Guenther Lecture Part 2

Auditory, Somatosensory, and Motor Interactions in Speech Production
Frank H. Guenther
Department of Cognitive and Neural Systems, Boston University Division of Health Sciences and Technology, Harvard University / M.I.T. Research Laboratory of Electronics, Massachusetts Institute of Technology
Collaborators
Satrajit Ghosh Alfonso Nieto-Castanon Jason Tourville Oren Civier Kevin Reilly Jason Bohland Jonathan Brumberg Michelle Hampson Joseph Perkell Virgilio Villacorta Majid Zandipour Melanie Matthies Shinji Maeda
HST 722 Speech Motor Control
Supported by NIDCD, NSF. 1
CNS Speech Lab at Boston University

Primary goal is to elucidate the neural processes underlying: Learning of speech in children Normal speech in adults Breakdowns of speech in disorders such as stuttering and apraxia of speech Methods of investigation include: Neural network modeling Functional brain imaging Motor and auditory psychophysics These studies are organized around the DIVA model, a neural network model of speech acquisition and production developed in our lab.
HST 722 Speech Motor Control 2
Talk Outline
Overview of the DIVA model Mirror neurons in the model Learning in the model Simulating a hemodynamic response from the model
Feedback control subsystem Auditory perturbation fMRI experiment Somatosensory perturbation fMRI experiment
Feedforward control subsystem Sensorimotor adaptation to F1 perturbation

Summary
Schematic of the DIVA Model
Boxes in the schematic correspond to maps of neurons; arrows correspond to synaptic projections. The model controls movements of a virtual vocal tract, or articulatory synthesizer. Video shows random movements of the articulators in this synthesizer. Production of a speech sound in the model starts with activation of a speech sound map cell in left ventral premotor cortex (BA 44/6), which in turn activates feedforward and feedback control subsystems that converge on primary motor cortex.
Speech Sound Map Mirror Neurons

Since its inception in 1992, the DIVA model has included a speech sound map that contains cells which are active during both perception and production of a particular speech sound (phoneme or syllable). During perception, these neurons are necessary to learn an auditory target or goal for the sound, and to a lesser degree somatosensory targets (limited to the visible articulators such as lips).
Speech sound map during perception

Somatosensory Goal Region Speech Sound Map (Premotor Cortex) Auditory Goal Region Auditory State
Somatosensory State
Feedforward Command
Auditory Error (Auditory Cortex)
Articulator Velocity and Position Cells (Motor Cortex)
Auditory FeedbackBased Command
Somatosensory Error (Somatosensory Cortex)
Somatosensory FeedbackBased Command
To Muscles
Speech Sound Map Mirror Neurons

After a sound has been learned (described next), activating the speech sound map cells for the sound leads to readout of the learned feedforward commands (gestures) and auditory and somatosensory targets for the sound (red arrows at right). These targets are compared to incoming sensory signals to generate corrective commands if needed (blue). The overall motor command (purple) combines feedforward and feedback components.
Speech sound map during production

Somatosensory State
Feedforward Command
To Muscles
Learning in the Model Stage 1

In the first learning stage, the model learns the relationships between motor commands, somatosensory feedback, and auditory feedback. In particular, the model needs to learn how to transform sensory error signals into corrective motor commands.
Somatosensory State
Feedforward Command
This is done with babbling movements of the vocal tract which provide paired sensory and motor signals that can be used to tune these transformations.
To Muscles
Learning in the Model Stage 2 (Imitation)

The model then needs to learn auditory and somatosensory targets for individual speech sounds, and feedforward motor commands (gestures) for these sounds. This is done through an imitation process involving the speech sound map cells.
Model projections tuned during the imitation process are shown in red.
Somatosensory State
Feedforward Command
To Muscles
The Imitation Process

(1) The model learns an auditory target from a sound sample provided by a fluent speaker; this target is stored in the synaptic weights projecting from the speech sound map to the higher-order auditory cortical areas.
Auditory target for ba
(2) The model practices production of the sound to tune the feedforward commands and learn a somatosensory target.
Simulation Learning Feedforward Commands

The model first learns the auditory target for the sound by listening to someone produce it. Sound sample presented to model:
Then it tries to repeat the target, initially under auditory feedback control. With each repetition, the model relies less on feedback control and more on feedforward control, resulting in better and better productions.
Top panel: Spectrogram of target utterance presented to the model. Remaining panels: Spectrograms of the models first few attempts to produce the utterance.
Note improvement of auditory trajectories with each practice iteration due to improved feedforward commands.
12
Simulating a Hemodynamic Response in the Model

Each model component corresponds to a particular region of the brain:
Estimated Anatomical Locations of Model Components
Resp
Resp
Lip Jaw DS Tongue Palate* Tongue Palate* Larynx SSM Jaw
Lip
DS
DA Lat Cbm
Larynx Aud
DA Aud Lat Cbm
The anatomical locations of the models components have been finetuned by comparison to the results of previous neurophysiological and neuroimaging studies (Guenther, Ghosh, and Tourville, 2006).
The models cell activities during simulations can be directly compared to the results of fMRI and PET studies.
14
Talk Outline

Summary
15
Feedback Control Subsystem

The models feedback control subsystem compares learned auditory and somatosensory target regions for the current speech sound to incoming sensory information. If the current auditory or somatosensory state is outside the target region for the sound, error signals arise in higher-order auditory and/or somatosensory areas in the superior temporal lobe and parietal lobe.
Somatosensory State
Feedforward Command
To Muscles
16
Feedback Control Subsystem (continued)

Auditory and somatosensory error signals are then transformed into corrective motor commands via projections from the sensory areas to the motor cortex.
Somatosensory State
Feedforward Command
To Muscles
17
Prediction: Auditory Error Cells

The model hypothesizes an auditory error map in the higher-order auditory cortex of the posterior superior temporal gyrus and planum temporale. Cells in this map should become active if a subjects auditory feedback of his/her own speech is perturbed so that it mismatches the subjects auditory target.
The model also predicts that this auditory error cell activation will give rise to increased activity in motor areas, where corrective articulator commands are generated.
18
fMRI Study of Unexpected Auditory Perturbation During Speech

To test these predictions, we performed an fMRI study involving 11 subjects in which the first formant frequency (an important acoustic cue for speech) was unexpectedly perturbed upward or downward by 30% in of the production trials.
The perturbed feedback trials were randomly interspersed with normal feedback trials so the subject could not anticipate the perturbations. Perturbations were applied using a DSP device developed with colleagues at MIT (Villacorta, Perkell, Guenther, 2004) which feeds the modified speech signal back to the subject in near real-time (~16 ms delay, not noticeable to subject).
Sound before shift: Sound after shift:
19
Unexpectedly shifting the feedback caused subjects to compensate within the same syllable as the shift (gray 95% confidence intervals):
Response to downshift
Normalized F1
Response to upshift
Time (sec)
DIVA model productions in response to unexpected upward (dashed line) and downward (solid line) perturbations of F1 fall within the distribution of productions of the speakers in the fMRI study (shaded HST 722 Speech Motor Control 20 regions).
=> Auditory feedback control is right-lateralized in the frontal cortex.

Prediction: Somatosensory Error Cells

The model also predicts that a sudden, unexpected perturbation of the jaw should cause an increase in error cell activity in somatosensory and (perhaps) auditory cortical areas.
Somatosensory State
Feedforward Command
This in turn should lead to increased activity in motor areas where corrective commands are generated.
To Muscles
22
fMRI Study of Unexpected Jaw Perturbation During Speech

13 subjects produced /aCi/ utterances while in the MRI scanner (e.g., abi, ani, agi). An event-triggered paradigm was used to avoid movement artifacts and scanner noise issues:
On 1 in 7 utterances, a small balloon was rapidly inflated between the teeth during the initial vowel. The balloon inhibits upward jaw movement for the consonant and final vowel, causing the subject to compensate with larger tongue and/or lip movements.
23
Perturbed Unperturbed Speech (p<0.001):
L L
R R
24
Talk Outline

Summary
25
Feedforward Control in the Model

In addition to activating the feedback control subsystem, activating speech sound map cells also causes the readout of feedforward commands for the sound to be produced.
Feedforward commands in the DIVA model
These commands are encoded in synaptic projections from premotor cortex to primary motor cortex, including both corticocortical (blue) and transcerebellar (purple) projections.
Combining Feedforward and Feedback Commands

The commands generated by the feedforward system (red) and feedback system (blue) are combined in motor cortex to form the overall motor command to the speech articulators (purple)
Somatosensory State
Feedforward Command
To Muscles
27
Tuning Feedforward Commands

Early in development, the feedforward commands are poorly tuned, so the feedback control subsystem is needed to correct the commands.
On each attempt to produce a sound, the feedforward controller incorporates these feedback-based corrections into the feedforward command for the next attempt, resulting in better and better feedforward control with practice.
28
The interactions between the feedforward and feedback control subsystems in the model lead to the following predictions:
If a speakers auditory feedback is perturbed consistently over many consecutive productions of a syllable, corrective commands issued by the auditory feedback control subsystem will become incorporated into the feedforward commands for that syllable.
Speakers with better hearing (auditory acuity) will adapt more than speakers with worse hearing. If the perturbation is then removed, the speaker will show after-effects due to these adjustments to the feedforward command.
This was investigated by Villacorta, Perkell, & Guenther (2004).

Sensorimotor Adaptation Study F1 Perturbation

In each epoch of the adaptation study, the subject read a short list of words involving the vowel eh (e.g., bet, peck). After a baseline phase of 15 epochs of reading with normal feedback, a shift of F1 was gradually applied to the subjects auditory feedback during the next 5 epochs. The shift was then held at the maximum level (30% shift) for 25 epochs. Finally, feedback was returned to normal in a 20-epoch post-test phase. The entire experimental session lasted approximately 60-90 minutes.
Results for 20 subjects shown by lines with standard error bars. Shaded region is the 95% confidence interval for model simulation results (one simulation per speaker, target region size determined by speakers auditory acuity).
Sensorimotor Adaptation Study Results

Sustained auditory perturbation leads to adjustments in feedforward commands for speech in order to cancel out the effects of the perturbation. Amount of adaptation is correlated to speakers auditory acuity: high acuity speakers adapt more completely to the perturbation. When perturbation is removed, speech only gradually returns to normal values; i.e., there is an after-effect in the first few trials after hearing returns to normal (evidence for feedforward command adaptation).
The model provides a close quantitative fit to these processes.
32
Summary
The DIVA model elucidates several types of learning in speech acquisition, e.g.: Learning of relationships between articulations and their acoustic and somatosensory consequences Learning of auditory targets for speech sounds in the native language from externally presented examples Learning of feedforward commands for new sounds through practice The model elucidates the interactions between motor, somatosensory, and auditory areas responsible for speech motor control. The model spans behavioral and neural levels and makes predictions that are being tested using a variety of experimental techniques.
Reconciling Gestural and Auditory Views of Speech Production

In our view, the gestural score is the feedforward component of speech production, stored in the projections from premotor to motor cortex, which consists of an optimized set of motor programs for the most frequently produced phonemes, syllables and syllable strings (in Levelts terms, the syllabary). These gestural scores are shaped by auditory experience in order to adhere to acceptable auditory bounds of speech sounds in the native language(s). They are supplemented by auditory and somatosensory feedback control systems that constantly adjust the gestures when they detect errors in performance, e.g. due to growth of the vocal tract, addition of a false palate, or auditory feedback perturbation.
34
Collaborators
Alfonso NietoCastanon Satrajit Ghosh Jason Tourville Kevin Reilly Oren Civier Jonathan Brumberg Jason Bohland Michelle Hampson Joseph Perkell Majid Zandipour Virgilio Villacorta Melanie Matthies Shinji Maeda
Supported by NSF and NIDCD
35
Simulating a Hemodynamic Response from the Model

Model cell activities during simulations of speech are convolved with an idealized hemodynamic response function, generated using default settings of the function spm_hrf from the SPM toolbox. This function was designed to characterize the transformation from cell activity to hemodynamic response in the brain. A brain volume is then constructed with the appropriate hemodynamic response values at each position and smoothed with a Gaussian kernel (FWHM=12mm). This smoothing process approximates the smoothing carried out during standard SPM analysis of human subject fMRI data. The resultant volume is then rendered using routines from the SPM toolbox.
36
An event-triggered paradigm was used to avoid movement artifacts and scanner noise issues:
37
Hypothesis regarding onset of stuttering:
Threshold for motor reset due to sensory error signals
Error signal magnitude of stuttering individual
Error signal magnitude for non-stuttering individual
Onset of stuttering
Age
38
Brain regions active during cued production of 3syllable strings
PreSMA
Frame R ep Frame S ignals
SMA
T rigger Cells T rigger S ignals Next S ound
Overt speech only
IFS
S equence WM
BA 44
S peech S ound Map
T o m otor c ortex (DIVA m odel)
39
Auditory Target Regions

The models use of sensory target regions provides a unified account for a number of speech production phenomena, including aspects of: Economy of effort (cf. Lindblom)
Tongue Body Height
Carryover Coarticulation
Articulatory variability Anticipatory coarticulation Carryover coarticulation
Target for /k/
Speaking rate effects
Config. for /u/
Config. for /i/
Schematized at right is the models Tongue Body Horiz. Position explanation of carryover coarticulation and economy of effort during production of /k/ in luke and leak. HST 722 Speech Motor Control
40
Two factors that could influence target region size: (1) Perceptual acuity of speaker: better perceptual acuity => smaller regions (2) Speaking condition: clear speech (vs. fast speech) => smaller regions
Contrast distance /i/ Second formant frequency
/e/
First formant frequency
41
L L L L
Articulatory
LO
2 0
Results of EMMA studies:
Whod- Hood
Acoustic Contrast Distance (Hz)
400
Cod-Cud
H H
(1) Speakers with high perceptual acuity show greater contrast distance in production of neighboring sound categories.
(2) General tendency for greater contrast distance in clear speech, less in fast speech.
350
HI 300
H
HI LO
*
L L L
LO
250
H L L
200
F N C Speaking Condition
F N C Speaking Condition
Acoustic contrast distance (Hz)
Sod-shod 2000 1500 1000 500 0
Discrimination
Said-shed -
*
H L H L H L
H L
H L
H L L
H L
These results support the predictions on the preceding slide. Perkell et al. (2004a,b).
Speaking condition
Speaking condition
42
Ellipses indicating the range of formant frequencies (+/-1 s.d.) used by a speaker to produce five vowels (iy, eh, aa, uh, uw) during fast speech (light gray) and clear speech (dark gray) in a variety of phonetic contexts.
Motor Equivalence in American English /r/ Production

It has long been known that the American English phoneme /r/ is produced with a large amount of articulatory variability, both within a subject and between subjects. Delattre and Freeman (1968):
44
Despite large articulatory variability, the key acoustic cue for /r/ remains relatively stable across phonetic contexts. Boyce and Espy-Wilson (1997):
45
Motor Equivalence in the DIVA Model

Models use of auditory target for /r/ and directional mapping between auditory and articulatory spaces leads to different articulatory gestures, and different vocal tract shapes, for /r/ depending on phonetic context: Producing /r/ after /g/: Producing /r/ after /a/:
EMMA/Modeling Study: (1) Collect EMMA data from speakers producing /r/ in different contexts (2) Build speaker-specific vocal tract models (articulatory synthesizers) for two of the speakers (3) Train the DIVA model to produce sounds with the speaker-specific vocal tracts (4) Compare the models /r/ productions to those of the EMMA subjects
Sketch of hypothesized trading relations for /r/:

S4
Lips
S1 S5
Tongue constriction
Acoustic effect of the larger front cavity (blue) is compensated by the effect of the longer and narrower constriction (red).
1 cm
S2
S6
This yields similar acoustics for bunched (red) and retroflex (blue) tongue configurations for /r/ (Stevens, 1998; Boyce & Espy-Wilson, 1997).
All seven subjects in the EMMA study utilized similar trading relations (Guenther et al., 1999).
S3
S7
BACK
FRONT
BACK
FRONT
/wagrav/ /warav/ or /wabrav/
47
Building Speaker-Specific Vocal Tract Models from MRI Images
changing
F1
changing
F2
changing
F3
Subject 2
Subject 1
48
Comparison of models articulations using speaker-specific vocal tracts to those speakers actual articulations:
/ar/
subject data
Subject 1 /dr/
/gr/
/ar/
Subject 2 /dr/
/gr/
DIVA simulations
[Nieto-Castanon, Guenther, Perkell, and Curtin (2005), J Acoust Soc Am.]
Somatosensory State
Feedforward Command
To Muscles
50

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Auditory, Somatosensory, and Motor Interactions in Speech Production

HST 722 Speech Motor Control

Supported by NIDCD, NSF. 1

CNS Speech Lab at Boston University

Feedforward control subsystem Sensorimotor adaptation to F1 perturbation

HST 722 Speech Motor Control

Schematic of the DIVA Model

HST 722 Speech Motor Control

Speech Sound Map Mirror Neurons

Speech sound map during perception

Auditory Error (Auditory Cortex)

Articulator Velocity and Position Cells (Motor Cortex)

Auditory FeedbackBased Command

Somatosensory Error (Somatosensory Cortex)

Somatosensory FeedbackBased Command

HST 722 Speech Motor Control

Speech Sound Map Mirror Neurons

Speech sound map during production

Auditory Error (Auditory Cortex)

Articulator Velocity and Position Cells (Motor Cortex)

Auditory FeedbackBased Command

Somatosensory Error (Somatosensory Cortex)

Somatosensory FeedbackBased Command

HST 722 Speech Motor Control

Learning in the Model Stage 1

Auditory Error (Auditory Cortex)

Articulator Velocity and Position Cells (Motor Cortex)

Auditory FeedbackBased Command

Somatosensory Error (Somatosensory Cortex)

Somatosensory FeedbackBased Command

HST 722 Speech Motor Control

Learning in the Model Stage 2 (Imitation)

Auditory Error (Auditory Cortex)

Articulator Velocity and Position Cells (Motor Cortex)

Auditory FeedbackBased Command

Somatosensory Error (Somatosensory Cortex)

Somatosensory FeedbackBased Command

HST 722 Speech Motor Control

The Imitation Process

Auditory target for ba

Simulation Learning Feedforward Commands

HST 722 Speech Motor Control

Simulating a Hemodynamic Response in the Model

Lip Jaw DS Tongue Palate* Tongue Palate* Larynx SSM Jaw

DA Aud Lat Cbm

HST 722 Speech Motor Control

Feedforward control subsystem Sensorimotor adaptation to F1 perturbation

HST 722 Speech Motor Control

Feedback Control Subsystem

Auditory Error (Auditory Cortex)

Articulator Velocity and Position Cells (Motor Cortex)

Auditory FeedbackBased Command

Somatosensory Error (Somatosensory Cortex)

Somatosensory FeedbackBased Command

HST 722 Speech Motor Control

Feedback Control Subsystem (continued)

Auditory Error (Auditory Cortex)

Articulator Velocity and Position Cells (Motor Cortex)

Auditory FeedbackBased Command

Somatosensory Error (Somatosensory Cortex)

Somatosensory FeedbackBased Command

HST 722 Speech Motor Control

Prediction: Auditory Error Cells