A central spectrum model: a synthesis of auditory-nerve timing and place cues in monaural communication of frequency spectrum

A probabilistic psychophysical model for monaural communication from the auditory nerve to the brain is given in the form of a tonotopic display of stimulus spectrum, termed central spectrum. The model builds upon prior research demonstrating the potential of neural timing cues from the auditory nerve for conveying information on complex spectra, and was designed to meet the quantified demands of the psychophysics of frequency measurement. The central spectrum magnitude at each frequency is determined by the response of the auditory-nerve fibre with characteristic frequency matching that frequency. An interval histogram from each fiber is passed through a filter matched to the characteristic frequency of the fiber. This output versus characteristic frequency defines the central spectrum. Detailed analysis demonstrates that efficient probabilistic processing of the central spectrum described known psychophysical properties of frequency measurement in discrimination and periodicity pitch experiments. Psychophysical models based upon the central spectrum model followed by optimum probabilistic pattern recognition are potentially relevant for predicting human communication limits in response to arbitrary sounds of speech and music.     

Connectivities and synchronous firing in cortical neuronal networks

Network connectivities ((-)k) of cortical neural cultures are studied by synchronized firing and determined from measured correlations between fluorescence intensities of firing neurons. The bursting frequency (f) during synchronized firing of the networks is found to be an increasing function of (-)k. With f taken to be proportional to (-)k, a simple random model with a (-)k dependent connection probability p((-)k).has been constructed to explain our experimental findings successfully.     

Synchronized discharge rate representation of voice-onset time in the chinchilla auditory nerve

Responses of chinchilla auditory nerve fibers to synthesized stop consonant syllables differing in voice-onset time (VOT) were obtained. The syllables, heard as /ga/-/ka/ or /da/-/ta/, were similar to those previously used by others in psychophysical experiments with human and chinchilla subjects. Synchronized discharge rates of neurons tuned to frequencies near the first formant increased at the onset of voicing for VOTs longer than 20 ms. Stimulus components near the formant or the neuron's characteristic frequency accounted for the increase. In these neurons, synchronized response changes were closely related to the same neuron's average discharge rates [D. G. Sinex and L. P. McDonald, J. Acoust. Soc. Am. 83, 1817-1827 (1988)]. Neurons tuned to frequency regions near the second and third formants usually responded to components near the second formant prior to the onset of voicing. These neurons' synchronized discharges could be captured by the first formant at the onset of voicing or with a latency of 50-60 ms, whichever was later. Since these neurons' average rate responses were unaffected by the onset of voicing, the latency of the synchronized response did provide as much additional neural cue to VOT. Overall, however, discharge synchrony did not provide as much information about VOT as was provided by the best average rate responses. The results are compared to other measurements of the peripheral encoding of speech sounds and to aspects of VOT perception.     

Masking of short stimuli by noises with spiked spectra: II. Time summation and frequency selectivity of hearing in a narrow frequency range

The thresholds of masking of short high-frequency pulses with either different durations (1.25–25 ms) and similar central frequency or different central frequencies (3.6–4.4 kHz) but similar durations were measured to reveal manifestations of the properties of peripheral encoding in auditory perception. Noises with a spiked amplitude spectrum structure were used as maskers. The central frequency and the frequency band of a masker were 4 and 1 kHz, respectively. The central frequencies of a stimulus and a masker being equal, the noise the central frequency of which coincided with the frequency corresponding to a dip of an indented spectrum was called an off_(rip)-frequency masker. Owing to the off_(rip)-masker, stimuli-induced masking thresholds were formed taking into account excitation in a narrow region of a basila membrane and auditory nerve fibers with characteristic frequencies from a narrow range. High-frequency pulses with an envelope in the form of the Gaussian function and sinusoidal filling were used as stimuli. At masker levels of 30 dB above the auditory threshold, frequencies of off_(rip)-masker spectra spikes of 500–2000 Hz, and a central stimulus frequency of 4 kHz, the thresholds of tonal stimuli (25 ms in duration) masking in two out of three probationers were higher than the thresholds of masking of compact stimuli (1.25 ms in duration). In the third probationer, on the contrary, the thresholds of tonal stimuli masking were lower than the thresholds of compact stimuli masking. At masker levels of 50 dB, individual threshold differences disappeared. The obtained results were interpreted in the context of implementation of different methods of auditory encoding of the intensity. The methods were based on either the average frequency of auditory nerve pulsations or the number of fibers participating in the response. The interpretation was also carried out in the context of revealing manifestations of nonlinear properties of basila membrane displacements in auditory thresholds. The fact that the dependence of detection thresholds of compact stimuli on their central frequency in one of the two probationers did not reveal the minimum in case of coincidence of off_(rip)-masker and stimulus frequencies pointed to the presence of an auditory “problem zone” that was likely to be localized at the periphery of the auditory system.     

Pitch discrimination of patterned electric stimulation

One reason for the poor pitch performance in current cochlear-implant users may be the highly synchronized neural firing in electric hearing that lacks stochastic properties of neural firing in normal acoustic hearing. This study used three different electric stimulation patterns, jittered, probabilistic, and auditory-model-generated pulses, to mimic some aspects of the normal neural firing pattern in acoustic hearing. Pitch discrimination was measured at standard frequencies of 100, 250, 500, and 1000 Hz on three Nucleus-24 cochlear-implant users. To test the utility of the autocorrelation pitch perception model in electric hearing, one, two, and four electrodes were stimulated independently with the same patterned electric stimulation. Results showed no improvement in performance with any experimental pattern compared to the fixed-rate control. Pitch discrimination was actually worsened with the jittered pattern at low frequencies (125 and 250 Hz) than that of the control, suggesting that externally introduced stochastic properties do not improve pitch perception in electric stimulation. The multiple-electrode stimulation did not improve performance but did not degrade performance either. The present results suggest that both "the right time and the right place" may be needed to restore normal pitch perception in cochlear-implant users.     

Speech processing in the auditory system. II: Lateral inhibition and the central processing of speech evoked activity in the auditory nerve

A biologically realistic model of a uniform lateral inhibitory network (LIN) is shown capable of extracting from the complex spatio-temporal firing patterns of the cat's auditory nerve the formants and low-order harmonics of synthetic voiced speech stimuli. The model provides a realistic mechanism to utilize the temporal aspects of the firing and thus supports the hypothesis that the neural coding of complex sounds in terms of average rates can be supplemented by the information coded in the synchronous firing. At low levels of intensity the LIN can sharpen the average rate profiles. At moderate and high levels the LIN uses the cues available in the distribution of phases of the synchronous activity which exhibit rapid relative phase shifts at specific characteristic frequency (CF) locations (corresponding to the frequencies of the low-order harmonics in the stimulus). These temporal phase shifts manifest themselves at the input of the LIN as steep and localized spatial discontinuities in the instantaneous pattern of activity across the fiber array. The LIN enhances its output from these spatially steep input regions while suppressing its output from spatially smooth input regions (where little phase shifts occur). In this manner the LIN recreates from the response patterns a representation of the stimulus spectrum using the temporal cues as spatial markers of the stimulus components rather than as absolute measures of their frequencies. Similar results are obtained with various lateral inhibitory topologies, e.g., recurrent versus nonrecurrent, single versus double layer, and linear versus nonlinear.     

Modeling “preattention” and “attention” information processing by synchronization of neural activity

Oscillatory neural-network preattention and attention models are examined. A two-layer network of Wilson-Cowan oscillators is used to show that two-frequency oscillations can appear in response to a compound stimulus. It is shown that these oscillations can be synchronized at the low frequency, which can be interpreted as feature binding. Partial synchronization is studied in a model of a network of phased oscillators with a central element. Formulas are given for approximation of the average frequency of the central oscillator for small and large networks. The results describe the effect of a distracting stimulus on attention focus.Institute of Mathematical Problems of Biology, Russian Academy of Sciences. Translated from Izvestiya Vysshikh Uchebnykh Zavedenii, Radiofizika, Vol. 37, No. 8, pp. 933–944, August, 1994.     

Synchronized firing of FitzHugh-Nagumo neurons by noise

We investigate the influence of noise on synchronization between the spiking activities of neurons with external impulsive forces. We first analyze the dependence of the synchronized firing on the amplitude and the angular frequency of the impulsive force in the noise-free system. Three cases (regular spiking, traveling wave, and chaotic spiking) with low synchronized firing are chosen to study effects due to noise. In each case we find that small noise can be a promoter of synchronization phenomena in neural activities, by choosing an appropriate noise intensity acting on some of the neurons.     

基于全相位滤波器的电子耳蜗汉语音调感知及改进研究

针对电子耳蜗音调信息感知差的问题,在多导电子耳蜗机理模型的基础上,利用零(或全)相位滤波器以及希尔伯特变换,提取并分解16个通道信号的包络和精细结构,用多种方式嵌合成声音,对正常人耳测听合成音的音调信息,以确定信号的包络和精细结构对音调感知的影响。测试结果表明:精细结构相对包络对音调感知起着更重要的作用,且该作用主要表现在低频段(约1.2 kHz以内)通道上。研究发现,在固定通道上,精细结构与通道中心频率和相位信息决定的包络出现时刻对应,其音调感知与低通道电极刺激的脉冲发放时间有关。研究结论:在低频段电极上增加刺激脉冲发放时刻的控制对提高电子耳蜗音调感知是重要的,同时应注意滤波器的相位保真。      

Diversity of adaptation patterns in responses of eighth nerve fibers in the bullfrog, Rana catesbeiana

The firing patterns of eighth nerve fibers in the bullfrog, Rana catesbeiana, were analyzed for responses to long duration tone bursts at best excitatory frequency ( BEF ) and at frequencies along the upper and lower boundaries of the excitatory tuning curve of each fiber. These firing patterns were used as an index of the degree of short-term adaptation of each fiber. Amphibian papilla fibers (with BEFs 100-1000 Hz) exhibited marked diversity in their firing patterns to BEF tones, ranging from very flat or tonic (sustained responses throughout the duration of the stimulus) to very peaked or phasic (responding primarily or exclusively to stimulus onset). Moreover, the degree of short-term adaptation shown by an individual fiber varied with stimulating frequency. The firing patterns of amphibian papilla fibers tended to become more tonic as stimulus frequency was lowered below BEF ; conversely, as stimulus frequency was increased above BEF , firing patterns either showed little change from that at BEF , or became more phasic. A similar frequency dependence of adaptation has not been reported in responses of mammalian eighth nerve fibers with comparable BEFs . The firing patterns of basilar papilla fibers ( BEFs greater than 1000 Hz) remained similar in response to both BEF and non- BEF tones. These data reveal that the firing patterns and degrees of short-term adaptation of amphibian papilla fibers vary considerably across the tuning curve, whereas those of basilar papilla fibers remain relatively more constant with changes in stimulating frequency.(ABSTRACT TRUNCATED AT 250 WORDS)     

Self-synchronization of coupled oscillators with hysteretic responses

We analyze a large system of nonlinear phase oscillators with sinusoidal nonlinearity, uniformly distributed natural frequencies and global all-to-all coupling, which is an extension of Kuramoto's model to second-order systems. For small coupling, the system evolves to an incoherent state with the phases of all the oscillators distributed uniformly. As the coupling is increased, the system exhibits a discontinuous transition to the coherently synchronized state at a pinning threshold.of the coupling strength, or to a partially synchronized oscillation coherent state at a certain threshold below the pinning threshold. If the coupling is decreased from a strong coupling with all the oscillators synchronized coherently, this coherence can persist until the depinning threshold which is less than the pinning threshold, resulting in hysteretic synchrony depending on the initial configuration of the oscillators. We obtain analytically both the pinning and depinning threshold and also expalin the discontinuous transition at the thresholds for the underdamped case in the large system size limit. Numerical exploration shows the oscillatory partially coherent state bifurcates at the depinning threshold and also suggests that this state persists independent of the system size. The system studied here provides a simple model for collective behaviour in damped driven high-dimensional Hamiltonian systems which can explain the synchronous firing of certain fireflies or neural oscillators with frequency adaptation and may also be applicable to interconnected power systems.     

Neuronal responses to amplitude-modulated and pure-tone stimuli in the guinea pig inferior colliculus, and their modification by broadband noise

Neuronal responses were recorded to pure and to sinusoidally amplitude-modulated (AM) tones at the characteristic frequency (CF) in the central nucleus of the inferior colliculus of anesthetized guinea pigs. Temporal (synchronized) and mean-rate measures were derived from period histograms locked to the stimulus modulation waveform to characterize the modulation response. For stimuli presented in quiet, the modulation gain at low frequencies of modulation (approx less than 50 Hz) was inversely proportional to the neuron's mean firing rate in response to both the modulated stimulus and to a pure tone at an equivalent level. In 43% of units the mean discharge rates in response to the AM stimuli were greatest for those modulation frequencies that generated the largest temporal responses. These discharge-rate maxima occurred at signal intensities corresponding to the steeply sloping part of the neuron's pure-tone rate-intensity function (RIF). The change in mean-rate response to modulated stimuli, as a function of intensity, was qualitatively similar to the pure-tone RIF. Adding broadband noise to the modulated stimulus increased the neuron's temporal response to low modulation frequencies. This increase in modulation gain was correlated with mean firing rate in response to the modulation but did not bear a simple relationship to the noise-induced shift in the RIF measured for a pure tone.     

The representation of the spectra and fundamental frequencies of steady-state single- and double-vowel sounds in the temporal discharge patterns of guinea pig cochlear-nerve fibers

Psychophysical results using double vowels imply that subjects are able to use the temporal aspects of neural discharge patterns. To investigate the possible temporal cues available, the responses of fibers in the cochlear nerve of the anesthetized guinea pig to synthetic vowels were recorded at a range of sound levels up to 95 dB SPL. The stimuli were the single vowels /i/ [fundamental frequency (f0) 125 Hz], /a/ (f0, 100 Hz), and /c/ (f0, 100 Hz) and the double vowels were /a(100),i(125)/ and /c(100),i(125)/. Histograms synchronized to the period of the double vowels were constructed, and locking of the discharge to individual harmonics was estimated from them by Fourier transformation. One possible cue for identifying the f0's of the constituents of a double vowel is modulation of the neural discharge with a period of 1/f0. Such modulation was found at frequencies between the formant peaks of the double vowel, with modulation at the periods of 100 and 125 Hz occurring at different places in the fiber array. Generation of a population response based on synchronized responses [average localized synchronized rate (ALSR): see Young and Sachs [J. Acoust. Soc. Am. 66, 1381-1403 (1979)] allowed estimation of the f0's by a variety of methods and subsampling the population response at the harmonics of the f0 of the constituent vowel achieved a good reconstruction of its spectrum. Other analyses using interval histograms and autocorrelation, which overcome some problems associated with the ALSR approach, also allowed f0 identification and vowel segregation. The present study has demonstrated unequivocally that the timing of the impulses in auditory-nerve fibers provides copious possible cues for the identification of the fundamental frequencies and spectra associated with each of the constituents of double vowels.     

The synergy between speech production and perception

Speech intelligibility is known to be relatively unaffected by certain deformations of the acoustic spectrum. These include translations, stretching or contracting dilations, and shearing of the spectrum (represented along the logarithmic frequency axis). It is argued here that such robustness reflects a synergy between vocal production and auditory perception. Thus, on the one hand, it is shown that these spectral distortions are produced by common and unavoidable variations among different speakers pertaining to the length, cross-sectional profile, and losses of their vocal tracts. On the other hand, it is argued that these spectral changes leave the auditory cortical representation of the spectrum largely unchanged except for translations along one of its representational axes. These assertions are supported by analyses of production and perception models. On the production side, a simplified sinusoidal model of the vocal tract is developed which analytically relates a few "articulatory" parameters, such as the extent and location of the vocal tract constriction, to the spectral peaks of the acoustic spectra synthesized from it. The model is evaluated by comparing the identification of synthesized sustained vowels to labeled natural vowels extracted from the TIMIT corpus. On the perception side a "multiscale" model of sound processing is utilized to elucidate the effects of the deformations on the representation of the acoustic spectrum in the primary auditory cortex. Finally, the implications of these results for the perception of generally identifiable classes of sound sources beyond the specific case of speech and the vocal tract are discussed.     

Vowel processing by a model of the auditory periphery: a comparison to eighth-nerve responses

A model of peripheral auditory processing that incorporates processing steps describing the conversion from the acoustic pressure-wave signal at the eardrum to the time course activity in auditory neurons has been developed. It can process arbitrary time domain waveforms and yield the probability of neural firing. The model consists of a concatenation of modules, one for each anatomical section of the periphery. All modules are based on published algorithms and current experimental data, except that the basilar membrane is assumed to be linear. The responses of this model to vowels alone and vowels in noise are compared to neural population responses, as determined by the temporal and average rate response measures of Sachs and Young [J. Acoust. Soc. Am. 66, 470-479, (1979)] and Young and Sachs [J. Acoust. Soc. Am. 66, 1381-1403, (1979)]. Despite the exclusion of nonlinear membrane mechanics, the model accurately predicts the vowel formant representations in the average localized synchronized rate (ALSR) responses and the saturating characteristics of the normalized average rate responses in quiet. When vowels are presented in background noise, the modeled ALSR responses are less robust than the neural data.     

Auditory nerve representation of a complex communication sound in background noise.

A population study of auditory nerve responses in the bullfrog, Rana catesbeiana, analyzed the relative contributions of spectral and temporal coding in representing a complex, species-specific communication signal at different stimulus intensities and in the presence of background noise. At stimulus levels of 70 and 80 dB SPL, levels which approximate that received during communication in the natural environment, average rate profiles plotted over fiber characteristic frequency do not reflect the detailed spectral fine structure of the synthetic call. Rate profiles do not change significantly in the presence of background noise. In ambient (no noise) and low noise conditions, both amphibian papilla and basilar papilla fibers phase lock strongly to the waveform periodicity (fundamental frequency) of the synthetic advertisement call. The higher harmonic spectral fine structure of the synthetic call is not accurately reflected in the timing of fiber firing, because firing is "captured" by the fundamental frequency. Only a small number of fibers synchronize preferentially to any harmonic in the call other than the first, and none synchronize to any higher than the third, even when fiber characteristic frequency is close to one of these higher harmonics. Background noise affects fiber temporal responses in two ways: It can reduce synchronization to the fundamental frequency, until fiber responses are masked; or it can shift synchronization from the fundamental to the second or third harmonic of the call. This second effect results in a preservation of temporal coding at high noise levels. These data suggest that bullfrog eighth nerve fibers extract the waveform periodicity of multiple-harmonic stimuli primarily by a temporal code.     

Factors that influence rate-versus-intensity relations in single cochlear nerve fibers of the gerbil.

The relationship between characteristic frequency (CF) and rate-intensity curve shape was examined in 144 cochlear nerve fibers obtained from 39 Mongolian gerbils. Quasi-steady-state firing rates were measured in response to tone bursts at the CF. From each intensity curve, estimates of slope, firing rate at saturation, and dynamic range were derived using nonlinear curve fitting. Saturation firing rate was depressed for stimuli with a high duty cycle, especially for units with low rates of spontaneous discharge. The distributions of slope and saturation firing rate differed for fibers with CFs above and below 3 to 4 kHz. The interrelation of slope, dynamic range, maximum driven rate, and spontaneous firing rate was also different for fibers with CFs above and below this band. This mid-CF transition is discussed in terms of possible longitudinal changes in the function of the gerbil cochlea.     

感知听觉场景分析的说话人识别

针对低信噪比说话人识别中缺失数据特征方法鲁棒性下降的问题,提出了一种采用感知听觉场景分析的缺失数据特征提取方法。首先求取语音的缺失数据特征谱,并由语音的感知特性求出感知特性的语音含量。含噪语音经过感知特性的语音增强和对其语谱的二维增强后求解出语音的分布,联合感知特性语音含量和缺失强度参数提取出感知听觉因子。再结合缺失数据特征谱把特征的提取过程分解为不同听觉场景进行区分地分析和处理,以增强说话人识别系统的鲁棒性能。实验结果表明,在-10 dB到10 dB的低信噪比环境下,对于4种不同的噪声,提出的方法比5种对比方法的鲁棒性均有提高,平均识别率分别提高26.0%,19.6%,12.7%,4.6%和6.5%。论文提出的方法,是一种在时-频域中寻找语音鲁棒特征的方法,更适合于低信噪比环境下的说话人识别。      

The temporal representation of speech in a nonlinear model of the guinea pig cochlea

The temporal representation of speechlike stimuli in the auditory-nerve output of a guinea pig cochlea model is described. The model consists of a bank of dual resonance nonlinear filters that simulate the vibratory response of the basilar membrane followed by a model of the inner hair cell/auditory nerve complex. The model is evaluated by comparing its output with published physiological auditory nerve data in response to single and double vowels. The evaluation includes analyses of individual fibers, as well as ensemble responses over a wide range of best frequencies. In all cases the model response closely follows the patterns in the physiological data, particularly the tendency for the temporal firing pattern of each fiber to represent the frequency of a nearby formant of the speech sound. In the model this behavior is largely a consequence of filter shapes; nonlinear filtering has only a small contribution at low frequencies. The guinea pig cochlear model produces a useful simulation of the measured physiological response to simple speech sounds and is therefore suitable for use in more advanced applications including attempts to generalize these principles to the response of human auditory system, both normal and impaired.     

Noise-induced synchronization in bidirectionally coupled type-I neurons

We present here some studies on noise-induced order and synchronous firing in a system of bidirectionally coupled generic type-I neurons. We find that transitions from unsynchronized to completely synchronized states occur beyond a critical value of noise strength that has a clear functional dependence on neuronal coupling strength and input values. For an inhibitory-excitatory (IE) synaptic coupling, the approach to a partially synchronized state is shown to vary qualitatively depending on whether the input is less or more than a critical value. We find that introduction of noise can cause a delay in the bifurcation of the firing pattern of the excitatory neuron for IE coupling.