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Parrell, B.

Publications and source records attributed to Parrell, B..

At least 19 recordsLinked to original sources

Modeling framework disentangles cerebellar mechanisms in speech feedback control, revealing trade-offs reshaped by degeneration

A central challenge in systems neuroscience is understanding how computational mechanisms--including those implemented within a single brain region--interact to produce behavior. For example, prior work in the literature attributes many computational functions to the cerebellum, but these functions have been tested in isolation and it remains unclear how they jointly contribute to motor control. Here, we test several established hypotheses of cerebellar function: internal modeling, timing of movement dynamics, sensory-error processing, delay processing, and multimodal integration. We first formalize these functions as mechanistic parameters within a computational model of speech motor control. We then use this formalism to investigate the relative contribution of each function to the abnormal speech corrective response seen in adults with cerebellar degeneration during perturbed auditory feedback. We find the following functions explain most of the behavioral differences: internal modeling, timing of movement dynamics, and multimodal integration. We also show that the key mechanisms have a trade-off relationship, and that cerebellar degeneration modulates those trade-off strengths and boundaries. These results both elaborate the mechanistic function of the cerebellum in speech feedback control and, more broadly, demonstrate the promise of using this paradigm to simultaneously test competing theories of neural function underlying behavior.

neuroscience↗

Sparse Stimulus Generation Improves Reverse Correlation Efficiency and Interpretability

Reverse correlation is a widely-used and well-established method for probing latent perceptual representations in which subjects render subjective preference responses to ambiguous stimuli. Stimuli are purposefully designed to have no direct relationship with the target representation (e.g., they are randomly-generated), a property which makes each individual stimulus minimally informative toward reconstructing the target, and often difficult to interpret for subjects. As a result, a large number of stimulus-response pairs must be gathered from a given subject in order for reconstructions to be of sufficient quality, making the task fatiguing. Recent work has demonstrated that the number of trials needed can be substantially reduced using a compressive sensing framework that incorporates the assumption that the target representation can be sparsely represented in some basis into the reconstruction process. Here, we introduce an alternative method that incorporates the sparsity assumption directly into stimulus generation, which holds promise not only for improving efficiency, but also for improving the interpretability of stimuli from subjects perspective. We develop this new method as a mathematical variation of the compressive sensing approach, before conducting one simulation study and two human subjects experiments to assess the benefits of this method to reconstruction quality, sample size efficiency, and subjective interpretability. Results show that sparse stimulus generation improves all three of these areas relative to conventional reverse correlation approaches, and also relative to compressive sensing in most conditions.

neuroscience↗

Sensorimotor adaptation of vocal pitch is severely impaired in cerebellar ataxia

Sensory errors, mismatches between predicted sensory outcomes of movement and reafferent sensory feedback, drive changes in the feedforward control of future motor behavior that correct for those errors. Across a wide variety of motor behaviors, individuals with cerebellar damage show impairments in these corrections, strongly suggesting a key role of the cerebellum in sensorimotor adaptation. However, the extent to which the cerebellum is involved in controlling vocal pitch is currently unknown. Crucially, vocal pitch differs in several ways from other systems that suggest it relies more on feedback than feedforward control. Adaptation itself also differs in vocal pitch: rather than the gradual build-up/decay of learning seen in other systems, pitch adaptation and de-adaptation are almost immediate. Together, this questions whether adaptation in vocal pitch relies on the same mechanism as other motor domains. Here, we test the hypothesis that the cerebellum underlies sensorimotor adaptation in vocal pitch, testing the domain-generality of this neurocomputational process. In both sustained vocalization and a more natural word production task, individuals with cerebellar ataxia fail to adapt to external auditory perturbation of vocal pitch. The complete lack of adaptation observed, compared to the impaired but present adaptation seen in other systems, suggest that the cerebellum plays an especially critical role in maintaining accurate control of vocal pitch. Conversely, we failed to detect a previously observed increase in online compensation to vocal pitch errors in ataxia, potentially suggesting this may be an idiosyncratic change in control rather than a common trait in this population. Significance statementWhen exposed to external perturbations of vocal pitch, individuals with cerebellar ataxia fail to adapt their produced pitch to oppose the perturbation. These results highlight the critical role of the cerebellum in driving adaptation across motor domains, even for motor behaviors that are thought to rely more on feedback compared to feedforward control, such as pitch. Conversely, and despite a large cohort, we failed to replicate previous findings of enhanced online corrections for pitch perturbations in ataxia, suggesting that any such increases are likely due to individual-specific changes in motor function in response to cerebellar-induced control deficits rather than being directly related to cerebellar damage itself.

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Preserved cerebellar functions despite structural degeneration in older adults

Aging is frequently perceived negatively due to its association with the decline of various brain and bodily functions. While it is evident that motor abilities deteriorate with age, it is incorrect to assume that all aspects of movement execution are equally affected. The cerebellum, a brain region that is closely involved in motor control among other functions, undergoes clear structural changes with aging. While several studies suggest that cerebellar degeneration causes age-related motor control deficits, other studies suggest that the cerebellum might act as a motor reserve and compensate for its structural degeneration, leaving cerebellar motor function intact despite cerebellar degeneration. The present study aims at thoroughly investigating the impact of age on cerebellar function across an array of tasks and domains. We investigated cerebellar motor and cognitive functions across the lifespan by examining 50 young adults (20-35 years), 80 older adults (55-70 years), and 30 older-old adults (>80 years). Participants completed a test battery comprising seven motor control tasks and one cognitive task, each designed to probe cerebellar function through different paradigms. This multi-task approach allowed for a comprehensive evaluation of performance patterns, providing a balanced perspective on cerebellar function across the different age groups. In addition, we analyzed outcomes from the same tasks that, while related to movement, were not specifically linked to cerebellar function. Structural magnetic resonance imaging was also conducted to assess whether cerebellar atrophy was present in the older and older-old groups compared to the young. Our results revealed that, despite age-related cerebellar degeneration, cerebellar functions in older adults remained intact compared to young adults, even in adults above 80 years old. In contrast, the sensorimotor measures that were not directly linked to cerebellar function exhibited a clear pattern of decline in older adults, and were further deteriorated in the older-old adults compared to the older adults. These findings indicate that cerebellar motor control functions remain largely preserved with age, providing compelling evidence that the cerebellum possesses a remarkable degree of functional resilience and redundancy. This suggests that cerebellar circuits may be uniquely equipped to preserve function despite structural degeneration.

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Subject Instructions for Improved Characterization of Auditory Representations Using Reverse Correlation

Reverse Correlation (RC) is an established method for reconstructing auditory representations, that has recently emerged as a tool for characterizing the sounds experienced by tinnitus patients. Toward further optimizing RC for auditory research, the present work investigated the influence of subject instructions on characterization quality of tinnitus-like sounds. A validation study was conducted in which eighteen normal-hearing subjects were randomly assigned one of three candidate instruction sets, each inspired by the RC literature. Results show a significant effect of instruction set on characterization quality, and reveal that instructing subjects to detect a hidden signal in the RC stimuli resulted the best reconstruction.

bioengineering↗

Audiomotor prediction errors drive speech adaptation even in the absence of overt movement

Observed outcomes of our movements sometimes differ from our expectations. These sensory prediction errors recalibrate the brains internal models for motor control, reflected in alterations to subsequent movements that counteract these errors (motor adaptation). While leading theories suggest that all forms of motor adaptation are driven by learning from sensory prediction errors, dominant models of speech adaptation argue that adaptation results from integrating time-advanced copies of corrective feedback commands into feedforward motor programs. Here, we tested these competing theories of speech adaptation by inducing planned, but not executed, speech. Human speakers were prompted to speak a word and, on a subset of trials, were rapidly cued to withhold the prompted speech. On standard trials, speakers were exposed to real-time playback of their own speech with an auditory perturbation of the first formant to induce single-trial speech adaptation. Speakers experienced a similar sensory error on movement cancellation trials, hearing a perturbation applied to a recording of their speech from a previous trial at the time they would have spoken. Speakers adapted to auditory prediction errors in both contexts, altering the spectral content of spoken vowels to counteract formant perturbations even when no actual movement coincided with the perturbed feedback. Such adaptation was not observed when participants passively listened to perturbed feedback without the intention to speak, ruling out observational learning as the cause of adaptation in movement cancellation trials. These results build upon recent findings in reaching, and suggest that prediction errors, rather than corrective motor commands, drive audiomotor adaptation in speech.

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Increased vowel contrast and intelligibility in connected speech induced by sensorimotor adaptation

Alterations to sensory feedback can drive robust adaptive changes to the production of consonants and vowels, but these changes often have no behavioral relevance or benefit to communication (e.g., making "head" more like "had"). This work aims to align the outcomes of adaptation with changes known to increase speech intelligibility - specifically, adaptations that increase the acoustic contrast between vowels in running speech. To this end, we implemented a vowel centralization feedback perturbation paradigm that pushes all vowels towards the center of vowel space, making them sound less distinct from one another. Speakers across the adult lifespan adapted to the centralization perturbation during sentence production, increasing the global acoustic contrast among vowels and the articulatory excursions for individual vowels. These changes persisted after the perturbation was removed, including after a silent delay, and showed robust transfer to words that were not present in the sentences. Control analyses demonstrated that these effects were unlikely to be due to explicit pronunciation strategies and occurred in the face of increasingly more rapid and less distinct production of familiar sentences. Finally, sentence transcription by crowd-sourced listeners showed that speakers vowel contrast predicted their baseline intelligibility and that experimentally-induced increases in contrast predicted intelligibility gains. These findings establish the validity of a sensorimotor adaptation paradigm to implicitly increase vowel contrast and intelligibility in connected speech, an outcome that has the potential to enhance rehabilitation in individuals who present with a reduced vowel space due to motor speech disorders, such as the hypokinetic dysarthria associated with Parkinsons disease. Significance statementDuring speech production, humans learn from perceived errors by adjusting their speech to compensate, a process known as sensorimotor adaptation. Here we leveraged sensorimotor adaptation to cause an increase in the distinctiveness of speech sounds. Specifically, speakers rapidly and implicitly learned vowel-specific articulation changes during sentence production, with positive effects on their intelligibility. This study is important for models of sensorimotor integration across all motor domains, as it demonstrates the simultaneous learning of multiple opposing transformations in the course of continuous movement. Our findings show that sensorimotor adaptation can be successfully applied to drive changes relevant to a vital ecological behavior: intelligible communication.

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Theta-burst stimulation over primary somatosensory cortex modulates tactile acuity of tongue

BackgroundEmerging studies in humans have established the modulatory effects of repetitive transcranial magnetic stimulation (rTMS) over primary somatosensory cortex (S1) on somatosensory cortex activity and perception. However, to date, research in this area has primarily focused on the hand and fingers, leaving a gap in our understanding of the modulatory effects of rTMS on somatosensory perception of the orofacial system and speech articulators. ObjectiveThe present study aimed to examine the effects of different types of theta-burst stimulation--continuous TBS (cTBS), intermittent TBS (iTBS), or sham--over the tongue representation of left S1 on tactile acuity of the tongue. MethodsIn a repeated-measures design, fifteen volunteers participated in four separate sessions, where cTBS, iTBS, sham, or no stimulation was applied over the tongue representation of left S1. Effects of TBS were measured on both temporal and spatial perceptual acuity of tongue using a custom vibrotactile stimulator. ResultsCTBS significantly impaired spatial amplitude threshold at the time window of 16-30 minutes after stimulation, while iTBS improved it at the same time window. The effect of iTBS, however, was smaller than cTBS. In contrast, neither cTBS nor iTBS had any effect on the temporal discrimination threshold. ConclusionsThe current study establishes the validity of using TBS to modulate somatosensory perception of the orofacial system. Directly modifying somatosensation in the orofacial system has the potential to benefit clinical populations with abnormal tactile acuity, improve our understanding of the role of sensory systems in speech production, and enhance speech motor learning and rehabilitation. HighlightsO_LITheta-burst stimulation (TBS) can modulate somatosensation in the orofacial system C_LIO_LIcTBS over S1 impaired spatial acuity of tongue C_LIO_LIiTBS over S1 improved spatial acuity of tongue C_LI

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Bayesian inference of state feedback control parameters for fo perturbation responses in cerebellar ataxia

Behavioral speech tasks have been widely used to understand the mechanisms of speech motor control in healthy speakers as well as in various clinical populations. However, determining which neural functions differ between healthy speakers and clinical populations based on behavioral data alone is difficult because multiple mechanisms may lead to the same behavioral differences. For example, individuals with cerebellar ataxia (CA) produce abnormally large compensatory responses to pitch perturbations in their auditory feedback, compared to controls, but this pattern could have many explanations. Here, computational modeling techniques were used to address this challenge. Bayesian inference was used to fit a state feedback control (SFC) model of voice fundamental frequency (fo) control to the behavioral pitch perturbation responses of individuals with CA and healthy controls. This fitting process resulted in estimates of posterior likelihood distributions of five model parameters (sensory feedback delays, absolute and relative levels of auditory and somatosensory feedback noise, and controller gain), which were compared between the two groups. Results suggest that the CA group may proportionally weight auditory and somatosensory feedback differently from the control group. Specifically, the CA group showed a greater relative sensitivity to auditory feedback than the control group. There were also large group differences in the controller gain parameter, suggesting increased motor output responses to target errors in the CA group. These modeling results generate hypotheses about how CA may affect the speech motor system, which could help guide future empirical investigations in CA. This study also demonstrates the overall proof-of-principle of using this Bayesian inference approach to understand behavioral speech data in terms of interpretable parameters of speech motor control models. Author summaryCerebellar ataxia is a condition characterized by a loss of coordination in the control of muscle movements, including those required for speech, due to damage in the cerebellar region of the brain. Behavioral speech experiments have been used to understand this disorders impact on speech motor control, but the results can be ambiguous to interpret. In this study, we fit a computational model of the neural speech motor control system to the speech data of individuals with cerebellar ataxia and that of healthy controls to determine what differences in model parameters best explain how the two groups differ in their control of vocal pitch. We found that group differences may be explained by increased sensitivity to auditory feedback prediction errors (differences between the actual sound speakers hear of their own speech as they produce it and the sound they expected to hear) and increased motor response in individuals with cerebellar ataxia. These computational results help us understand how cerebellar ataxia impacts speech motor control, and this general approach can also be applied to study other neurological speech disorders.

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Reverse Correlation Characterizes More Complete Tinnitus Spectra in Patients

GoalWe validate a recent reverse correlation approach to tinnitus characterization by applying it to individuals with clinically-diagnosed tinnitus. MethodsTwo tinnitus patients assessed the subjective similarity of their non-tonal tinnitus percepts and random auditory stimuli. Regression of the responses onto the stimuli yielded reconstructions which were evaluated qualitatively by playing back resynthesized waveforms to the subjects and quantitatively by response prediction analysis. ResultsSubject 1 preferred their resynthesis to white noise; subject 2 did not. Response prediction balanced accuracies were significantly higher than chance across subjects: subject 1: 0.5963, subject 2: 0.6922. ConclusionReverse correlation can provide the foundation for reconstructing accurate representations of complex, non-tonal tinnitus in clinically diagnosed subjects. Further refinements may yield highly similar waveforms to individualized tinnitus percepts. Impact StatementCharacterization of tinnitus sounds can help clarify the heterogeneous nature of the condition and link etiology to subtypes and treatments.

bioengineering↗

The brain's sensitivity to sensory error can be modulated by altering perceived variability

When individuals make a movement that produces an unexpected outcome, they learn from the resulting error. This process, essential in both acquiring new motor skills and adapting to changing environments, critically relies on error sensitivity, which governs how much behavioral change results from a given error. Although behavioral and computational evidence suggests error sensitivity can change in response to task demands, neural evidence regarding the flexibility of error sensitivity in the human brain is lacking. Critically, the sensitivity of the nervous system to auditory errors during speech production, a complex and well-practiced motor behavior, has been extensively studied by examining the prediction-driven suppression of auditory cortical activity. Here, we tested whether the nervous systems sensitivity to errors, as measured by this suppression, can be modulated by altering speakers perceived variability. Our results showed that error sensitivity was increased after exposure to an auditory perturbation that increased participants perceived variability, consistent with predictions generated from previous behavioral data and state-space modeling. Conversely, we observed no significant changes in error sensitivity when perceived variability was unaltered or artificially reduced. The current study establishes the validity of behaviorally modulating the nervous systems sensitivity to errors. As sensitivity to sensory errors plays a critical role in sensorimotor adaptation, modifying error sensitivity has the potential to enhance motor learning and rehabilitation in speech and, potentially, more broadly across motor domains. Significance StatementThe process of learning from error is essential for both the acquisition of new skills and successful adaptation to changing environments. Such error-based learning critically relies on error sensitivity, which determines how much we learn from a given error. Although evidence from behavioral studies suggests error sensitivity is malleable, neural evidence regarding the flexibility of error sensitivity in the human brain is lacking. Here, we showed that the nervous systems sensitivity to errors can be modulated by altering perceived variability. The present study establishes the validity of behaviorally modulating neural sensitivity to sensory errors. Improving our ability to learn from error can play a critical role in applied settings such as rehabilitation.

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Reverse Correlation Uncovers More Complete Tinnitus Spectra

GoalThis study validates an approach to characterizing the sounds experienced by tinnitus patients via reverse correlation, with potential for characterizing a wider range of sounds than currently possible. MethodsTen normal-hearing subjects assessed the subjective similarity of random auditory stimuli and target tinnitus-like sounds ("buzzing" and "roaring"). Reconstructions of the targets were obtained by regressing subject responses on the stimuli, and were compared for accuracy to the frequency spectra of the targets using Pearsons r. ResultsReconstruction accuracy was significantly higher than chance across subjects: buzzing (M = 0.53, SD = 0.27): t(9) = 5.766, p < 0.001; roaring (M = 0.57, SD = 0.30): t(9) = 5.76, p < 0.001. ConclusionReverse correlation can accurately reconstruct nontonal tinnitus-like sounds in normal-hearing subjects, indicating its potential for characterizing the sounds experienced by patients with non-tonal tinnitus. Impact StatementCharacterization of tinnitus sounds can inform treatment by facilitating individualized sound therapies, leading to better outcomes for patients suffering from the cognitive and psychological effects of tinnitus.

bioengineering↗

Stimulus Whitening Improves the Efficiency of Reverse Correlation

Human perception depends upon internal representations of the environment that help to organize the raw information available from the senses by acting as reference patterns. Internal representations are widely characterized using reverse correlation, a method capable of producing unconstrained estimates of the representation itself, all on the basis of simple responses to random stimuli. Despite its advantages, reverse correlation is often infeasible to apply because the number of stimulus-response trials needed to provide an accurate estimate is typically very large. Prior approaches have aimed to overcome this sampling inefficiency by incorporating prior knowledge of the representation, which biases the estimate and ultimately limits the essential power of reverse correlation. The present approach, however, improves efficiency via stimulus whitening, a statistical procedure that decorrelates stimuli, making them less redundant, and commensurately more favorable for efficient estimation of an arbitrary target. We provide a mathematical justification for whitening, and demonstrate in simulation that whitening provides greater than 85% improvement in efficiency for a given estimation accuracy, and also a two- to five-fold increase in accuracy for a given sample size. Improving the efficiency of reverse correlation may enable a broader scope of investigations into individual variability and potential universality of perceptual mechanisms.

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Variability is actively regulated in speech

Although movement variability is often attributed to unwanted noise in the motor system, recent work has demonstrated that variability may be actively controlled. To date, research on regulation of motor variability has relied on relatively simple, laboratory-specific reaching tasks. It is not clear how these results translate to complex, well-practiced and real-world tasks. Here, we test how variability is regulated during speech production, a complex, highly over-practiced and natural motor behavior that relies on auditory and somatosensory feedback. Specifically, in a series of four experiments, we assessed the effects of auditory feedback manipulations that modulate perceived speech variability, shifting every production either towards (inward-pushing) or away from (outward-pushing) the center of the distribution for each vowel. Participants exposed to the inward-pushing perturbation (Experiment 1) increased produced variability while the perturbation was applied as well as after it was removed. Unexpectedly, the outward-pushing perturbation (Experiment 2) also increased produced variability during exposure, but variability returned to near baseline levels when the perturbation was removed. Outward-pushing perturbations failed to reduce participants produced variability both with larger perturbation magnitude (Experiment 3) or after their variability had increased above baseline levels as a result of the inward-pushing perturbation (Experiment 4). Simulations of the applied perturbations using a state space model of motor behavior suggest that the increases in produced variability in response to the two types of perturbations may arise through distinct mechanisms: an increase in controlled variability in response to the inward-pushing perturbation, and an increase in sensitivity to auditory errors in response to the outward-pushing perturbation. Together, these results suggest that motor variability is actively regulated even in complex and well-practiced behaviors, such as speech.

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A Compressive Sensing Approach to Inferring Cognitive Representations with Reverse Correlation

Uncovering high-level cognitive representations of categories such as faces is an elusive goal that has been frequently sought using reverse correlation, a technique employed in fields ranging from neurophysiology to cognitive psychology. In reverse correlation, subjects are asked to make perceptual judgments (e.g., "do you see a face?") about richly varying stimuli (e.g., white noise), and observed responses are then regressed against stimuli to yield reconstructions of the underlying cognitive representation. However, many thousands of stimulus-response pairs are frequently required, which severely limits the breadth of studies that are feasible using this powerful method. Techniques that are currently employed to improve efficiency, such as filtering the reconstruction, nevertheless bias the outcome. Here, we show that an advanced signal processing technique for improving sampling efficiency - compressive sensing - is directly compatible with reverse correlation. A trio of simulations are performed to demonstrate that compressive sensing can reduce the required stimulus-response pairs by up to 90% without biasing the reconstruction or can retrospectively improve the accuracy of the reconstructions on existing data. This work concludes by outlining the potential of compressive sensing to improve representation reconstruction throughout the field of neuroscience and beyond. Significance StatementUncovering cognitive representations is an elusive goal that is increasingly pursued using the reverse correlation method, wherein human subjects make judgments about vague stimuli. Employing reverse correlation often entails collecting thousands of stimulus-response pairs, which severely limits the breadth of studies that are feasible using the method. Here we show that this methodological barrier can be overcome using compressive sensing, an advanced signal processing technique designed to improve sampling efficiency. Three sets of simulations are performed to demonstrate that compressive sensing can improve the accuracy of reconstructed cognitive representations and dramatically reduce the required number of stimulus-response pairs. This work concludes by outlining the potential of compressive sensing to improve representation reconstruction throughout the field of neuroscience and beyond.

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A single exposure to altered auditory feedback causes observable sensorimotor adaptation in speech

Sensory errors caused by perturbations to movement-related feedback induce two types of behavioral changes that oppose the perturbation: rapid compensation within a movement, as well as longer-term adaptation of subsequent movements. Although adaptation is hypothesized to occur whenever a sensory error is perceived (including after a single exposure to altered feedback), adaptation of articulatory movements in speech has only been observed after repetitive exposure to auditory perturbations, questioning both current theories of speech sensorimotor adaptation as well as the universality of more general theories of adaptation. Thus, positive evidence for the hypothesized single-exposure or "one-shot" learning would provide critical support for current theories of speech sensorimotor learning and control and align adaptation in speech more closely with other motor domains. We measured one-shot learning in a large dataset in which participants were exposed to intermittent, unpredictable auditory perturbations to their vowel formants (the resonant frequencies of the vocal tract that distinguish between different vowels). On each trial, participants spoke a word out loud while their first formant was shifted up, shifted down, or remained unshifted. We examined whether the perturbation on a given trial affected speech on the subsequent, unperturbed trial. We found that participants adjusted their first formant in the opposite direction of the preceding shift, demonstrating that learning occurs even after a single auditory perturbation as predicted by current theories of sensorimotor adaptation. While adaptation and the preceding compensation responses were correlated, this was largely due to differences across individuals rather than within-participant variation from trial to trial. These findings are more consistent with theories that hypothesize adaptation is driven directly by updates to internal control models than those that suggest adaptation results from incorporation of feedback responses from previous productions.

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Differential effects of cerebellar degeneration on feedforward versus feedback control across speech and reaching movements

Errors that result from a mismatch between predicted movement outcomes and sensory afference are used to correct ongoing movements through feedback control and to adapt feedforward control of future movements. The cerebellum has been identified as a critical part of the neural circuit underlying implicit adaptation across a wide variety of movements (reaching, gait, eye movements, and speech). The contribution of this structure to feedback control is less well understood: although it has recently been shown in the speech domain that individuals with cerebellar degeneration produce even larger online corrections for sensory perturbations than control participants, similar behavior has not been observed in other motor domains. Currently, comparisons across domains are limited by different population samples and potential ceiling effects in existing tasks. To assess the relationship between changes in feedforward and feedback control associated with cerebellar degeneration across motor domains, we evaluated adaptive (feedforward) and compensatory (feedback) responses to sensory perturbations in reaching and speech production in human participants of both sexes with cerebellar degeneration and neurobiologically healthy controls. As expected, the cerebellar group demonstrated impaired adaptation in both reaching and speech. In contrast, the groups did not differ in their compensatory response in either domain. Moreover, compensatory and adaptive responses in the cerebellar group were not correlated within or across motor domains. These results point to a general impairment in feedforward control with spared feedback control in cerebellar degeneration. However, the magnitude of feedforward impairments and potential changes in feedback-based control manifest in a domain-specific manner across individuals. Significance StatementThe cerebellum contributes to feedforward updating of movement in response to sensory errors, but its role in feedback control is less understood. Here, we tested individuals with cerebellar degeneration (CD), using sensory perturbations to assess adaptation of feedforward control and feedback gains during reaching and speech production tasks. The results confirmed that CD leads to reduced adaption in both domains. However, feedback gains were unaffected by CD in either domain. Interestingly, measures of feedforward and feedback control were not correlated across individuals within or across motor domains. Together, these results indicate a general impairment in feedforward control with spared feedback control in CD. However, the magnitude of feedforward impairments manifests in a domain-specific manner across individuals.

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Speech motor behavior can be altered through reinforcement learning

Reinforcement learning, the ability to change motor behavior based on external reward, has been suggested to play a critical role in early stages of speech motor development and is widely used in clinical rehabilitation for speech motor disorders. However, no current evidence exists that demonstrates the capability of reinforcement to drive changes in human speech behavior. Speech provides a unique test of the universality of reinforcement learning across motor domains: speech is a complex, high-dimensional motor task whose goals do not specify a task to be performed in the environment but ultimately must be self-generated by each speaker such that they are understood by those around them. Across four experiments, we examine whether reinforcement learning alone is sufficient to drive changes in speech behavior and parametrically test two features known to affect reinforcement learning in reaching: how informative the reinforcement signal is as well as the availability of sensory feedback about the outcomes of ones motor behavior. We show that learning does occur and is more likely when participants receive auditory feedback that gives an implicit target for production, even though they do not explicitly imitate that target. Contrary to results from upper limb control, masking feedback about movement outcomes has no effect on speech learning. Together, our results suggest a potential role for reinforcement learning in speech but that it likely operates differently than in other motor domains.

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