Sound and vibration sensitivity of VIIIth nerve fibers in the frogs Leptodactylus albilabris and Rana pipiens pipiens

1. Responses of 73 fibers to dorso-ventral vibration were recorded in the saccular and utricular branchlets of Rana pipiens pipiens using a ventral approach. The saccular branchlet contained nearly exclusively vibration-sensitive fibers (33 out of 36) with best frequencies (BFs) between 10 and 70 Hz, whereas none of the 37 fibers encountered in the utricular branchlet responded to dorso-ventral vibrations. 2. Using a dorsal approach we recorded from the VIIIth nerve near its entry in the brainstem and analyzed responses to both sound and vibration stimuli for 65 fibers in R. pipiens pipiens and 25 fibers in Leptodactylus albilabris. The fibers were classified as amphibian papilla (AP), basilar papilla (BP), saccular or vestibular fibers based on their location in the nerve. Only AP and saccular fibers responded to vibrations. The AP-fibers responded to vibrations from 0.01 cm/s2 and to sound from 40 dB SPL by increasing their spike rate. Best frequencies (BFs) ranged from 60 to 900 Hz, and only fibers with BFs below 500 Hz responded to vibrations. The fibers had identical BF's for sound and vibration. The saccular fibers had BFs ranging from 10 to 80 Hz with 22 fibers having BFs at 40-50 Hz. The fibers responded to sound from 70 dB SPL and to vibrations from 0.01 cm/s2. 3. No differences in sensitivity, tuning or phase-locking were found between the two species, except that most BP-fibers in R. pipiens pipiens had BFs from 1.2 to 1.4 kHz, whereas those in L. albilabris had BFs from 2.0 to 2.2 kHz (matching the energy peak of L. albilabris' mating call). 4. The finding that the low-frequency amphibian papilla fibers are extremely sensitive to vibrations raises questions regarding their function in the behaving animal. They may be substrate vibration receptors, respond to sound-induced vibrations or bone-conducted sound.     

Mechanical responses to two-tone distortion products in the apical and basal turns of the mammalian cochlea

Mechanical responses to one- and two-tone acoustic stimuli were recorded from the cochlear partition in the apical turn of the chinchilla cochlea, the basal turn of the guinea pig cochlea, and the hook region of the guinea pig cochlea. The most sensitive or "best" frequencies (BFs) for the sites studied were approximately 500 Hz, 17 kHz, and 30 kHz, respectively. Responses to the cubic difference tone (CDT), 2F1 - F2 (where F1 and F2 are the frequencies of the primary stimuli), were characterized at each site. Responses to the quadratic difference tone (QDT), F2 - F1, were also characterized in the apical turn preparations (QDT responses were too small to measure in the basal cochlea). The observed responses to BF QDTs and CDTs and to BF CDTs at each site appeared similar in many ways; the relative magnitudes of the responses were highest at low-to-moderate sound pressure levels (SPLs), for example, and the absolute magnitudes grew nonmonotonically with increases in the level of either primary (L1 or L2) alone. The peak effective levels of the CDT and QDT responses were also similar, at around -20 dB re L1 and/or L2. In other respects, however, the responses to CDTs and QDTs and to BF CDTs at each site behaved quite differently. At low-to-moderate SPLs, for example, most CDT phase leads decreased with increases in either L1 or L2, whereas most QDT phase leads increased with increasing L1 and varied little with L2. Most CDT responses also varied monotonically with equal-level primaries (i.e., when L1 = L2), whereas most QDT responses varied nonmonotonically. Different responses also varied in different ways when F1 and F2 were varied. Apical turn QDT responses were observed over a very wide F1/F2 range (F1 = 1-12 kHz), but were usually largest for stimuli <2-4 kHz. Apical turn CDT levels decreased (at rates of approximately 40-80 dB/octave) only when the frequency ratio F2/F1 increased beyond approximately 1.4-1.5. In the basal turn and hook regions, the CDT levels depended nonmonotonically on F2/F1 with the eventual rates of decrease being approximately 200 dB/octave. Optimal frequency ratios for the CDT increased from (F2 < 1.1F1) to (F2 approximately 1.2F1) with increasing SPL in the basal turn, but were stable at around F2/F1 approximately 1.05 in the hook region. CDT phase leads tended to increase with increasing F2/F1 in all three regions of the cochlea, particularly at low-to-moderate SPLs. These findings are discussed in relation to previous studies of cochlear mechanics, physiology, and psychophysics.     

Discriminability of vowel representations in cat auditory-nerve fibers after acoustic trauma

This paper attempts to connect deficits seen in the neural representation of speech with perceptual deficits. Responses of auditory-nerve fibers were studied in cats exposed to acoustic trauma. Four synthetic steady-state vowels were used as test signals; these stimuli are identical, except that the second format (F2) resonator in the synthesizer was set to 1.4, 1.5, 1.7, or 2 kHz, producing four spectra that differ mainly in the vicinity of the F2 frequency. These stimuli were presented to a large population (523) of auditory-nerve fibers in four cats with sloping high-frequency threshold shifts that reached 50-70 dB at 2-4 kHz. In normal animals, May et al. [Auditory Neurosci 3, 135-162 (1996)] showed previously that the discharge rates of fibers with best frequencies near the F2 frequencies provide enough information to allow discrimination of these stimuli at the performance levels shown by cats in behavioral experiments. Here it is shown that, after acoustic trauma, there is essentially no rate information which would allow the vowels with different F2 frequencies to be discriminated. However, information that could allow discrimination remains in the temporal (phase-locked) aspects of the responses.     

Effects of stapedius-muscle contractions on the masking of auditory-nerve responses

There has been little exploration of the mechanisms by which stapedius muscle contractions reduce the masking of responses to high-frequency sounds by low-frequency sounds. To fill this gap in knowledge, controlled stapedius contractions were elicited with direct shocks in anesthetized cats, and measurements were made of the effects of these contractions on the masking of single auditory-nerve fibers and on the attenuation of middle-ear transmission. The results show that the stapedius-induced reductions of masking can be much larger than the attenuations of low-frequency sound. With a 300-Hz band of masking noise centered at 500 Hz, and signal tones at 6 or 8 kHz, unmasking effects over 40 dB were observed for sounds 100 dB SPL or less. The data suggest that much larger unmasking might occur. The observed unmasking can be explained completely by a linear stapedius-induced attenuation of sound transmission through the middle ear and a nonlinear growth rate of masking for auditory-nerve fibers. No central effects are required. It is argued that the reduction of the upward spread of masking is probably one of the most important functions of the stapedius muscle.     

Effects of bilateral olivocochlear lesions on vowel formant discrimination in cats

Operant conditioning procedures were used to measure the effects of bilateral olivocochlear lesions on the cat's discrimination thresholds for changes in the second formant frequency (deltaF2) of the vowel /epsilon/. Three cats were tested with the formant discrimination task under quiet conditions and in the presence of continuous broadband noise at signal-to-noise ratios (S/Ns) of 23, 13, and 3 dB. In quiet, vowel levels of 50 and 70 dB produced average deltaF2s of 42 and 47 Hz, respectively, and these thresholds did not change significantly in low levels of background noise (S/Ns = 23 and 13 dB). Average deltaF2s increased to 94 and 97 Hz for vowel levels of 50 and 70 dB in the loudest level of background noise (S/N = 3 dB). Average deltaF2 thresholds in quiet and in lower noise levels were only slightly affected when the olivocochlear bundle was lesioned by making bilateral cuts into the floor of the IVth ventricle. In contrast, post-lesion deltaF2 thresholds in the highest noise level were significantly larger than pre-lesion values; the most severely affected subject showed post-lesion discrimination thresholds well over 200 Hz for both 50 and 70 dB vowels. These results suggest that olivocochlear feedback may enhance speech processing in high levels of ambient noise.     

The missing fundamental in vowel height perception

Traunmüller (1981) suggested that the tonotopic distance between the first formant (F1) and the fundamental frequency (F0) is a major determinant of perceived vowel height. In the present study, subjects identified a vowel-height continuum ranging in formant pattern from /I/to/epsilon/, at five F0 values. Increasing F0 led to an increased probability of /I/responses (i.e., the phoneme boundary shifted toward the /epsilon/ end of the continuum). Various conditions of filtering out the lower harmonics of the stimuli caused only marginal shifts of the phoneme boundary. The experiments provide evidence against interpretations of Traunmüller's (1981) results that claim that vowel height is determined by the distance between F1 and the lowest harmonic that is present in the basilar membrane excitation pattern.     

Modeling inhibition of type II units in the dorsal cochlear nucleus

Type II units in the dorsal cochlear nucleus (DCN) are characterized by vigorous but nonmonotonic responses to best frequency tones as a function of sound pressure level, and relatively weak responses to noise. A model of DCN neural circuitry was used to explore two hypothetical mechanisms by which neurons may be endowed with type II unit response properties. Both mechanisms assume that type II units receive excitatory input from auditory nerve (AN) fibers and inhibitory input from an unspecified class of cochlear nucleus interneurons that also receive excitatory AN input. The first mechanism, a lateral inhibition (LI) model, supposes that type II units receive inhibitory input from a number of narrowly tuned interneurons whose best frequencies (BFs) flank the BF of the type II unit. Tonal stimuli near BF result in only weak inhibitory input, but broadband stimuli recruit enough lateral inhibitors to greatly weaken the type II unit response. The second mechanism, a wideband inhibition (WBI) model, supposes that type II units receive inhibitory input from interneurons that are broadly tuned so that they respond more vigorously to broadband stimuli than to tones. Physiological and anatomical evidence points to the possible existence of such a class of neurons in the cochlear nucleus. The model extends an earlier computer model of an iso-frequency DCN patch to multiple frequency slices and adds a population of interneurons to provide the inhibition to model type II units (called 12-cells). The results show that both mechanisms accurately simulate responses of type II units to tones and noise. An experimental paradigm for distinguishing the two mechanisms is proposed.     

A note on vowel centralization in stuttering and nonstuttering individuals

Inferences were made regarding vocal tract vowel space during fluently produced utterances through examination of the first two formant frequencies. Fifteen adult males served as subjects, representing separate groups of untreated and treated individuals who stutter and nonstuttering controls. The steady-state portion of formant one (F1) and formant two (F2) was examined in the production of various CVC tokens containing the vowels /i/, /u/, and /a/. Vocal tract vowel space was estimated three ways. The first analysis scheme involved measurement of formant frequency spacing. The second measure involved calculating the area of the vowel space triangle. The third measure was based on calculating the average Euclidean distance from each subject's midpoint "centroid" vocal tract position to the corresponding /i/, /u/, and /a/ points on the vowel triangle. The formant frequency spacing measures proved to be most revealing of group differences, with the untreated stutterers showing significantly greater vowel centralization than the treated group and control group. Discussion focuses on the vocal tract articulation characterizing fluent speech productions and possible treatment implications for persons who stutter.     

Difference limens for formant patterns of vowel sounds

Studies of thresholds for discrimination of formant frequency variation in synthetic vowel sounds have been predominantly limited to variations in a single formant. Here, differences limens (DLs) are presented for multiformant variations expressed in measures of delta F and as distances in the auditory-perceptual space (APS) proposed by J. D. Miller [J. Acoust. Soc. Am. 85, 2114-2134 (1989)]. DLs for four subjects were estimated along 102 synthetic vowel continua representing five patterns of formant variation [(1) single variation in F1; (2) single variation in F2; (3) parallel simultaneous variation in F1 and F2; (4) opposing simultaneous variation in F1 and F2; and parallel simultaneous variation in F1, F2, and F3] and 17 within- or between-category vowel sounds. Minimal uncertainty methodology was employed utilizing an adaptive up-down procedure with a cued, two-interval forced-choice (2IFC) task. The results of this experiment reflect smaller DLs for both single- and multiple-formant changes than have been found in the past and also suggest that discrimination of parallel multiformant variation is significantly better than opposing multiformant or single-formant variation.     

The effects of decreased audibility produced by high-pass noise masking on cortical event-related potentials to speech sounds/ba/and/da

This study investigated the effects of decreased audibility produced by high-pass noise masking on cortical event-related potentials (ERPs) N1, N2, and P3 to the speech sounds /ba/and/da/presented at 65 and 80 dB SPL. Normal-hearing subjects pressed a button in response to the deviant sound in an oddball paradigm. Broadband masking noise was presented at an intensity sufficient to completely mask the response to the 65-dB SPL speech sounds, and subsequently high-pass filtered at 4000, 2000, 1000, 500, and 250 Hz. With high-pass masking noise, pure-tone behavioral thresholds increased by an average of 38 dB at the high-pass cutoff and by 50 dB one octave above the cutoff frequency. Results show that as the cutoff frequency of the high-pass masker was lowered, ERP latencies to speech sounds increased and amplitudes decreased. The cutoff frequency where these changes first occurred and the rate of the change differed for N1 compared to N2, P3, and the behavioral measures. N1 showed gradual changes as the masker cutoff frequency was lowered. N2, P3, and behavioral measures showed marked changes below a masker cutoff of 2000 Hz. These results indicate that the decreased audibility resulting from the noise masking affects the various ERP components in a differential manner. N1 is related to the presence of audible stimulus energy, being present whether audible stimuli are discriminable or not. In contrast, N2 and P3 were absent when the stimuli were audible but not discriminable (i.e., when the second formant transitions were masked), reflecting stimulus discrimination. These data have implications regarding the effects of decreased audibility on cortical processing of speech sounds and for the study of cortical ERPs in populations with hearing impairment.     

Temporal patterns of the responses of auditory-nerve fibers to low-frequency tones

The temporal response patterns of auditory-nerve fibers to low-frequency tones were studied in anesthetized cats using period histograms. 'Peak-splitting' was observed mostly in fibers with lower characteristic frequencies (CF < 2 kHz) and with lower-frequency stimulation (< or = 500 Hz). The occurrence of peak-splitting, the number of peaks, and the time between the peaks were all dependent upon the stimulus frequency. The phases of responses, although complex functions of stimulus frequency, intensity, and the fiber's CF, clearly showed traveling-wave characteristics for all frequencies at or above 100 Hz. The amount of phase change with intensity was generally small for lower-frequency stimuli (< approximately 50 degrees), although larger phase changes (e.g., approximately 180 degrees) were occasionally seen with higher-frequency stimuli. At 50 and 100 Hz, the phase of neural responses in the basal region roughly corresponds to the maximum velocity of the basilar membrane towards scala tympani (as inferred from cochlear microphonic recordings).     

A multidimensional scaling analysis of vowel discrimination in humans and monkeys

Multidimensional scaling (MDS) was used to compare perceptual maps for 10 synthetic English vowels in humans and Old World monkeys (Macaca fuscata and Cercopithecus albogularis). Subjects discriminated among the vowels using a repeating background procedure, and reaction times were submitted to an MDS analysis to derive measures of perceive similarity. The dimensions that emerged related to the frequencies of the first (F1), second (F2), and third (F3) formants. Human data indicated a good match to previous MDS studies using rating procedures or confusion matrices: The dominant dimension mapped onto vowel F2, the phonetically most important formant, and the second and third dimensions mapped onto F1 and F3, respectively. For monkeys, equal weightings occurred for F1 and F2, and F3 was not clearly represented. Monkey sensitivity to the formants appeared to relate to formant amplitudes. If monkeys are giving an accurate representation of the psychoacoustic relations among the formants, then our human results suggest that species-specific mechanisms, reflecting the salience of the phonetic feature of advancement, may contribute to vowel coding in humans.     

Exploration and development of China's large-scale blast furance

Large-scale BF is the tendency of domestic and foreign Iron-making technology.Due to the limit of resource,technology and equipment,we can not reach the planned goal after putting the big BF into production,it is of great significance to explore the key technologies again for the sound development of the large-scale BF.The paper highlights the unique advantages of large-scale BF and the practical performance of BFs in Baosteel,and explores the related parameter choice,comprehensively matched longevity te...     

General characteristics and suppression tuning properties of the distortion-product otoacoustic emission 2f1-f2 in the barn owl

The distortion-product otoacoustic emission (DPOAE) 2f1-f2 was measured in the ear canal of the barn owl. DPOAE were elicited by primary tones in 11 frequency regions from 1 to 9 kHz. The highest DPOAE output levels and best thresholds were found for f1 frequencies of 4 to 7 kHz and additionally at the lowest f1 frequency investigated. In some cases, the DPOAE sound pressures were only 37 dB below the primary-tone levels (PTL). The optimal primary-tone frequency ratios ranged from 1.05 to 1.45 and varied strongly among the different frequency regions investigated. The largest optimal ratios were measured in the middle frequency range for f1. At lower and higher f1, the optimal ratios decreased. DPOAE levels could be suppressed in a frequency-selective way by adding a third tone. As in other non-mammals, the best suppressive frequencies were near f1, suggesting DPOAE generation near the frequency place of this primary tone. This is in contrast to what is known for mammalian species, where the DPOAE is thought to be generated near f2. To obtain 6 dB of suppression of the DPOAE level, suppressor-tone levels ranging from 13 dB below to 4 dB above the primary-tone level were necessary. The Q10dB-values of suppression tuning curves increased as a function of frequency up to a value of 15.8. This tendency resembled the increase in frequency selectivity of auditory nerve fibers in this species.     

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.     

Estimating cochlear filter response properties from distortion product otoacoustic emission (DPOAE) phase delay measurements in normal hearing human adults

This study examined cochlear filter response properties derived from f1- and f2-sweep phase delay difference measures in 60 normal hearing human adults. Seven different f2 frequencies ranging from 1.1 to 9.2 kHz were presented (f2/f1 ratios of 1.1-1.3). F2 intensity level was varied in 5 dB steps from 30 to 50 dB SPL (the level of f1 was 15 dB above the level of f2). DPOAE delay estimates in a f2-sweep paradigm are longer than in a f1-sweep paradigm at the same frequency and intensity. This indicates that the f2-sweep DPOAE phase delay is composed of a greater proportion of the filter response time at the site of DPOAE generation than the f1-sweep delay. This proportion was isolated by subtracting f1-sweep DPOAE delays from f2-sweep delays at similar f2 frequencies and intensities. Under the assumption of linearity and minimum phase the impulse response of the filter at each f2 stimulus level was calculated from the mean phase delay difference. Frequency response properties were calculated by Fourier transformation of the impulse response at each f2 frequency and intensity. High frequency low intensity impulse responses had longer response times and narrower frequency bandwidths than low frequency high intensity responses. The Q10dB values of DPOAE derived tuning curves ranged from 2.4 (1.5 kHz) to 7.3 (8.5 kHz).     

Speaking-rate-induced variability in F2 trajectories

This study examined speaking-rate-induced spectral and temporal variability of F2 formant trajectories for target words produced in a carrier phrase at speaking rates ranging from fast to slow. F2 onset frequency measured at the first glottal pulse following the stop consonant release in target words was used to quantify the extent to which adjacent consonantal and vocalic gestures overlapped; F2 target frequency was operationally defined as the first occurrence of a frequency minimum or maximum following F2 onset frequency. Regression analyses indicated 70% of functions relating F2 onset and vowel duration were statistically significant. The strength of the effect was variable, however, and the direction of significant functions often differed from that predicted by a simple model of overlapping, sliding gestures. Results of a partial correlation analysis examining interrelationships among F2 onset, F2 target frequency, and vowel duration across the speaking rate range indicated that covariation of F2 target with vowel duration may obscure the relationship between F2 onset and vowel duration across rate. The results further suggested that a sliding based model of acoustic variability associated with speaking rate change only partially accounts for the present data, and that such a view accounts for some speakers' data better than others.     

A model for the responses of low-frequency auditory-nerve fibers in cat

A computational model was developed for the responses of low-frequency auditory-nerve (AN) fibers in cat. The goal was to produce realistic temporal response properties and average discharge rates in response to simple and complex stimuli. Temporal and average-rate properties of AN responses change as a function of sound-pressure level due to nonlinearities in the auditory periphery. The input stage of the AN model is a narrow-band filter that simulates the mechanical tuning of the basilar membrane. The parameters of this filter vary continuously as a function of stimulus level via a feedback mechanism, simulating the compressive nonlinearity associated with the mechanics of the basilar membrane. A memoryless, saturating nonlinearity and two low-pass filters simulate transduction and membrane properties of the inner hair cell (IHC). A diffusion model for the IHC-AN synapse introduces adaptation. Finally, a nonhomogeneous Poisson process, modified by absolute and relative refractoriness, provides the output discharge times. Responses to several different stimuli are presented. These responses illustrate nonlinear temporal response properties that cannot be achieved with linear models for AN fibers.     

Transient-evoked and 2F1-F2 distortion product oto-acoustic emissions in dogs: preliminary findings

Transient (click)-evoked oto-acoustic emissions (TEOAEs) and distortion product oto-acoustic emissions (DPOAEs) were recorded in a feasibility study in 7 healthy mixed-breed dogs using the ILO 92 OAE analyser (Otodynamics, Hartfield, UK). Five dogs were found to have normal hearing in both ears and 2 dogs in the left ear only following otoscopy, tympanometry and auditory brainstem response audiometry. Twelve sets of TEOAEs (click-evoked) to 80 dB peSPL click stimulus and 9 sets of DPOAEs (2F1-F2) to 8 different stimulus levels of the primary tones (L1/L2) were collected at 11 test frequencies (F2) in these normal-hearing dogs. TEOAEs were successfully recorded in 11 of the 12 ears using the default user setting and in all 12 ears using the quickscreen program. DPOAEs were successfully recorded in all 9 ears tested. While the TEOAEs parameters matched those for humans, the average signal-to-noise ratio of DPOAEs was considerably higher in the dogs. Stimulus levels at 55/55, 55/45 and 55/35 dB SPL were demonstrated to produce DPOAEs that seem to reflect the active dynamic status of the outer hair cell system. Postmortem DPOAEs at these stimulus levels and TEOAEs at 80 Db peSPL could not be elicited 5 min following euthanasia of dogs. However, DPOAEs could still be recorded albeit with reduced amplitude at stimulus levels where L1 > 55 dB SPL. The results suggest that TEOAEs and DPOAEs in dogs have the potential to provide valuable insights into their mechanisms of generation, and the specific role and behaviour of outer hair cells of the cochlea in certain pathological conditions, particularly in drug-induced ototoxicity, in humans.     

湘钢高炉与芬兰拉赫高炉生产的比较分析

湘钢1号高炉与芬兰拉赫高炉虽然外形相近.但原料条件、高炉操作、炉前工作、人员配置等方面有很大的差异,结果是湘钢1号高炉的主要技术经济指标居全国的中下游,而芬兰拉赫高炉的主要技术经济指标在世界领先.