Kevin J. Reilly

Neural mechanisms underlying auditory feedback control of speech

The neural substrates underlying auditory feedback control of speech were investigated using a combination of functional magnetic resonance imaging (fMRI) and computational modeling. Neural responses were measured while subjects spoke monosyllabic words under two conditions: (i) normal auditory feedback of their speech and (ii) auditory feedback in which the first formant frequency of their speech was unexpectedly shifted in real time. Acoustic measurements showed compensation to the shift within approximately 136 ms of onset. Neuroimaging revealed increased activity in bilateral superior temporal cortex during shifted feedback, indicative of neurons coding mismatches between expected and actual auditory signals, as well as right prefrontal and Rolandic cortical activity. Structural equation modeling revealed increased influence of bilateral auditory cortical areas on right frontal areas during shifted speech, indicating that projections from auditory error cells in posterior superior temporal cortex to motor correction cells in right frontal cortex mediate auditory feedback control of speech.

Speech serial control in healthy speakers and speakers with hypokinetic or ataxic dysarthria: effects of sequence length and practice

The current study investigated the processes responsible for selection of sounds and syllables during production of speech sequences in 10 adults with hypokinetic dysarthria from Parkinson’s disease, five adults with ataxic dysarthria, and 14 healthy control speakers. Speech production data from a choice reaction time task were analyzed to evaluate the effects of sequence length and practice on speech sound sequencing. Speakers produced sequences that were between one and five syllables in length over five experimental runs of 60 trials each. In contrast to the healthy speakers, speakers with hypokinetic dysarthria demonstrated exaggerated sequence length effects for both inter-syllable intervals (ISIs) and speech error rates. Conversely, speakers with ataxic dysarthria failed to demonstrate a sequence length effect on ISIs and were also the only group that did not exhibit practice-related changes in ISIs and speech error rates over the five experimental runs. The exaggerated sequence length effects in the hypokinetic speakers with Parkinson’s disease are consistent with an impairment of action selection during speech sequence production. The absent length effects observed in the speakers with ataxic dysarthria is consistent with previous findings that indicate a limited capacity to buffer speech sequences in advance of their execution. In addition, the lack of practice effects in these speakers suggests that learning-related improvements in the production rate and accuracy of speech sequences involves processing by structures of the cerebellum. Together, the current findings inform models of serial control for speech in healthy speakers and support the notion that sequencing deficits contribute to speech symptoms in speakers with hypokinetic or ataxic dysarthria. In addition, these findings indicate that speech sequencing is differentially impaired in hypokinetic and ataxic dysarthria.

Sequence Complexity Effects on Speech Production in Healthy Speakers and Speakers with Hypokinetic or Ataxic Dysarthria

The present study investigated the effects of sequence complexity, defined in terms of phonemic similarity and phonotoactic probability, on the timing and accuracy of serial ordering for speech production in healthy speakers and speakers with either hypokinetic or ataxic dysarthria. Sequences were comprised of strings of consonant-vowel (CV) syllables with each syllable containing the same vowel, /a/, paired with a different consonant. High complexity sequences contained phonemically similar consonants, and sounds and syllables that had low phonotactic probabilities; low complexity sequences contained phonemically dissimilar consonants and high probability sounds and syllables. Sequence complexity effects were evaluated by analyzing speech error rates and within-syllable vowel and pause durations. This analysis revealed that speech error rates were significantly higher and speech duration measures were significantly longer during production of high complexity sequences than during production of low complexity sequences. Although speakers with dysarthria produced longer overall speech durations than healthy speakers, the effects of sequence complexity on error rates and speech durations were comparable across all groups. These findings indicate that the duration and accuracy of processes for selecting items in a speech sequence is influenced by their phonemic similarity and/or phonotactic probability. Moreover, this robust complexity effect is present even in speakers with damage to subcortical circuits involved in serial control for speech.

The Effects of Fluency Enhancing Conditions on Sensorimotor Control of Speech in Typically Fluent Speakers: An EEG Mu Rhythm Study

Objective: To determine whether changes in sensorimotor control resulting from speaking conditions that induce fluency in people who stutter (PWS) can be measured using electroencephalographic (EEG) mu rhythms in neurotypical speakers. Methods: Non-stuttering (NS) adults spoke in one control condition (solo speaking) and four experimental conditions (choral speech, delayed auditory feedback (DAF), prolonged speech and pseudostuttering). Independent component analysis (ICA) was used to identify sensorimotor μ components from EEG recordings. Time-frequency analyses measured μ-alpha (8–13 Hz) and μ-beta (15–25 Hz) event-related synchronization (ERS) and desynchronization (ERD) during each speech condition. Results: 19/24 participants contributed μ components. Relative to the control condition, the choral and DAF conditions elicited increases in μ-alpha ERD in the right hemisphere. In the pseudostuttering condition, increases in μ-beta ERD were observed in the left hemisphere. No differences were present between the prolonged speech and control conditions. Conclusions: Differences observed in the experimental conditions are thought to reflect sensorimotor control changes. Increases in right hemisphere μ-alpha ERD likely reflect increased reliance on auditory information, including auditory feedback, during the choral and DAF conditions. In the left hemisphere, increases in μ-beta ERD during pseudostuttering may have resulted from the different movement characteristics of this task compared with the solo speaking task. Relationships to findings in stuttering are discussed. Significance: Changes in sensorimotor control related feedforward and feedback control in fluency-enhancing speech manipulations can be measured using time-frequency decompositions of EEG μ rhythms in neurotypical speakers. This quiet, non-invasive, and temporally sensitive technique may be applied to learn more about normal sensorimotor control and fluency enhancement in PWS.

Vowel generalization and its relation to adaptation during perturbations of auditory feedback

Repeated perturbations of auditory feedback during vowel production elicit changes not only in the production of the perturbed vowel (adaptation) but also in the production of nearby vowels that were not perturbed (generalization). The finding that adaptation generalizes to other, nonperturbed vowels suggests that sensorimotor representations for vowels are not independent; instead, the goals for producing any one vowel may depend in part on the goals for other vowels. The present study investigated the dependence or independence of vowel representations by evaluating adaptation and generalization in two groups of speakers exposed to auditory perturbations of their first formant (F1) during different vowels. The speakers in both groups who adapted to the perturbation exhibited generalization in two nonperturbed vowels that were produced under masking noise. Correlation testing was performed to evaluate the relations between adaptation and generalization as well as between the generalization in the two nonperturbed vowels. These tests identified significant coupling between the F1 changes of adjacent vowels but not nonadjacent vowels. The pattern of correlation findings indicates that generalization was due in part to feedforward representations that are partly shared across adjacent vowels, possibly to maintain their acoustic contrast.NEW & NOTEWORTHY Speech adaptations to alterations, or perturbations, of auditory feedback have provided important insights into sensorimotor representations underlying speech. One finding from these studies that is yet to be accounted for is vowel generalization, which describes the effects of repeated perturbations to one vowel on the production of other vowels that were not perturbed. The present study used correlation testing to quantify the effects of changes in a perturbed vowel on neighboring (i.e., similar) nonperturbed vowels. The results identified significant correlations between the changes of adjacent, but not nonadjacent, vowel pairs. This finding suggests that generalization is partly a response to adaptation and not solely due to the auditory perturbation.

Trait related sensorimotor deficits in people who stutter: An EEG investigation of μ rhythm dynamics during spontaneous fluency

Stuttering is associated with compromised sensorimotor control (i.e., internal modeling) across the dorsal stream and oscillations of EEG mu (μ) rhythms have been proposed as reliable indices of anterior dorsal stream processing. The purpose of this study was to compare μ rhythm oscillatory activity between (PWS) and matched typically fluent speakers (TFS) during spontaneously fluent overt and covert speech production tasks. Independent component analysis identified bilateral μ components from 24/27 PWS and matched TFS that localized over premotor cortex. Time-frequency analysis of the left hemisphere μ clusters demonstrated significantly reduced μ-α and μ-β ERD (pCLUSTER < 0.05) in PWS across the time course of overt and covert speech production, while no group differences were found in the right hemisphere in any condition. Results were interpreted through the framework of State Feedback Control. They suggest that weak forward modeling and evaluation of sensory feedback across the time course of speech production characterizes the trait related sensorimotor impairment in PWS. This weakness is proposed to represent an underlying sensorimotor instability that may predispose the speech of PWS to breakdown.

EEG mu rhythms: Rich sources of sensorimotor information in speech processing

a Department of Audiology and Speech-Language Pathology, University of Tennessee Health Sciences, Knoxville, TN 37996, USA University of Arkansas, Epley Center for Health Professions, 606 N. Razorback Road, Fayetteville, AR 72701, USA c Department of Speech and Hearing Sciences, Elson S. Floyd College of Medicine, Spokane, WA 99210-1495, USA d Department of Hearing, Speech, and Language Sciences, Gallaudet University, 800 Florida Avenue NE, Washington, DC 20002, USA

Vowel and sibilant sound production in noise

The present study investigated speech production in noise and whether speakers modulate the spectral contrasts of vowel and sibilant sounds depending on the frequency characteristics of the background noise signal. Twelve speakers participated in the present study. Speakers were presented with three-word sequences one word at a time on a computer monitor and, after a variable delay, produced the three words in the order they were presented. Word sequences were comprised of CVC words containing either or both of the vowels, /a/ and /ae/, and either or both of the sibilants, /s/ and /ʃ/, in word-initial position. During the presentation and production of each sequence, speakers were exposed to one of four possible noise conditions: (1) silence; (2) vowel masking; (3) sibilant masking; and (4) speech-shaped noise. The sibilant masker contained energy at frequencies associated with a speaker’s productions of /s/ and /ʃ/ and the vowel masker at frequencies associated with their productions of /a/ and /ae/. The...

A neuroimaging investigation of auditory‐motor learning

Brain activity was measured in 11 subjects while learning a novel speech‐like auditory‐motor task. Subjects learned to move a dot on a screen to one of three target locations by changing their formant frequencies during production of vowel‐like sounds. The targets corresponded to regions of the formant space not associated with American English vowels. Functional images were acquired using a sparse sampling technique that allowed performance of the task in the absence of scanner noise and avoided artifactual signal changes resulting from articulator movements. The accuracy of subjects’ production of the novel vowel sounds increased significantly over the course of the scanning session. Significant correlations between performance error and brain activity were observed in a number of areas, including superior temporal gyrus, inferior frontal gyrus, primary motor cortex, supplementary motor area, and medial and lateral cerebellum. Brain regions exhibiting a positive correlation with performance error were g...