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Neurophysiologic bases of speech discrimination.

The mismatch negativity (MMN) is an automatic cortical evoked potential that signifies the brain's detection of acoustic change. In other words, the MMN reflects the neurophysiologic processes that underlie auditory discrimination. As such, the MMN provides an objective tool for evaluating central auditory mechanisms involved in speech perception. We are using the MMN to study the central auditory processes that encode acoustic changes important for speech perception in 1) normal-hearing adults and children, 2) individuals with impaired auditory systems (including persons with learning disabilities, attention deficit disorders, cochlear implants), and 3) an animal model. Specifically, we have demonstrated that the MMN provides information about the central processing of fine acoustic differences, the neuroanatomic pathways that encode acoustic change, central auditory processing in the presence of peripheral hearing deficits, and central auditory system plasticity. In addition, we have considered methodological challenges associated with measuring the MMN in individual subjects. Several methodological issues--including appropriate stimuli, stimulus presentation variables, the recording protocol and environment, and validation of the MMN in individuals--are discussed.

Child↗

Infants' and adults' perception of scale structure.

Adults and 9-month-old infants were required to detect mistuned tones in multitone sequences. When 7-tone versions of a common nursery tune were generated from the Western major scale (unequal scale steps) or from an alternative scale (equal steps), infants detected the mistuned tones more accurately in the unequal-step context than in the equal-step context (Experiment 1). Infants and adults were subsequently tested with 1 of 3 ascending-descending scales (15 tones): (a) a potentially familiar scale (major) with unequal steps, (b) an unfamiliar scale with unequal steps, and (c) an unfamiliar scale with equal steps. Infants detected mistuned tones only in the scales with unequal steps (Experiment 2). Adults performed better on the familiar (major) unequal-step scale and equally poorly on both unfamiliar scales (Experiments 3 and 4). These findings are indicative of an inherent processing bias favoring unequal-step scales.

Age Factors↗

Neuronal populations in the human brain extracting invariant relationships from acoustic variance.

The ability to extract invariant relationships from physically varying stimulation is critical for example to categorical perception of complex auditory information such as speech and music. Human subjects were presented with tone pairs randomly varying over a wide frequency range, there being no physically constant tone pair at all. Instead, the invariant feature was either the direction of the tone pairs (ascending: the second tone was higher in frequency than the first tone) or the frequency ratio (musical interval) of the two tones. The subjects ignored the tone pairs, and instead attended a silent video. Occasional deviant pairs (either descending in direction or having a different frequency ratio) elicited the mismatch negativity (MMN) of the event-related potential, demonstrating the existence of neuronal populations which automatically (independently of attention) extract invariant relationships from acoustical variance.

Acoustic Stimulation↗

Vowel and consonant judgments are not independent when cued by the same information.

Despite many attempts to define the major unit of speech perception, none has been generally accepted. In a unique study, Mermelstein (1978) claimed that consonants and vowels are the appropriate units because a single piece of information (duration, in this case) can be used for one distinction without affecting the other. In a replication, this apparent independence was found, instead, to reflect a lack of statistical power: The vowel and consonant judgments did interact. In another experiment, interdependence of two phonetic judgments was found in responses based on the fricative noise and the vocalic formants of a fricative-vowel syllable. These results show that each judgment made on speech signals must take into account other judgments that compete for information in the same signal. An account is proposed that takes segments as the primary units, with syllables imposing constraints on the shape they may take.

Adult↗

Critical modulation frequency based on detection of AM versus FM tones.

The ratios between the modulation index (eta) for just noticeable FM of a sinusoidally modulated pure tone and the degree of modulation (m) for just noticeable AM at the same carrier and the same modulation frequency were measured at carrier frequencies of 0.125, 0.25, 0.5, 1, 2, 4, and 8 kHz. Signal levels were 20 dB SL and 50 dB SPL or 80 dB SPL. At low modulation frequencies, for example, 8 Hz, AM and FM elicit very different auditory sensations (i.e., a fluctuation in loudness or pitch, respectively). In this case, eta and m show different values for just noticeable modulation. Since both stimuli have almost equal amplitude spectra if eta equals m (m less than 0.3), the difference in detection thresholds reflects differences in the phase relation between carrier and sidebands in AM and FM. With increasing modulation frequency, the eta-m ratio decreases and reaches unity at a modulation frequency called the "critical modulation frequency" (CMF). At modulation frequencies above the CMF, the same modulation thresholds are obtained for AM and FM. Therefore, it can be concluded that the difference in phase between the two types of stimuli is not perceived in this range. At center frequencies below 1 kHz, where phase errors caused by headphones and ear canal presumably are small, the CMF is useful in estimating critical bandwidth.

Adult↗

Binaural sensitivity as a function of interaural electrode position with a bilateral cochlear implant user.

Experiments were conducted with a single, bilateral cochlear implant user to examine interaural level and time-delay cues that putatively underlie the design and efficacy of bilateral implant systems. The subject's two implants were of different types but custom equipment allowed presentation of controlled bilateral stimuli, particularly those with specified interaural time difference (ITD) and interaural level difference (ILD) cues. A lateralization task was used to measure the effect of these cues on the perceived location of the sensations elicited. For trains of fixed-amplitude, biphasic current pulses at 100 pps, the subject demonstrated sensitivity to an ITD of 300 micros, providing evidence of access to binaural information. The choice of bilateral electrode pair greatly influenced ITD sensitivity, suggesting that electrode pairings are likely to be an important consideration in the effort to provide binaural advantages. The selection of bilateral electrode pairs showing sensitivity to ITD was partially aided by comparisons of the pitch elicited by individual electrodes in each ear (when stimulated alone with fixed-amplitude current pulses at 813 pps): specifically, interaural electrodes with similar pitches were more likely (but not certain) to show ITD sensitivity. Significant changes in lateral position occurred with specific electrode pairs. With five bilateral electrode pairs of 14 tested, ITDs of 300 and 600 micros moved an auditory image significantly from right to left. With these same pairs, ILD changes of approximately 11% of the dynamic range (in microApp) moved an auditory image from the far left to the far right-significantly farther than the nine pairs not showing significant ITD sensitivity. However, even these nine pairs did show response changes as a function of the interaural (or confounding monaural) level cue. Overall, insofar as the access to bilateral cues demonstrated herein generalizes to other subjects, it provides hope that the normal binaural advantages for speech recognition and sound localization can be made available to bilateral implant users.

Acoustic Stimulation↗

Amplitude compression in cochlear implants artificially restricts the perception of temporal asymmetry.

This paper presents a study in which five cochlear implantees were asked to discriminate the timbre of stimuli with temporally asymmetric envelopes. Stimuli were damped and ramped sinusoids presented acoustically. They were transformed by the speech processor of the implant and were presented through one electrode. All cochlear implantees could discriminate the damped and ramped sinusoids when the half-life was 4 ms, the carrier frequency was 400 Hz, and the period of the envelope was 50 ms. In a second experiment, timbre discrimination performance was measured as a function of half-life for two cochlear implantees. Both showed that timbre discrimination was possible over the range 1-24 ms. In normal-hearing listeners, the range is 1-64 ms and in cochlear implantees, stimulated directly without the speech processor, the range is 1-300 ms. At long half-lives, the decrease in discrimination performance observed with the speech processor appears to be due to the amplitude compression applied by the device. The present results suggest that it may be important to ensure that cochlear implants do not restrict temporal asymmetry unduly when applying compression to control level.

Adult↗

Effect of stimulus bandwidth on the perception of /s/ in normal- and hearing-impaired children and adults.

Recent studies with adults have suggested that amplification at 4 kHz and above fails to improve speech recognition and may even degrade performance when high-frequency thresholds exceed 50-60 dB HL. This study examined the extent to which high frequencies can provide useful information for fricative perception for normal-hearing and hearing-impaired children and adults. Eighty subjects (20 per group) participated. Nonsense syllables containing the phonemes /s/, /f/, and /O/, produced by a male, female, and child talker, were low-pass filtered at 2, 3, 4, 5, 6, and 9 kHz. Frequency shaping was provided for the hearing-impaired subjects only. Results revealed significant differences in recognition between the four groups of subjects. Specifically, both groups of children performed more poorly than their adult counterparts at similar bandwidths. Likewise, both hearing-impaired groups performed more poorly than their normal-hearing counterparts. In addition, significant talker effects for /s/ were observed. For the male talker, optimum performance was reached at a bandwidth of approximately 4-5 kHz, whereas optimum performance for the female and child talkers did not occur until a bandwidth of 9 kHz.

Adult↗

Alteration of frequency range for binaural beats in acute low-tone hearing loss.

The effect of acute low-tone sensorineural hearing loss (ALHL) on the interaural frequency difference (IFD) required for perception of binaural beats (BBs) was investigated in 12 patients with unilateral ALHL and 7 patients in whom ALHL had lessened. A continuous pure tone of 30 dB sensation level at 250 Hz was presented to the contralateral, normal-hearing ear. The presence of BBs was determined by a subjective yes-no procedure as the frequency of a loudness-balanced test tone was gradually adjusted around 250 Hz in the affected ear. The frequency range in which no BBs were perceived (FRNB) was significantly wider in the patients with ALHL than in the controls, and FRNBs became narrower in the recovered ALHL group. Specifically, detection of slow BBs with a small IFD was impaired in this limited (10 s) observation period. The significant correlation between the hearing level at 250 Hz and FRNBs suggests that FRNBs represent the degree of cochlear damage caused by ALHL.

Acute Disease↗

Measuring the critical band for speech.

The current experiments were designed to measure the frequency resolution employed by listeners during the perception of everyday sentences. Speech bands having nearly vertical filter slopes and narrow bandwidths were sharply partitioned into various numbers of equal log- or ERBN-width subbands. The temporal envelope from each partition was used to amplitude modulate a corresponding band of low-noise noise, and the modulated carriers were combined and presented to normal-hearing listeners. Intelligibility increased and reached asymptote as the number of partitions increased. In the mid- and high-frequency regions of the speech spectrum, the partition bandwidth corresponding to asymptotic performance matched current estimates of psychophysical tuning across a number of conditions. These results indicate that, in these regions, the critical band for speech matches the critical band measured using traditional psychoacoustic methods and nonspeech stimuli. However, in the low-frequency region, partition bandwidths at asymptote were somewhat narrower than would be predicted based upon psychophysical tuning. It is concluded that, overall, current estimates of psychophysical tuning represent reasonably well the ability of listeners to extract spectral detail from running speech.

Adolescent↗

Vibrotactile perception of suprasegmental features of speech: a comparison of single-channel and multichannel instruments.

The recognition of three suprasegmental aspects of speech--the number of syllables in a word, the stress pattern of a word, and rising or falling intonation patterns--through a single-channel tactile device and through a 24-channel tactile vocoder, using two groups of normal-hearing subjects, was compared. All subjects received an initial pretest on three recognition tasks, one for each prosodic feature. Half the subjects from each group then received 12 h of training with feedback on the tasks and stimuli used in the pretest. All subjects received a post-test which contained physically different stimuli from those previously tested. Performance was significantly better on the syllable-number and syllabic stress tasks with the single-channel than with the multichannel device on both the pre- and post-tests; no difference was found for the intonation task. Performance on the post-test was poorer for all trained subjects compared to their final training results, suggesting that cues learned in training were not readily transferable to new stimuli, even those with similar prosodic characteristics. Overall, the results provide support for the notion that certain prosodic features of speech may be conveyed more readily when the waveform envelope is preserved.

Adult↗

Ontogenetic change in the analysis of sound frequency in the infant rat.

Previous research has shown that there is an ontogenetic shift in the spatial code of sound frequency in the cochlea and central auditory nuclei. During ontogenesis, a given area of the basilar membrane or central auditory nuclei is maximally stimulated by sounds of progressively higher frequencies. Here, a similar ontogenetic shift in the rat's "perception" of sound frequency is reported. Animals were trained with a tone (CS) paired with footshock (US) and generalization of their conditioned response (CR) to different frequencies was assessed either 0.5 or 72 hr later. When 15-day-old animals were trained with an 8-kHz tone and tested 0.5 hr later, greater suppression of activity (CR) was evoked by the 8-kHz tone than by either a 12-kHz or a 6-kHz tone. When tested 72 hr after training, however, greater suppression of activity was evoked by a 12-kHz tone than by the 8-kHz training stimulus or by a 24-kHz tone. This shift in the peak of the generalization gradient could not be accounted for by changes in unconditioned responses to the test stimuli. Additionally, the observed shift in the generalization gradient was age-dependent. It was not observed when pups were trained at 18 days old and tested 72 hr later. The implications of these data for studies of the ontogenesis of learning and memory are discussed.

Age Factors↗

Speech processing studies using an acoustic model of a multiple-channel cochlear implant.

The speech perception of two multiple-channel cochlear implant patients was compared with that of three normally hearing listeners using an acoustic model of the implant for 22 different speech tests. The tests used included a minimal auditory capabilities battery, both closed-set and open-set word and sentence tests, speech tracking and a 12-consonant confusion study using nonsense syllables. The acoustic model represented electrical current pulses by bursts of noise and the effects of different electrodes were represented by using bandpass filters with different center frequencies. All subjects used a speech processor that coded the fundamental voicing frequency of speech as a pulse rate and the second formant frequency of speech as the electrode position in the cochlea, or the center frequency of the bandpass filter. Very good agreement was found for the two groups of subjects, indicating that the acoustic model is a useful tool for the development and evaluation of alternative cochlear implant speech processing strategies.

Aged↗

Hemispheric specialization for processing auditory nonspeech stimuli.

The left hemisphere specialization for speech perception might arise from asymmetries at more basic levels of auditory processing. In particular, it has been suggested that differences in "temporal" and "spectral" processing exist between the hemispheres. Here we used functional magnetic resonance imaging to test this hypothesis further. Fourteen healthy volunteers listened to sequences of alternating pure tones that varied in the temporal and spectral domains. Increased temporal variation was associated with activation in Heschl's gyrus (HG) bilaterally, whereas increased spectral variation activated the superior temporal gyrus (STG) bilaterally and right posterior superior temporal sulcus (STS). Responses to increased temporal variation were lateralized to the left hemisphere; this left lateralization was greater in posteromedial HG, which is presumed to correspond to the primary auditory cortex. Responses to increased spectral variation were lateralized to the right hemisphere specifically in the anterior STG and posterior STS. These findings are consistent with the notion that the hemispheres are differentially specialized for processing auditory stimuli even in the absence of linguistic information.

Acoustic Stimulation↗

Is poor frequency modulation detection linked to literacy problems? A comparison of specific reading disability and mild to moderate sensorineural hearing loss.

Specific reading disability (SRD) is now widely recognised as often being caused by phonological processing problems, affecting analysis of spoken as well as written language. According to one theoretical account, these phonological problems are due to low-level problems in auditory perception of dynamic acoustic cues. Evidence for this has come from studies showing poor discrimination of frequency-modulated from unmodulated tones. We measured frequency modulation detection limens (FMDLs) in 16 children with specific reading disability (SRD group), 16 children with mild to moderate hearing loss (SNH group) and 16 age-matched controls (CA group) aged 8-14. To obtain information about possible mechanisms used in frequency modulation detection, FMDLs were measured at modulation rates of 2 and 20 Hz, both in the absence and the presence of amplitude modulation, intended to force listeners to rely, if possible, upon phase-locking cues. Although both the SNH and SRD groups showed a trend for elevated FMDLs at both 2 and 20 Hz, these differences reached statistical significance for the SNH group alone. However, the SNH group had no evidence of literacy impairments. This study thus shows that impairments in perceiving dynamically modulated auditory stimuli do not necessarily lead to difficulty in learning to read.

Adolescent↗

Tonal perceptions in normal laryngeal, esophageal, and electrolaryngeal speech of Mandarin.

The present study attempted to investigate if alaryngeal speakers of Mandarin Chinese could differentially produce the four tone levels of Mandarin: high-level, mid-rising, falling-rising and high-falling, as perceived by native listeners. Syllables /ma/ and /ba/ produced at four different tone levels by 8 normal laryngeal (NL), 7 standard esophageal (SE), and 8 electrolaryngeal (EL) speakers of Mandarin were perceived by 8 naïve listeners. Results from the listening experiments showed a higher percent correct identification of tone for NL speech than SE and EL speech. Perceptual data showed different patterns of tone confusion associated with the three types of speech. SE and EL speakers were not able to produce the four tone levels as accurately as were NL speakers. NL speakers achieved a near-perfect level of accuracy in signaling tonal contrasts. SE speakers produced the falling-rising and high-falling tones more accurately than high-level and mid-rising tones, but tonal confusions existed between mid-rising tone and falling-rising tone, and between high-level tone and high-falling tone. In EL phonation, high-level tone was produced more accurately than the other tones which were often misidentified as a high-level tone.

Aged↗

Dynamic causal modeling of evoked responses in EEG and MEG.

Neuronally plausible, generative or forward models are essential for understanding how event-related fields (ERFs) and potentials (ERPs) are generated. In this paper, we present a new approach to modeling event-related responses measured with EEG or MEG. This approach uses a biologically informed model to make inferences about the underlying neuronal networks generating responses. The approach can be regarded as a neurobiologically constrained source reconstruction scheme, in which the parameters of the reconstruction have an explicit neuronal interpretation. Specifically, these parameters encode, among other things, the coupling among sources and how that coupling depends upon stimulus attributes or experimental context. The basic idea is to supplement conventional electromagnetic forward models, of how sources are expressed in measurement space, with a model of how source activity is generated by neuronal dynamics. A single inversion of this extended forward model enables inference about both the spatial deployment of sources and the underlying neuronal architecture generating them. Critically, this inference covers long-range connections among well-defined neuronal subpopulations. In a previous paper, we simulated ERPs using a hierarchical neural-mass model that embodied bottom-up, top-down and lateral connections among remote regions. In this paper, we describe a Bayesian procedure to estimate the parameters of this model using empirical data. We demonstrate this procedure by characterizing the role of changes in cortico-cortical coupling, in the genesis of ERPs. In the first experiment, ERPs recorded during the perception of faces and houses were modeled as distinct cortical sources in the ventral visual pathway. Category-selectivity, as indexed by the face-selective N170, could be explained by category-specific differences in forward connections from sensory to higher areas in the ventral stream. We were able to quantify and make inferences about these effects using conditional estimates of connectivity. This allowed us to identify where, in the processing stream, category-selectivity emerged. In the second experiment, we used an auditory oddball paradigm to show that the mismatch negativity can be explained by changes in connectivity. Specifically, using Bayesian model selection, we assessed changes in backward connections, above and beyond changes in forward connections. In accord with theoretical predictions, there was strong evidence for learning-related changes in both forward and backward coupling. These examples show that category- or context-specific coupling among cortical regions can be assessed explicitly, within a mechanistic, biologically motivated inference framework.

Auditory Pathways↗

The perception of temporal structure and auditory evoked brain activity.

The processing and perception of auditory signals depends on the temporal structure of stimulus characteristics. We studied 26 healthy subjects who participated in psychophysical experiments and in electrophysiological recordings of auditory evoked potentials from C2, C3, C4, T3 and T4. Stimuli consisted of tone series presented binaurally as tones or gaps with a base duration of 100 ms. In the psychophysical experiments, difference thresholds as indicators of temporal discrimination performance were significantly lower for tones than for gaps. In the electrophysiological recordings, gaps often failed to elicit N100 components. Tones produced shortest component latencies with largest amplitudes. In addition, brain activity was strongest at C2, and showed a symmetrical fall-off over both hemispheres. N100 components had significantly longer latencies and smaller amplitudes when they were evoked by the end of the gap (i.e. with the continuation of the tone) than by tones. Our data illustrate how the temporal structure of auditory stimuli affects neuronal responses of the brain. Similar effects were observed in psychophysical and electrophysiological experiments, and we were able to demonstrate a direct relationship between subjective sensory thresholds and auditory evoked brain activity.

Adult↗