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Biomedical subjects

C R Larson

Publications and source records attributed to C R Larson.

At least 37 records · Page 2Linked to original sources

Cerebral averaged potentials preceding oral movement.

The "readiness potential" is an event-related potential that shows increasing negativity at vertex and motor strip scalp recording sites prior to voluntary, unilateral limb movements. Though speech involves movement on both sides of the midline, recent recordings of prespeech potentials suggest a pattern of bilateral activation that lateralizes to the dominant hemisphere just prior to the onset of articulatory movement. To determine whether this pattern of dominant hemisphere activation is present prior to a stereotyped, nonspeech movement of the mouth, the averaged potentials preceding a lip protrusion task were recorded at the cranial vertex and over the right and left motor cortex. Results were compared to potentials preceding a right finger extension task performed by the same subjects. Both the finger and the lip movements were initially preceded by slow negative potentials. Prior to the finger extension task, the negative amplitude became greatest over the left motor cortex, contralateral to the side of movement. Prior to the lip protrusion task, the amplitude of the potential remained even over the right and left motor cortices. The results suggest that, for this nonspeech movement of a midline structure, bilateral cortical control takes place. Control of lip movement is apparently not necessarily a dominant hemisphere function, though dominance may become part of the motor control strategy for more complex movements such as those used during speech.

Action Potentials↗

Stutterers' and nonstutterers' voice fundamental frequency changes in response to auditory stimuli.

An acoustic analysis of certain aspects of voice fundamental frequency (Fo) in response to auditory clicks may provide a tool for studying short-latency auditory-laryngeal reflexes in humans (Sapir, McClean, & Larson, 1983). The purpose of this investigation was to compare 18 adult male stutterers' and 19 adult male nonstutterers' auditory-laryngeal reflexes. Subjects sustained phonation at constant pitch and intensity levels while receiving bilateral auditory click stimuli. Fo signal averages were generated and measured. Data analyses indicated that there were no differences between stutterers and nonstutterers with respect to the frequency of reflex occurrence. Also, there were no significant group differences with respect to the mean reflex temporal measures or the variability of the reflex temporal measures.

Acoustic Stimulation↗

Changes in voice fundamental frequency following discharge of single motor units in cricothyroid and thyroarytenoid muscles.

This investigation was designed to measure voice Fo changes related to single motor unit (SMU) contractions in the cricothyroid and thyroarytenoid muscles. Four subjects (3 men and 1 woman) were recorded producing a prolonged vowel at modal pitch and loudness levels while simultaneous recordings of electromyograms (EMG) from the muscles were obtained. Voice Fo changes unrelated to SMU firings in the muscles were eliminated using an averaging method previously described by Baer (1979). Results indicate that the time between discharge of the SMU and the peak in Fo change ("Fo Latency") was variable and ranged from 5 to 20 ms for the thyroarytenoid and 6 to 75 ms for the cricothyroid muscle. Distinct oscillations in Fo were always present in recordings from the woman subject and from the men when they phonated at higher-than-modal pitch levels. The findings are discussed in relation to SMU contraction times, biomechanics of the vocal folds, and the presence of jitter in normal voices.

Adult↗

The relationship of periaqueductal gray neurons to vocalization and laryngeal EMG in the behaving monkey.

The midbrain periaqueductal gray (PAG) of most higher animals has been shown by stimulation and lesion methods to be important in vocalization. In order to learn how the PAG is involved in vocalization, activity from single PAG neurons was recorded from 3 awake, vocalizing monkeys. From a population of 149 units that were temporally related to vocalization, 91 were analyzed with respect to specific parameters of vocalization and laryngeal EMG activity. Measures of the activity of 52 units were significantly correlated with vocalization or EMG activity. Units tended to be correlated with only a few measures of vocalization or EMG activity suggesting rather specific relationships between PAG units and vocalization measures. Microstimulation near recorded cells usually did not excite every muscle sampled, suggesting PAG projections to brainstem motor nuclei may be somewhat specific. The results confirm previous suggestions that the PAG may be involved in the coordination of brainstem motor nuclei during vocalization.

Animals↗

The midbrain periaqueductal gray: a brainstem structure involved in vocalization.

Experiments were performed on one Macaca fascicularis and three Macaca mulatta monkeys. Electrical stimulation of the midbrain periaqueductal gray (PAG) elicited phonation closely resembling that naturally produced by such animals. Electromyographic (EMG) recordings from laryngeal, thoracic, tongue, and facial muscles showed increased levels of activity following PAG stimulation. Recordings were made from single neurons located within the PAG in one monkey. These neurons increased their firing rates prior to the onset of laryngeal electromyographic activity or vocalization. Spike-triggered averaging and microstimulation indicated that some of the PAG neurons may project to the laryngeal system. The function of the PAG is discussed with regard to vocal initiation, voice intensity, coordination of different muscle systems during vocalization, and for emotional vocalizations.

Animals↗

Periaqueductal gray neuronal activity associated with laryngeal EMG and vocalization in the awake monkey.

Extracellular recordings were made from 73 periaqueductal gray (PAG) neurons in an awake, vocalizing monkey (Macaca fascicularis). About 50% of the neurons were inactive and only began discharging prior to laryngeal EMG activity and vocalization; they reached their peak firing rate near the onset of vocalization and then became inactive again. Spike-triggered averaging of laryngeal EMG from the PAG unit discharge revealed in most cases an increase in laryngeal EMG immediately following the unit spike. Other cells appeared to be related to other oral-facial behaviors. The role of the PAG in vocalization is discussed.

Animals↗

Human laryngeal responses to auditory stimulation.

In spite of various claims for the presence and functional significance of brainstem reflexes in phonatory control, very little is known about these reflexes in humans. The present study was aimed at studying auditory-laryngeal reflexes in human subjects during sustained phonation. Fourteen subjects sustained phonation at constant voice pitch and voice intensity while receiving auditory stimulation with clicks of different intensity levels. Electromyographic (EMG) recordings from the cricothyroid muscle were obtained in two of the subjects. The voice signal was transduced with a throat microphone and bandpass filtered to isolate the fundamental frequency (FO). The filtered signal was fed to a frequency meter, which yielded a voltage proportional to FO. This FO voltage and the rectified EMG were signal averaged relative to the click. Short-latency changes in FO (50 ms) and in cricothyroid EMG (11 ms) were observed in response to auditory stimulation. The magnitude of the EMG and FO responses were well correlated with each other and with the intensity of the stimulus. The EMG response did not habituate with repeated stimulation. The neuromuscular origins of the auditory-laryngeal reflexes and their significance to phonatory control are discussed.

Acoustic Stimulation↗

Jaw muscle afferent firing during an isotonic jaw-positioning task in the monkey.

The activity of jaw muscle receptors was studied by recording neurons in the mesencephalic nucleus of the trigeminal nerve in monkeys trained to control the position and movement of their mandible. Jaw position was measured by a weighted lever resting on the mandibular incisors. The force required to maintain the position of the lever was varied; in most cases it was either 25 or 360 g. Firing rates of neurons were related to stationary mandibular positions and to the velocity of movements during intervals when the movement velocity was constant. Of 49 neurons studied in detail, 21 fired at rates that were consistently and linearly related to static incisal openings. This static position sensitivity was typically about 5 spikes/mm of incisal opening. Most position-sensitive neurons fired at higher rates during opening movements and at lower rates during closing movements than would be accounted for by their position sensitivity. This sensitivity to the velocity of movement was not linear, however; slow closing movements sometimes did not produce a decrease in firing rate, and an actual increase during muscle shortening was seen in a few instances. The position sensitivity of eight neurons was evaluated during different loading conditions; in no case did it change substantially. Of the remaining 28 neurons, 26 fired at high rates during all opening movements and either stopped firing or fired at low, sporadic rates during closing movements. The static position sensitivity of these neurons was weak and variable both within and between neurons. The velocity sensitivity of these stretch-sensitive neurons was very nonlinear. Except for a range of slow movements (+/- 5 mm/s), the firing rate was maximal (200 spikes/s or higher) for most opening movements and zero for most closing movements. Maximal firing rates were higher when the loads being moved were increased from 25 to 360 g. The majority of position-sensitive neurons exhibited a large interspike-interval variability at wide incisal opening. In most of these neurons, this interspike-interval variability was periodic, usually at a rate of about 10 periods/s, and took the form of "saw-tooth" modulation on a record of instantaneous firing rate. Neurons that exhibited this modulation in a very prominent form also exhibited, in many instances, a substantial increase in firing rate during closing jaw movements.

Afferent Pathways↗

Evidence that the human jaw stretch reflex increases the resistance of the mandible to small displacements.

1. Small 'step' or sinusoidal displacements were imposed on the mandible while human subjects maintained an average biting force of 10 N. Phase-related changes in the force resisting sinusoidal displacement were used to determine the mechanical stiffness of the human mandibular system as a function of the frequency of stretching. 2. Jaw-muscle electromyographic (e.m.g.) responses to 'step' stretches were of 8 msec latency and generated a very substantial force response. Jaw-muscle e.m.g. responses having longer latency were not observed. 3. The mechanical stiffness of the human mandible was relatively constant as a function of the frequency of stretching, having a typical magnitude of about 15 N/mm (+/- 200 micrometers stretch) or 10 N/mm (+/- 1500 micrometers stretch) at mean biting forces of 10 N. The force resisting displacement was phase-advanced at all frequencies. 4. Modulation of jaw muscle electrical activity evoked by sinusoidal stretches increased in amplitude as a function of increasing stretch frequency. E.m.g. modulation was 60--100 degrees advanced at frequencies of 1--10 Hz, but the phase decreased at higher frequencies, becoming negative (lagging stretch) at frequencies of 30 Hz and above. These characteristics are consistent with the idea that the jaw stretch reflex is dependent on jaw muscle spindle afferent fibres exciting jaw-closing motoneurones by relatively direct (but not necessarily monosynaptic) connexions. 5. The relationship between jaw-muscle activity and voluntary fluctuations of isometric biting force suggests that human jaw muscles can be modelled as a second-order linear 'filter'. The corner frequency for human jaw muscle is about 3 Hz; thus it would appear to be considerably slower than jaw muscle of monkeys. 6. The reflex stiffness of the human mandible, estimated quantitatively on the assumption that human jaw muscle stiffness is similar to the intrinsic stiffness of the gastrocnemius of the cat, ranges between 5 and 9 N/mm at frequencies between 1 and 8 Hz. Since this reflex stiffness is about the same as muscle stiffness in this frequency range, we conclude that the stretch reflex of the human mandible contributes functionally to its postural stability. 7. Reflex stiffness appears to be greater in the monkey mandible relative to muscle stiffness than in the human mandible. The difference is argued to be a manifestation of the difference in jaw muscle contraction speed between the two species. 8. The fact that the mandibular stretch reflex appears to be stronger than the stretch reflex of the limbs of intact animals and humans is discussed in terms of the special anatomical and functional features of the mandible.

Adult↗

Cerebellar regulation of phonation in rhesus monkey (Macaca mulatta).

Five rhesus monkeys (Macaca mulatta) were trained to emit a "coo" vocalization with a duration of at least 500 msec. After stable performance was achieved cerebellar lesions were introduced, and various pre- and postlesion measures of phonation and of laryngeal EMG activity were compared to assess the effects of the lesions. The phonatory changes were interpreted with respect to possible laryngeal or respiratory modifications. The relation between fundamental frequency and intensity of phonation was changed in some animals, with no obvious alterations in either fundamental frequency or intensity considered separately. Intensity of phonation was decreased while duration was prolonged in two animals. Fundamental frequency of phonation was also affected in some animals. Reliable laryngeal EMG was obtained in two animals and was affected by cerebellar lesions. These results indicate that normal cerebellar function is involved in the control of fundamental frequency, intensity, duration, and the coordination of the laryngeal and respiratory systems for the control of phonation.

Acoustics↗

Effects of cerebellar lesions on monkey jaw-force control: implications for understanding ataxic dysarthria.

Eight macaques were trained on one of two isometric force regulation tasks. Six animals were trained to maintain a static force for a 2-sec period, and two animals were trained to make a rapid series of bites on a force transducer four times in succession. Electromyographic (EMG) electrodes were chronically implanted into temporalis and masseter muscles. Lesions were made in the cerebellum in seven monkeys and a control surgery with no neural damage was performed on one monkey. Following cerebellar lesions, the abilities of the monkeys to maintain a static isometric force for 2 sec were impaired. Animals overshot the correct force band and force tremors increased in amplitude following the lesions. Cerebellar lesions were followed also by a reduction in biting frequency from about 3 Hz to 2 Hz. Measurements of the temporalis EMG indicated that the reduced biting rate was associated primarily with a prolongation of the inter-EMG interval and secondarily with a prolongation of the EMG duration. Changes in jaw-force regulation following the lesions appear to be similar to the types of changes observed in arm control in monkeys, and in the speech of persons with ataxic dysarthria.

Action Potentials↗