patients with tinnitus: four with unilateral tinnitus, two with bilateral tinnitus. Of the patients with unilateral tinnitus, three had right-sided tinnitus and one had left-sided tinnitus. The average age was 57.3 years (range 43-69). Average tinnitus duration was 5.3 years (range 1-10). The differences in visual analogue scale (VAS) scores before and after the amytal tests were analysed. RESULTS: Amytal injection ipsilateral to the side where the tinnitus was perceived resulted in a maximum of 30% tinnitus suppression, whereas amytal injection contralateral to the tinnitus side yielded a 60-70% tinnitus suppression in three patients with unilateral chronic tinnitus (>4 years). Only pure tone tinnitus was suppressed, white noise was not. Two patients with bilateral tinnitus had no suppression, irrespective of the tinnitus type. A third patient without clinical tinnitus suppression had tinnitus of more recent origin (1.5 years).

Adult↗

Why "sounds are judged longer than lights": application of a model of the internal clock in humans.

Three experiments, using temporal generalization and verbal estimation methods, studied judgements of durations of auditory (500-Hz tone) and visual (14-cm blue square) stimuli. With both methods, auditory stimuli were judged longer, and less variable, than visual ones. The verbal estimation experiments used stimuli from 77 to 1183 msec in length, and the slope of the function relating mean estimate to real length differed between modalities (but the intercept did not), consistent with the idea that a pacemaker generating duration representations ran faster for auditory than for visual stimuli. The different variability of auditory and visual stimuli was attributed to differential variability in the operation of a switch of a pacemaker-accumulator clock, and experimental data suggested that such switch effects were separable from changes in pacemaker speed. Overall, the work showed how a clock model consistent with scalar timing theory, the leading account of animal timing, can address an issue derived from the classical literature on human time perception.

Attention↗

Hemispheric differences in processing tone frequency and amplitude modulations.

Transient frequency and amplitude modulations (FMs, AMs) of sound are requisite to speech recognition. We recorded whole-head magnetoencephalographic signals from seven subjects to binaural 620 ms 667 Hz tones, with 3, 30, or 300 ms FMs or AMs in the beginning or middle of the tone. Responses were significantly larger and earlier for FMs than AMs, for rapid than slow modulations, and for modulations at the beginning (BEG) than in the middle (MID) of the sound. BEG 3 ms FMs elicited strongest signals in the left and MID 3 ms FMs in the right hemisphere. Fast MID modulations produced significantly stronger responses in the right than left hemisphere. These differences may reflect different functions of the left and right hemisphere in speech perception.

Adult↗