Association of orofacial with laryngeal and respiratory motor output during speech

Speech motor coordination most likely involves synaptic coupling among neural systems that innervate orofacial, laryngeal, and respiratory muscles. The nature and strength of coupling of the orofacial with the respiratory and laryngeal systems was studied indirectly by correlating orofacial speeds with fundamental frequency, vocal intensity, and inspiratory volume during speech. Fourteen adult subjects repeated a simple test utterance at varying rates and vocal intensities while recordings were obtained of the acoustic signal and movements of the upper lip, lower lip, tongue, jaw, rib cage, and abdomen. Across subjects and orofacial speed measures (14 subjects x 4 structures), significant correlations were obtained for fundamental frequency in 42 of 56 cases, for intensity in 35 of 56 cases, and for inspiratory volume in 14 of 56 cases. These results suggest that during speech production there is significant neural coupling of orofacial muscle systems with the laryngeal and respiratory systems as they are involved in vocalization. Comparisons across the four orofacial structures revealed higher correlations for the jaw relative to other orofacial structures. This suggests stronger connectivity between neural systems linking the jaw with the laryngeal and respiratory systems. This finding may be relevant to the frame/content theory of speech production, which suggests that the neural circuitry involved in jaw motor control for speech has evolved to form relatively strong linkages with systems involved in vocalization.     

Subglottal pressure, tracheal airflow, and intrinsic laryngeal muscle activity during rat ultrasound vocalization

Vocal production requires complex planning and coordination of respiratory, laryngeal, and vocal tract movements, which are incompletely understood in most mammals. Rats produce a variety of whistles in the ultrasonic range that are of communicative relevance and of importance as a model system, but the sources of acoustic variability were mostly unknown. The goal was to identify sources of fundamental frequency variability. Subglottal pressure, tracheal airflow, and electromyographic (EMG) data from two intrinsic laryngeal muscles were measured during 22-kHz and 50-kHz call production in awake, spontaneously behaving adult male rats. During ultrasound vocalization, subglottal pressure ranged between 0.8 and 1.9 kPa. Pressure differences between call types were not significant. The relation between fundamental frequency and subglottal pressure within call types was inconsistent. Experimental manipulations of subglottal pressure had only small effects on fundamental frequency. Tracheal airflow patterns were also inconsistently associated with frequency. Pressure and flow seem to play a small role in regulation of fundamental frequency. Muscle activity, however, is precisely regulated and very sensitive to alterations, presumably because of effects on resonance properties in the vocal tract. EMG activity of cricothyroid and thyroarytenoid muscle was tonic in calls with slow or no fundamental frequency modulations, like 22-kHz and flat 50-kHz calls. Both muscles showed brief high-amplitude, alternating bursts at rates up to 150 Hz during production of frequency-modulated 50-kHz calls. A differentiated and fine regulation of intrinsic laryngeal muscles is critical for normal ultrasound vocalization. Many features of the laryngeal muscle activation pattern during ultrasound vocalization in rats are shared with other mammals.     

A Songbird Animal Model for Dissecting the Genetic Bases of Autism Spectrum Disorder

The neural and genetic bases of human language development and associated neurodevelopmental disorders, including autism spectrum disorder (ASD), in which language impairment represents a core deficit, are poorly understood. Given that no single animal model can fully capture the behavioral and genetic complexity of ASD, work in songbird, an experimentally tractable animal model of vocal learning, can complement the valuable tool of rodent genetic models and contribute important insights to our understanding of the communication deficits observed in ASD. Like humans, but unlike traditional laboratory animals such as rodents or non-human primates, songbirds exhibit the capacity of vocal learning, a key subcomponent of language. Human speech and birdsong reveal important parallels, highlighting similar developmental critical periods, a homologous cortico-basal ganglia-thalamic circuitry, and a critical role for social influences in the learning of vocalizations. Here I highlight recent advances in using the songbird model to probe the cellular and molecular mechanisms underlying the formation and function of neural circuitry for birdsong and, by analogy, human language, with the ultimate goal of identifying any shared or human unique biological pathways underscoring language development and its disruption in ASD.     

Auditory detection and discrimination in deaf cats: psychophysical and neural thresholds for intracochlear electrical signals.

More than 30,000 hearing-impaired human subjects have learned to use cochlear implants for speech perception and speech discrimination. To understand the basic mechanisms underlying the successful application of contemporary speech processing strategies, it is important to investigate how complex electrical stimuli delivered to the cochlea are processed and represented in the central auditory system. A deaf animal model has been developed that allows direct comparison of psychophysical thresholds with central auditory neuronal thresholds to temporally modulated intracochlear electrical signals in the same animals. Behavioral detection thresholds were estimated in neonatally deafened cats for unmodulated pulse trains (e.g., 30 pulses/s or pps) and sinusoidal amplitude-modulated (SAM) pulse trains (e.g., 300 pps, SAM at 30 Hz; 300/30 AM). Animals were trained subsequently in a discrimination task to respond to changes in the modulation frequency of successive SAM signals (e.g., 300/8 AM vs. 300/30 AM). During acute physiological experiments, neural thresholds to pulse trains were estimated in the inferior colliculus (IC) and the primary auditory cortex (A1) of the anesthetized animals. Psychophysical detection thresholds for unmodulated and SAM pulse trains were virtually identical. Single IC neuron thresholds for SAM pulse trains showed a small but significant increase in threshold (0.4 dB or 15.5 microA) when compared with thresholds for unmodulated pulse trains. The mean difference between psychophysical and minimum neural thresholds within animals was not significant (mean = 0.3 dB). Importantly, cats also successfully discriminated changes in the modulation frequencies of the SAM signals. Performance on the discrimination task was not affected by carrier rate (100, 300, 500, 1,000, or 1,500 pps). These findings indicate that 1) behavioral and neural response thresholds are based on detection of the peak pulse amplitudes of the modulated and unmodulated signals, and 2) discrimination of successive SAM pulse trains is based on temporal resolution of the envelope frequencies. Overall, our animal model provides a robust framework for future studies of behavioral discrimination and central neural temporal processing of electrical signals applied to the deaf cochlea by a cochlear implant.     

Neuronal activity in the periaqueductal gray and bordering structures during vocal communication in the squirrel monkey

In seven freely moving squirrel monkeys (Saimiri sciureus), the neuronal activity in the periaqueductal gray (PAG) and bordering structures was registered during vocal communication, using a telemetric single-unit recording technique. In 9.3% of the PAG neurons, a vocalization-correlated activity was found. Four reaction types could be distinguished: a) neurons, showing an activity burst immediately before vocalization onset; b) neurons, firing during vocalization, and starting shortly before vocalization onset; c) neurons, firing exclusively during vocalization; d) neurons, firing in the interval between perceived vocalizations (i.e. vocalizations produced by group mates) and self-produced vocal response. All PAG neurons showed a marked vocalization-type specificity. None of the neurons reflected simple acoustic parameters, such as fundamental frequency or amplitude, in its discharge rate. None of the neurons reacted to vocalizations of other animals not responded to by the experimental animal. All four reaction types found in the PAG were also found in the reticular formation bordering the PAG, though in lower density.     

Neural firing in ventrolateral thalamic nucleus during conditioned vocal behavior in cats

 The ventrolateral (VL) thalamus in mammals is a site well-situated to show vocalization-related neural activity if there is general or classical motor system involvement in vocal production. It receives input from both the basal ganglia and cerebellum, and forms reciprocal connections with motor cortical areas. The current study examined the activity in cat VL thalamus neurons during instrumentally conditioned vocalization. Units in our sample showed irregular spontaneous firing which could be modulated by slowly occurring fluctuations in intensity of vocalization task performance. Two main types of behavioral events were associated with changes in neural firing rate. The first of these was the ingestion of food reward. More than half of all recordings showed phasic bursting patterns during licking; a similar number had increases in firing preparatory to this phasic activity. The second behavioral event modulating unit responses was vocalization. Approximately 60% of recordings showed activity changes time-locked to vocalization. These responses were almost always excitatory, and often involved changes in firing that preceded vocalization onset. No spatial organization of differences in firing pattern between neurons could be distinguished. Our results suggest that VL thalamus may well be involved in mediating vocal behavior, although its functional role remains an object of speculation. Results are compared with previous studies of vocalization-related activity and of VL thalamus activity. Received: 15 November 1995 / Accepted: 16 December 1996     

Task-specific sensorimotor interactions in speech production

Speaking involves the activity of multiple muscles moving many parts (articulators) of the vocal tract. In previous studies, it has been shown that mechanical perturbation delivered to one moving speech articulator, such as the lower lip or jaw, results in compensatory responses in the perturbed and other non-perturbed articulators, but not in articulators that are uninvolved in the specific speech sound being produced. These observations suggest that the speech motor control system may be organized in a task-specific manner. However, previous studies have not used the appropriate controls to address the mechanism by which this task-specific organization is achieved. A lack of response in a non-perturbed articulator may simply reflect the fact that the muscles examined were not active. Alternatively, there may be a specific gating of somatic sensory signals due to task requirements. The present study was designed to address the nature of the underlying sensorimotor organization. Unanticipated mechanical loads were applied to the upper lip during the "p" in "apa" and "f" in "afa" in six subjects. Both lips are used to produce "p", while only the lower lip is used for "f". For "apa", both upper lip and lower lip responses were observed following upper lip perturbation. For "afa", no upper lip or lower lip responses were observed following the upper lip perturbation. The differential response of the lower lip, which was phasically active during both speech tasks, indicates that the neural organization of these two speech tasks differs not only in terms of the different muscles used to produce the different movements, but also in terms of the sensorimotor interactions within and across the two lips.     

Speech sounds alter facial skin sensation

Interactions between auditory and somatosensory information are relevant to the neural processing of speech since speech processes and certainly speech production involves both auditory information and inputs that arise from the muscles and tissues of the vocal tract. We previously demonstrated that somatosensory inputs associated with facial skin deformation alter the perceptual processing of speech sounds. We show here that the reverse is also true, that speech sounds alter the perception of facial somatosensory inputs. As a somatosensory task, we used a robotic device to create patterns of facial skin deformation that would normally accompany speech production. We found that the perception of the facial skin deformation was altered by speech sounds in a manner that reflects the way in which auditory and somatosensory effects are linked in speech production. The modulation of orofacial somatosensory processing by auditory inputs was specific to speech and likewise to facial skin deformation. Somatosensory judgments were not affected when the skin deformation was delivered to the forearm or palm or when the facial skin deformation accompanied nonspeech sounds. The perceptual modulation that we observed in conjunction with speech sounds shows that speech sounds specifically affect neural processing in the facial somatosensory system and suggest the involvement of the somatosensory system in both the production and perceptual processing of speech.     

A method for measurement of the vocal tract impedance at the mouth

In this contribution a method is presented for the measurement of vocal tract resonances. The technique uses a non-invasive acoustic excitation of the vocal tract and a fast and robust detection. The method is an alternative to the linear predictive coding (LPC) analysis for patients with voice and speech disorders. Sweep signals are emitted and recorded simultaneously from the small end of a tube placed in front of the mouth opening. The use of a pressure sensor and a velocity sensor provides a direct measurement of the vocal tract impedance at the mouth (VTMI). For selected sustained German vowels, and some consonants, a comparison of results from LPC analysis and VTMI measurements is given. The results indicate a good agreement in the frequency range from 500 to 5,000 Hz. The feasibility of the VTMI method for diagnostic and therapeutic applications is subject to current research.     

Sensory-motor interaction in the primate auditory cortex during self-initiated vocalizations

Little is known about sensory-motor interaction in the auditory cortex of primates at the level of single neurons and its role in supporting vocal communication. The present study investigated single-unit activities in the auditory cortex of a vocal primate, the common marmoset (Callithrix jacchus), during self-initiated vocalizations. We found that 1) self-initiated vocalizations resulted in suppression of neural discharges in a majority of auditory cortical neurons. The vocalization-induced inhibition suppressed both spontaneous and stimulus-driven discharges. Suppressed units responded poorly to external acoustic stimuli during vocalization. 2) Vocalization-induced suppression began several hundred milliseconds prior to the onset of vocalization. 3) The suppression of cortical discharges reduced neural firings to below the rates expected from a unit's rate-level function, adjusted for known subcortical attenuation, and therefore was likely not entirely caused by subcortical attenuation mechanisms. 4) A smaller population of auditory cortical neurons showed increased discharges during self-initiated vocalizations. This vocalization-related excitation began after the onset of vocalization and is likely the result of acoustic feedback. Units showing this excitation responded nearly normally to external stimuli during vocalization. Based on these findings, we propose that the suppression of auditory cortical neurons, possibly originating from cortical vocal production centers, acts to increase the dynamic range of cortical responses to vocalization feedback for self monitoring. The excitatory responses, on the other hand, likely play a role in maintaining hearing sensitivity to the external acoustic environment during vocalization.     

Contextual cuing contributes to the independent modification of multiple internal models for vocal control

Research on the control of visually guided limb movements indicates that the brain learns and continuously updates an internal model that maps the relationship between motor commands and sensory feedback. A growing body of work suggests that an internal model that relates motor commands to sensory feedback also supports vocal control. There is evidence from arm-reaching studies that shows that when provided with a contextual cue, the motor system can acquire multiple internal models, which allows an animal to adapt to different perturbations in diverse contexts. In this study we show that trained singers can rapidly acquire multiple internal models regarding voice fundamental frequency (F(0)). These models accommodate different perturbations to ongoing auditory feedback. Participants heard three musical notes and reproduced each one in succession. The musical targets could serve as a contextual cue to indicate which direction (up or down) feedback would be altered on each trial; however, participants were not explicitly instructed to use this strategy. When participants were gradually exposed to altered feedback adaptation was observed immediately following vocal onset. Aftereffects were target specific and did not influence vocal productions on subsequent trials. When target notes were no longer a contextual cue, adaptation occurred during altered feedback trials and evidence for trial-by-trial adaptation was found. These findings indicate that the brain is exceptionally sensitive to the deviations between auditory feedback and the predicted consequence of a motor command during vocalization. Moreover, these results indicate that, with contextual cues, the vocal control system may maintain multiple internal models that are capable of independent modification during different tasks or environments.     

Concurrent encoding of frequency and amplitude modulation in human auditory cortex: encoding transition

Complex natural sounds (e.g., animal vocalizations or speech) can be characterized by specific spectrotemporal patterns the components of which change in both frequency (FM) and amplitude (AM). The neural coding of AM and FM has been widely studied in humans and animals but typically with either pure AM or pure FM stimuli. The neural mechanisms employed to perceptually unify AM and FM acoustic features remain unclear. Using stimuli with simultaneous sinusoidal AM (at rate f(AM) = 37 Hz) and FM (with varying rates f(FM)), magnetoencephalography (MEG) is used to investigate the elicited auditory steady-state response (aSSR) at relevant frequencies (f(AM), f(FM), f(AM) + f(FM)). Previous work demonstrated that for sounds with slower FM dynamics (f(FM) < 5 Hz), the phase of the aSSR at f(AM) tracked the FM; in other words, AM and FM features were co-tracked and co-represented by "phase modulation" encoding. This study explores the neural coding mechanism for stimuli with faster FM dynamics (< or =30 Hz), demonstrating that at faster rates (f(FM) > 5 Hz), there is a transition from pure phase modulation encoding to a single-upper-sideband (SSB) response (at frequency f(AM) + f(FM)) pattern. We propose that this unexpected SSB response can be explained by the additional involvement of subsidiary AM encoding responses simultaneously to, and in quadrature with, the ongoing phase modulation. These results, using MEG to reveal a possible neural encoding of specific acoustic properties, demonstrate more generally that physiological tests of encoding hypotheses can be performed noninvasively on human subjects, complementing invasive, single-unit recordings in animals.     

Factors influencing neural activity in parabrachial regions during cat vocalizations

Summary The parabrachial nucleus in mammals is intimately connected with other vocalization controlling brainstem structures. It, along with ventromedially adjacent structures, also has been identified as the pneumotaxic center, and as such shows strong respiratory related activity in the anesthetized cat. The current study examines the neuronal activity in cat parabrachial regions during production of instrumentally conditioned vocalizations. Most of the units in our sample show considerable activity during periods between vocalizations. For many units, firing rate fluctuates during the respiratory cycle, although apparently not as strongly as reported in the decerebrate cat. Also, there is often strong phasic activity during periods where animals are licking to ingest their food rewards. During the peri-vocalization period, various neural activity patterns can be recorded. Most common is an activity increase during the vocalization itself. Moreover, in some units, this activity increase has an auditory component. A smaller number of units show other activity patterns, including a suppression of activity during vocalization and activity increases preceding the vocalization. Overall, our results suggest that the parabrachial region's involvement in vocal control is quite complex, involving convergence of respiratory, acoustic, vocalization-related, and perhaps somatosensory influences.     

Speech motor learning in profoundly deaf adults

Speech production, like other sensorimotor behaviors, relies on multiple sensory inputs--audition, proprioceptive inputs from muscle spindles and cutaneous inputs from mechanoreceptors in the skin and soft tissues of the vocal tract. However, the capacity for intelligible speech by deaf speakers suggests that somatosensory input alone may contribute to speech motor control and perhaps even to speech learning. We assessed speech motor learning in cochlear implant recipients who were tested with their implants turned off. A robotic device was used to alter somatosensory feedback by displacing the jaw during speech. We found that implant subjects progressively adapted to the mechanical perturbation with training. Moreover, the corrections that we observed were for movement deviations that were exceedingly small, on the order of millimeters, indicating that speakers have precise somatosensory expectations. Speech motor learning is substantially dependent on somatosensory input.     

鸣禽古纹状体粗核的传出投射

用生物素结合的葡聚精胺顺行示踪技术对鸣禽斑胸草雀古纹状体粗核的传出投射进行了研究．结果表明：古纹状体粗核作为发声控制的前脑运动核团之一，不仅可直接投射至中脑丘间复合体背内侧核和延髓舌下神经核气管鸣管部，而且还进一步发现其投射至延髓喙端腹外侧核、疑核、后疑核。这些发现提示，古织状体租核与伴随发声的呼吸协调有关．     

Cutting the vagus nerve below the diaphragm prevents maternal potentiation of infant rat vocalization

In maternal potentiation, the rate of vocalization by a young organism during isolation is greatly enhanced if that isolation has been immediately preceded by an interaction with the mother (or other adult female in the case of rats). The enhancement in isolation-induced vocalization rate does not occur if the young animal had an interaction with other social companions like littermates or with familiar inanimate stimuli like home cage shavings. The present study demonstrates that pups whose vagus nerve is cut below the diaphragm do not potentiate vocalization after an interaction with their dam. The vocalization rates of denervated pups in a first isolation, in the presence of the dam, and during cold exposure do not differ from control pups. Their non-vocal behaviors also appear unaffected by the surgery. Similar to what has been shown in studies of fever-induced behavioral changes, an intact vagus nerve from the gut is necessary for young rat pups to show normal social mediation of their isolation-induced vocal responses.     

Virtual vocalization stimuli for investigating neural representations of species-specific vocalizations

Most studies investigating neural representations of species-specific vocalizations in non-human primates and other species have involved studying neural responses to vocalization tokens. One limitation of such approaches is the difficulty in determining which acoustical features of vocalizations evoke neural responses. Traditionally used filtering techniques are often inadequate in manipulating features of complex vocalizations. Furthermore, the use of vocalization tokens cannot fully account for intrinsic stochastic variations of vocalizations that are crucial in understanding the neural codes for categorizing and discriminating vocalizations differing along multiple feature dimensions. In this work, we have taken a rigorous and novel approach to the study of species-specific vocalization processing by creating parametric "virtual vocalization" models of major call types produced by the common marmoset (Callithrix jacchus). The main findings are as follows. 1) Acoustical parameters were measured from a database of the four major call types of the common marmoset. This database was obtained from eight different individuals, and for each individual, we typically obtained hundreds of samples of each major call type. 2) These feature measurements were employed to parameterize models defining representative virtual vocalizations of each call type for each of the eight animals as well as an overall species-representative virtual vocalization averaged across individuals for each call type. 3) Using the same feature-measurement that was applied to the vocalization samples, we measured acoustical features of the virtual vocalizations, including features not explicitly modeled and found the virtual vocalizations to be statistically representative of the callers and call types. 4) The accuracy of the virtual vocalizations was further confirmed by comparing neural responses to real and synthetic virtual vocalizations recorded from awake marmoset auditory cortex. We found a strong agreement between the responses to token vocalizations and their synthetic counterparts. 5) We demonstrated how these virtual vocalization stimuli could be employed to precisely and quantitatively define the notion of vocalization "selectivity" by using stimuli with parameter values both within and outside the naturally occurring ranges. We also showed the potential of the virtual vocalization stimuli in studying issues related to vocalization categorizations by morphing between different call types and individual callers.     

The highly specialized vocal tract of the male Mongolian gazelle (Procapra gutturosa Pallas, 1777--Mammalia, Bovidae)

The entire head and neck of a wild adult male Mongolian gazelle (Procapra gutturosa) was dissected with special reference to its enlarged larynx. Two additional adult male specimens taken from the wild were analysed by computer tomography. The sternomandibularis, omohyoideus, thyrohyoideus and hyoepiglotticus muscles are particularly enlarged and improve laryngeal suspension and stabilization. The epiglottis is exceptionally large. A permanent laryngeal descent is associated with the evolution of an unpaired palatinal pharyngeal pouch. A certain momentary descent seems to occur during vocalization. The high lateral walls of the thyroid cartilage are ventrally connected by a broad keel. The large thyroarytenoid muscle is divided into two portions: a rostral ventricularis and a caudal vocalis muscle. A paired lateral laryngeal ventricle projects between these two muscles. The massive vocal fold is large and lacks any rostrally directed flexible structures. It is supported by a large cymbal-like fibroelastic pad. Vocal tract length was measured in the course of dissection and in computer tomographic images. Two representative spectrograms, one of an adult male and one of a juvenile, recorded in the natural habitat of the Mongolian gazelle are presented. In the spectrograms, the centre frequency of the lowest band is about 500 Hz in the adult male and about 790 Hz in the juvenile. The low pitch of the adult male's call is ascribed to the evolutionary mass increase and elongation of the vocal folds. In the habitat of P. gutturosa a call with a low pitch and, thus, with an almost homogeneous directivity around the head of the vocalizing animal may be optimally suited for multidirectional advertisement calls during the rut. The signal range of an adult male's call in its natural habitat can therefore be expected to be larger than the high-pitched call of a juvenile.     

Motor timing learned without motor training

Improvements due to perceptual training are often specific to the trained task and do not generalize to similar perceptual tasks. Surprisingly, given this history of highly constrained, context-specific perceptual learning, we found that training on a perceptual task showed significant transfer to a motor task. This result provides evidence for a common neural architecture underlying analysis of sensory input and control of motor output, and suggests a potential role for perception in motor development and rehabilitation.     

Cutaneous reflexes during rhythmic arm cycling are insensitive to asymmetrical changes in crank length

The neural control of a movement depends upon the motor task performed. To further understand the neural regulation of different variations of the same type of movement, we created three dissimilar bilateral rhythmic arm cycling tasks by unilaterally manipulating crank length (CL). Modulation in the amplitude and sign of cutaneous reflexes was used as an index of neural control. Neurologically intact subjects performed three bilateral cycling trials at ∼1 Hz with the ipsilateral crank arm at one of three different lengths. Cutaneous reflexes were evoked during each trial with trains (5 × 1.0 ms pulses at 300 Hz) of electrical stimulation delivered to the superficial radial nerve at the ipsilateral wrist. EMG recordings were made bilaterally from muscles acting at the shoulder, elbow, and wrist. Analysis was conducted after phase-averaging contingent upon the timing of stimulation in the movement cycle. CL variation created an asymmetrical cycling pattern and produced significant changes in the range of motion at the ipsilateral shoulder and elbow. Background EMG amplitude in muscles of the contralateral arm generally increased significantly as CL decreased. Therefore at a given phase in the movement cycle, the background EMG was different between the three cycling trials. In contrast, cutaneous reflex amplitudes in muscles of both arms were similar at each phase of the movement cycle between the different CLs trials at both early and middle latencies. This was particularly evident in muscles ipsilateral to nerve stimulation. We suggest that variations of arm cycling that primarily yield significant changes in the amplitude of muscle activity do not require significant task-specific change in neural control.