Processing Of Speech Sounds Research Articles

Speech sound processing deficits and training-induced neural plasticity in rats with dyslexia gene knockdown.

In utero RNAi of the dyslexia-associated gene Kiaa0319 in rats (KIA-) degrades cortical responses to speech sounds and increases trial-by-trial variability in onset latency. We tested the hypothesis that KIA- rats would be impaired at speech sound discrimination. KIA- rats needed twice as much training in quiet conditions to perform at control levels and remained impaired at several speech tasks. Focused training using truncated speech sounds was able to normalize speech discrimination in quiet and background noise conditions. Training also normalized trial-by-trial neural variability and temporal phase locking. Cortical activity from speech trained KIA- rats was sufficient to accurately discriminate between similar consonant sounds. These results provide the first direct evidence that assumed reduced expression of the dyslexia-associated gene KIAA0319 can cause phoneme processing impairments similar to those seen in dyslexia and that intensive behavioral therapy can eliminate these impairments.

Dynamic modulation of shared sensory and motor cortical rhythms mediates speech and non-speech discrimination performance.

Oscillatory models of speech processing have proposed that rhythmic cortical oscillations in sensory and motor regions modulate speech sound processing from the bottom-up via phase reset at low frequencies (3–10 Hz) and from the top-down via the disinhibition of alpha/beta rhythms (8–30 Hz). To investigate how the proposed rhythms mediate perceptual performance, electroencephalographic (EEG) was recorded while participants passively listened to or actively identified speech and tone-sweeps in a two-force choice in noise discrimination task presented at high and low signal-to-noise ratios. EEG data were decomposed using independent component analysis and clustered across participants using principle component methods in EEGLAB. Left and right hemisphere sensorimotor and posterior temporal lobe clusters were identified. Alpha and beta suppression was associated with active tasks only in sensorimotor and temporal clusters. In posterior temporal clusters, increases in phase reset at low frequencies were driven by the quality of bottom-up acoustic information for speech and non-speech stimuli, whereas phase reset in sensorimotor clusters was associated with top-down active task demands. A comparison of correct discrimination trials to those identified at chance showed an earlier performance related effect for the left sensorimotor cluster relative to the left-temporal lobe cluster during the syllable discrimination task only. The right sensorimotor cluster was associated with performance related differences for tone–sweep stimuli only. Findings are consistent with internal model accounts suggesting that early efferent sensorimotor models transmitted along alpha and beta channels reflect a release from inhibition related to active attention to auditory discrimination. Results are discussed in the broader context of dynamic, oscillatory models of cognition proposing that top-down internally generated states interact with bottom-up sensory processing to enhance task performance.

Effects of Recurrent Acute Otitis Media on Cortical Speech-Sound Processing in 2-Year Old Children

To investigate at the age of 2 years the effects of childhood recurrent acute otitis media (RAOM) on central auditory processing by using cortical event-related potentials elicited by syllable stimuli. During a 1-year period, 22- to 26-month-old children fulfilling the criteria for tympanostomy tube insertion in Oulu University Hospital, Oulu, Finland, were recruited to the RAOM group (N = 20). The control group (N = 19) was matched by age, sex, and mother's educational level. In both groups, children were typically developing and had no family history of language disorder or developmental language problems. Finnish syllables /ke:/ and /pi:/ as standards and their variants with changes in frequency, intensity, vowel, consonant, and vowel duration as deviants were used to record P1, N2, and mismatch negativity (MMN) responses in the multifeature paradigm. The clinically healthy ears at the time of registration were a prerequisite for the participation. Children with RAOM and their controls showed the age-typical P1 and N2 responses with no differences in the amplitudes or latencies between the groups, which suggests unaffected basic encoding of sound features and sound representation formation. However, the groups showed different auditory discrimination profiles. In children with RAOM, frequency and vowel MMN amplitudes were increased. Furthermore, the MMN latency for the frequency change was shorter and the frequency MMN amplitude lateralized to the left hemisphere in the RAOM group instead of an adult-like right-hemispheric lateralization observed in the controls. The children with RAOM had a more anterior MMN amplitude scalp distribution for the intensity change than control children. In addition, the MMN amplitude elicited by consonant change was evenly distributed unlike in controls, who had a left-side preponderant lateralization. Taken together, these results suggest an elevated responsiveness for frequency, vowel, and intensity changes, and an immature pattern of discriminating small speech sound contrasts in children with RAOM. The results suggest that childhood RAOM does not affect the central auditory pathway integrity or sound encoding. However, RAOM may lead to aberrant preattentive discrimination of sound features even when the peripheral auditory input is normal. These results are clinically significant because even transient problems with auditory processing may delay language development.

Degraded speech sound processing in a rat model of fragile X syndrome

Fragile X syndrome is the most common inherited form of intellectual disability and the leading genetic cause of autism. Impaired phonological processing in fragile X syndrome interferes with the development of language skills. Although auditory cortex responses are known to be abnormal in fragile X syndrome, it is not clear how these differences impact speech sound processing. This study provides the first evidence that the cortical representation of speech sounds is impaired in Fmr1 knockout rats, despite normal speech discrimination behavior. Evoked potentials and spiking activity in response to speech sounds, noise burst trains, and tones were significantly degraded in primary auditory cortex, anterior auditory field and the ventral auditory field. Neurometric analysis of speech evoked activity using a pattern classifier confirmed that activity in these fields contains significantly less information about speech sound identity in Fmr1 knockout rats compared to control rats. Responses were normal in the posterior auditory field, which is associated with sound localization. The greatest impairment was observed in the ventral auditory field, which is related to emotional regulation. Dysfunction in the ventral auditory field may contribute to poor emotional regulation in fragile X syndrome and may help explain the observation that later auditory evoked responses are more disturbed in fragile X syndrome compared to earlier responses. Rodent models of fragile X syndrome are likely to prove useful for understanding the biological basis of fragile X syndrome and for testing candidate therapies.

Attention Fine-Tunes Auditory–Motor Processing of Speech Sounds

The earliest stages of cortical processing of speech sounds take place in the auditory cortex. Transcranial magnetic stimulation (TMS) studies have provided evidence that the human articulatory motor cortex contributes also to speech processing. For example, stimulation of the motor lip representation influences specifically discrimination of lip-articulated speech sounds. However, the timing of the neural mechanisms underlying these articulator-specific motor contributions to speech processing is unknown. Furthermore, it is unclear whether they depend on attention. Here, we used magnetoencephalography and TMS to investigate the effect of attention on specificity and timing of interactions between the auditory and motor cortex during processing of speech sounds. We found that TMS-induced disruption of the motor lip representation modulated specifically the early auditory-cortex responses to lip-articulated speech sounds when they were attended. These articulator-specific modulations were left-lateralized and remarkably early, occurring 60-100 ms after sound onset. When speech sounds were ignored, the effect of this motor disruption on auditory-cortex responses was nonspecific and bilateral, and it started later, 170 ms after sound onset. The findings indicate that articulatory motor cortex can contribute to auditory processing of speech sounds even in the absence of behavioral tasks and when the sounds are not in the focus of attention. Importantly, the findings also show that attention can selectively facilitate the interaction of the auditory cortex with specific articulator representations during speech processing.

The neural basis of sublexical speech and corresponding nonspeech processing: A combined EEG–MEG study

We addressed the neural organization of speech versus nonspeech sound processing by investigating preattentive cortical auditory processing of changes in five features of a consonant–vowel syllable (consonant, vowel, sound duration, frequency, and intensity) and their acoustically matched nonspeech counterparts in a simultaneous EEG–MEG recording of mismatch negativity (MMN/MMNm). Overall, speech–sound processing was enhanced compared to nonspeech sound processing. This effect was strongest for changes which affect word meaning (consonant, vowel, and vowel duration) in the left and for the vowel identity change in the right hemisphere also. Furthermore, in the right hemisphere, speech–sound frequency and intensity changes were processed faster than their nonspeech counterparts, and there was a trend for speech-enhancement in frequency processing. In summary, the results support the proposed existence of long-term memory traces for speech sounds in the auditory cortices, and indicate at least partly distinct neural substrates for speech and nonspeech sound processing.

Language Switching Makes Pronunciation Less Nativelike

It is well known that multilingual speakers’ nonnative productions are accented. Do these deviations from monolingual productions simply reflect the mislearning of nonnative sound categories, or can difficulties in processing speech sounds also contribute to a speaker’s accent? Such difficulties are predicted by interactive theories of production, which propose that nontarget representations, partially activated during lexical access, influence phonetic processing. We examined this possibility using language switching, a task that is well known to disrupt multilingual speech production. We found that these disruptions extend to the articulation of individual speech sounds. When native Spanish speakers are required to unexpectedly switch the language of production between Spanish and English, their speech becomes more accented than when they do not switch languages (particularly for cognate targets). These findings suggest that accents reflect not only difficulty in acquiring second-language speech sounds but also the influence of representations partially activated during on-line speech processing.

Action planning and predictive coding when speaking

Across the animal kingdom, sensations resulting from an animal's own actions are processed differently from sensations resulting from external sources, with self-generated sensations being suppressed. A forward model has been proposed to explain this process across sensorimotor domains. During vocalization, reduced processing of one's own speech is believed to result from a comparison of speech sounds to corollary discharges of intended speech production generated from efference copies of commands to speak. Until now, anatomical and functional evidence validating this model in humans has been indirect. Using EEG with anatomical MRI to facilitate source localization, we demonstrate that inferior frontal gyrus activity during the 300ms before speaking was associated with suppressed processing of speech sounds in auditory cortex around 100ms after speech onset (N1). These findings indicate that an efference copy from speech areas in prefrontal cortex is transmitted to auditory cortex, where it is used to suppress processing of anticipated speech sounds. About 100ms after N1, a subsequent auditory cortical component (P2) was not suppressed during talking. The combined N1 and P2 effects suggest that although sensory processing is suppressed as reflected in N1, perceptual gaps may be filled as reflected in the lack of P2 suppression, explaining the discrepancy between sensory suppression and preserved sensory experiences. These findings, coupled with the coherence between relevant brain regions before and during speech, provide new mechanistic understanding of the complex interactions between action planning and sensory processing that provide for differentiated tagging and monitoring of one's own speech, processes disrupted in neuropsychiatric disorders.

ERP correlates of word onset priming in infants and young children

Using word onset priming with early learned words, we tracked access to phonological representations and predictive phonological processing at 6, 12, 18, and 24 months after birth. Event-Related Potentials (ERPs) were recorded while participants heard German word onsets (primes) followed by disyllabic spoken words (targets). Primes and target onsets were either congruent or incongruent (ma – Mama vs. so – Mama [Engl. ‘mommy’]). For an adult control group, ERP differences were found for the N100 complex, which has been related to abstract auditory analysis; and for the P350 deflection, which has been related to lexical access. A combined analysis of all infants and young children revealed an immature instance of an N100 effect, suggesting adult-like abstract speech sound processing. A central negativity effect, which had formerly been obtained when adults or older children were engaged in a lexical decision task, suggests that adult-like predictive phonological processing is available early in infancy. However, the absence of a P350-like effect in the infant data suggests that adult-like access to phonological forms is not established in the first two years of life. Taken together, ERPs recorded in word onset priming proved useful in investigating early phonological processing without an explicit behavioral measure.

Neurophysiological evidence that musical training influences the recruitment of right hemispheric homologues for speech perception

Musicians have a more accurate temporal and tonal representation of auditory stimuli than their non-musician counterparts (Musacchia et al., 2007; Parbery-Clark et al., 2009a; Zendel and Alain, 2009; Kraus and Chandrasekaran, 2010). Musicians who are adept at the production and perception of music are also more sensitive to key acoustic features of speech such as voice onset timing and pitch. Together, these data suggest that musical training may enhance the processing of acoustic information for speech sounds. In the current study, we sought to provide neural evidence that musicians process speech and music in a similar way. We hypothesized that for musicians, right hemisphere areas traditionally associated with music are also engaged for the processing of speech sounds. In contrast we predicted that in non-musicians processing of speech sounds would be localized to traditional left hemisphere language areas. Speech stimuli differing in voice onset time was presented using a dichotic listening paradigm. Subjects either indicated aural location for a specified speech sound or identified a specific speech sound from a directed aural location. Musical training effects and organization of acoustic features were reflected by activity in source generators of the P50. This included greater activation of right middle temporal gyrus and superior temporal gyrus in musicians. The findings demonstrate recruitment of right hemisphere in musicians for discriminating speech sounds and a putative broadening of their language network. Musicians appear to have an increased sensitivity to acoustic features and enhanced selective attention to temporal features of speech that is facilitated by musical training and supported, in part, by right hemisphere homologues of established speech processing regions of the brain.

Examination of Complementarity in Speech and Emotional Vocalization Perception

Lateralization of cognitive functioning is a well-established principle of cerebral organization. The left and right hemispheres are known to play distinct and complementary roles in the processing of information. What is still unclear is whether these asymmetrically lateralized functions have a common or distinct developmental origin; are left and right processes lateralized through causal influences, or is the laterality of each function independently influenced? Left- and right-lateralized functions are commonly assessed in isolation, with little attention to the relationship in the degree and direction of lateralization within individuals. This relationship between left-hemisphere processing of speech sounds and right-hemisphere processing of emotional vocalizations was examined using dichotic listening tasks. An overall complementary pattern of lateralization was observed across participants, but no significant relationship was found for degree of lateralization of speech and emotional vocalization processing within individuals. These results support the view that functions in the left and right hemispheres are independently lateralized.

Personality, Category, and Cross-Linguistic Speech Sound Processing: A Connectivistic View

Category formation of human perception is a vital part of cognitive ability. The disciplines of neuroscience and linguistics, however, seldom mention it in the marrying of the two. The present study reviews the neurological view of language acquisition as normalization of incoming speech signal, and attempts to suggest how speech sound category formation may connect personality with second language speech perception. Through a questionnaire, (being thick or thin) ego boundary, a correlate found to be related to category formation, was proven a positive indicator of personality types. Following the qualitative study, thick boundary and thin boundary English learners native in Cantonese were given a speech-signal perception test using an ABX discrimination task protocol. Results showed that thick-boundary learners performed significantly lower in accuracy rate than thin-boundary learners. It was implied that differences in personality do have an impact on language learning.

Native language affects rhythmic grouping of speech

Perceptual attunement to one's native language results in language-specific processing of speech sounds. This includes stress cues, instantiated by differences in intensity, pitch, and duration. The present study investigates the effects of linguistic experience on the perception of these cues by studying the Iambic-Trochaic Law (ITL), which states that listeners group sounds trochaically (strong-weak) if the sounds vary in loudness or pitch and iambically (weak-strong) if they vary in duration. Participants were native listeners either of French or German; this comparison was chosen because French adults have been shown to be less sensitive than speakers of German and other languages to word-level stress, which is communicated by variation in cues such as intensity, fundamental frequency (F0), or duration. In experiment 1, participants listened to sequences of co-articulated syllables varying in either intensity or duration. The German participants were more consistent in their grouping than the French for both cues. Experiment 2 was identical to experiment 1 except that intensity variation was replaced by pitch variation. German participants again showed more consistency for both cues, and French participants showed especially inconsistent grouping for the pitch-varied sequences. These experiments show that the perception of linguistic rhythm is strongly influenced by linguistic experience.

Speech training alters tone frequency tuning in rat primary auditory cortex

Previous studies in both humans and animals have documented improved performance following discrimination training. This enhanced performance is often associated with cortical response changes. In this study, we tested the hypothesis that long-term speech training on multiple tasks can improve primary auditory cortex (A1) responses compared to rats trained on a single speech discrimination task or experimentally naïve rats. Specifically, we compared the percent of A1 responding to trained sounds, the responses to both trained and untrained sounds, receptive field properties of A1 neurons, and the neural discrimination of pairs of speech sounds in speech trained and naïve rats. Speech training led to accurate discrimination of consonant and vowel sounds, but did not enhance A1 response strength or the neural discrimination of these sounds. Speech training altered tone responses in rats trained on six speech discrimination tasks but not in rats trained on a single speech discrimination task. Extensive speech training resulted in broader frequency tuning, shorter onset latencies, a decreased driven response to tones, and caused a shift in the frequency map to favor tones in the range where speech sounds are the loudest. Both the number of trained tasks and the number of days of training strongly predict the percent of A1 responding to a low frequency tone. Rats trained on a single speech discrimination task performed less accurately than rats trained on multiple tasks and did not exhibit A1 response changes. Our results indicate that extensive speech training can reorganize the A1 frequency map, which may have downstream consequences on speech sound processing.

Increasing diversity of neural responses to speech sounds across the central auditory pathway

Neurons at higher stations of each sensory system are responsive to feature combinations not present at lower levels. As a result, the activity of these neurons becomes less redundant than lower levels. We recorded responses to speech sounds from the inferior colliculus and the primary auditory cortex neurons of rats, and tested the hypothesis that primary auditory cortex neurons are more sensitive to combinations of multiple acoustic parameters compared to inferior colliculus neurons. We independently eliminated periodicity information, spectral information and temporal information in each consonant and vowel sound using a noise vocoder. This technique made it possible to test several key hypotheses about speech sound processing. Our results demonstrate that inferior colliculus responses are spatially arranged and primarily determined by the spectral energy and the fundamental frequency of speech, whereas primary auditory cortex neurons generate widely distributed responses to multiple acoustic parameters, and are not strongly influenced by the fundamental frequency of speech. We found no evidence that inferior colliculus or primary auditory cortex was specialized for speech features such as voice onset time or formants. The greater diversity of responses in primary auditory cortex compared to inferior colliculus may help explain how the auditory system can identify a wide range of speech sounds across a wide range of conditions without relying on any single acoustic cue.

Brainstem auditory evoked potentials with the use of acoustic clicks and complex verbal sounds in young adults with learning disabilities

Purpose and backgroundAcoustic signals are transmitted through the external and middle ear mechanically to the cochlea where they are transduced into electrical impulse for further transmission via the auditory nerve. The auditory nerve encodes the acoustic sounds that are conveyed to the auditory brainstem. Multiple brainstem nuclei, the cochlea, the midbrain, the thalamus, and the cortex constitute the central auditory system. In clinical practice, auditory brainstem responses (ABRs) to simple stimuli such as click or tones are widely used. Recently, complex stimuli or complex auditory brain responses (cABRs), such as monosyllabic speech stimuli and music, are being used as a tool to study the brainstem processing of speech sounds. We have used the classic ‘click’ as well as, for the first time, the artificial successive complex stimuli ‘ba’, which constitutes the Greek word ‘baba’ corresponding to the English ‘daddy’. Patients and methodsTwenty young adults institutionally diagnosed as dyslexic (10 subjects) or light dyslexic (10 subjects) comprised the diseased group. Twenty sex-, age-, education-, hearing sensitivity-, and IQ-matched normal subjects comprised the control group. Measurements included the absolute latencies of waves I through V, the interpeak latencies elicited by the classical acoustic click, the negative peak latencies of A and C waves, as well as the interpeak latencies of A–C elicited by the verbal stimulus ‘baba’ created on a digital speech synthesizer. ResultsThe absolute peak latencies of waves I, III, and V in response to monoaural rarefaction clicks as well as the interpeak latencies I–III, III–V, and I–V in the dyslexic subjects, although increased in comparison with normal subjects, did not reach the level of a significant difference (p<0.05). However, the absolute peak latencies of the negative wave C and the interpeak latencies of A–C elicited by verbal stimuli were found to be increased in the dyslexic group in comparison with the control group (p=0.0004 and p=0.045, respectively). In the subgroup consisting of 10 patients suffering from ‘other learning disabilities’ and who were characterized as with ‘light’ dyslexia according to dyslexia tests, no significant delays were found in peak latencies A and C and interpeak latencies A–C in comparison with the control group. ConclusionsAcoustic representation of a speech sound and, in particular, the disyllabic word ‘baba’ was found to be abnormal, as low as the auditory brainstem. Because ABRs mature in early life, this can help to identify subjects with acoustically based learning problems and apply early intervention, rehabilitation, and treatment. Further studies and more experience with more patients and pathological conditions such as plasticity of the auditory system, cochlear implants, hearing aids, presbycusis, or acoustic neuropathy are necessary until this type of testing is ready for clinical application.

Neuroethology in the service of neurophonetics

Abstract Single-neuron recording methods, as commonly used in neuroethology studies, provide the needed spatial and temporal resolution capacities to generate explicit hypotheses addressing the ‘how’ of language processing. The goal of this article is to describe two well documented neural processing mechanisms that can provide insights into (1) the auditory decoding of speech sounds, and (2) disambiguation of context-induced variability in stop place perception. The neural unit underlying speech sound processing is the combination-sensitive neuron, and the neural entity best suited to resolve context-induced variability in the speech signal is the neural column. The ‘absorption’ of stimulus variability via signal-specific columnar encoding is contrasted to exemplar-based treatments of stimulus variability in neural systems.

Synchrony capture filterbank: Auditory-inspired signal processing for tracking individual frequency components in speech

A processing scheme for speech signals is proposed that emulates synchrony capture in the auditory nerve. The role of stimulus-locked spike timing is important for representation of stimulus periodicity, low frequency spectrum, and spatial location. In synchrony capture, dominant single frequency components in each frequency region impress their time structures on temporal firing patterns of auditory nerve fibers with nearby characteristic frequencies (CFs). At low frequencies, for voiced sounds, synchrony capture divides the nerve into discrete CF territories associated with individual harmonics. An adaptive, synchrony capture filterbank (SCFB) consisting of a fixed array of traditional, passive linear (gammatone) filters cascaded with a bank of adaptively tunable, bandpass filter triplets is proposed. Differences in triplet output envelopes steer triplet center frequencies via voltage controlled oscillators (VCOs). The SCFB exhibits some cochlea-like responses, such as two-tone suppression and distortion products, and possesses many desirable properties for processing speech, music, and natural sounds. Strong signal components dominate relatively greater numbers of filter channels, thereby yielding robust encodings of relative component intensities. The VCOs precisely lock onto harmonics most important for formant tracking, pitch perception, and sound separation.

Listening to an Audio Drama Activates Two Processing Networks, One for All Sounds, Another Exclusively for Speech

Earlier studies have shown considerable intersubject synchronization of brain activity when subjects watch the same movie or listen to the same story. Here we investigated the across-subjects similarity of brain responses to speech and non-speech sounds in a continuous audio drama designed for blind people. Thirteen healthy adults listened for ∼19 min to the audio drama while their brain activity was measured with 3 T functional magnetic resonance imaging (fMRI). An intersubject-correlation (ISC) map, computed across the whole experiment to assess the stimulus-driven extrinsic brain network, indicated statistically significant ISC in temporal, frontal and parietal cortices, cingulate cortex, and amygdala. Group-level independent component (IC) analysis was used to parcel out the brain signals into functionally coupled networks, and the dependence of the ICs on external stimuli was tested by comparing them with the ISC map. This procedure revealed four extrinsic ICs of which two–covering non-overlapping areas of the auditory cortex–were modulated by both speech and non-speech sounds. The two other extrinsic ICs, one left-hemisphere-lateralized and the other right-hemisphere-lateralized, were speech-related and comprised the superior and middle temporal gyri, temporal poles, and the left angular and inferior orbital gyri. In areas of low ISC four ICs that were defined intrinsic fluctuated similarly as the time-courses of either the speech-sound-related or all-sounds-related extrinsic ICs. These ICs included the superior temporal gyrus, the anterior insula, and the frontal, parietal and midline occipital cortices. Taken together, substantial intersubject synchronization of cortical activity was observed in subjects listening to an audio drama, with results suggesting that speech is processed in two separate networks, one dedicated to the processing of speech sounds and the other to both speech and non-speech sounds.

Sensorimotor integration during human self-vocalization: Insights from invasive electrophysiology

Effective human speech requires the neural integration of ongoing vocal production with the auditory and somatosensory feedback signals that are produced. We are using invasive electrophysiology techniques in patient volunteers undergoing neurosurgical treatment in order to gain insights into these mechanisms and underlying neural circuits. By using multi-contact electrode arrays chronically implanted over the perisylvian temporal lobe auditory cortex (e.g., area PLST) and the inferior frontal gyrus (IFG), we can examine local field potentials and frequency-specific responses from cortical areas important for both vocal production and speech sound processing. Our initial studies have found that during self-vocalization, focal areas within higher order auditory cortex on the superior temporal gyrus (STG) show response modulation compared to the responses of the same areas during passive listening. Manipulation of the auditory feedback that a speaker receives during vocalization (e.g., pitch-shifted or delayed auditory feedback) leads to further modulation of these PLST sites. Measures of functional connectivity including electrical stimulation tract tracing or phase-synchrony analysis demonstrate that portions of PLST are functionally connected to regions of the IFG. These findings support forward models for vocal control in which efference copies of premotor cortex activity modulate sub-regions of auditory cortex.