Domestic cat larynges can produce purring frequencies without neural input.

Most mammals produce vocal sounds according to the myoelastic-aerodynamic (MEAD) principle, through self-sustaining oscillation of laryngeal tissues.1,2 In contrast, cats have long been believed to produce their low-frequency purr vocalizations through a radically different mechanism involving active muscle contractions (AMC), where neurally driven electromyographic burst patterns (typically at 20-30 Hz) cause the intrinsic laryngeal muscles to actively modulate the respiratory airflow. Direct empirical evidence for this AMC mechanism is sparse.3 Here, the fundamental frequency (fo) ranges of eight domestic cats (Felis silvestris catus) were investigated in an excised larynx setup, to test the prediction of the AMC hypothesis that vibration should be impossible without neuromuscular activity, and thus unattainable in excised larynx setups, which are based on MEAD principles. Surprisingly, all eight excised larynges produced self-sustained oscillations at typical cat purring rates. Histological analysis of cat larynges revealed the presence of connective tissue masses, up to 4 mm in diameter, embedded in the vocal fold.4 This vocal fold specialization appears to allow the unusually low fo values observed in purring. While our data do not fully reject the AMC hypothesis for purring, they show that cat larynges can easily produce sounds in the purr regime with fundamental frequencies of 25 to 30 Hz without neural input or muscular contraction. This strongly suggests that the physical and physiological basis of cat purring involves the same MEAD-based mechanisms as other cat vocalizations (e.g., meows) and most other vertebrate vocalizations but is potentially augmented by AMC.

Subglottal pressure oscillations in anechoic and resonant conditions and their influence on excised larynx phonations.

Excised larynges serve as natural models for studying behavior of the voice source. Acoustic resonances inside the air-supplying tubes below the larynx (i.e., subglottal space), however, interact with the vibratory behavior of the larynges and obscure their inherent vibration properties. Here, we explore a newly designed anechoic subglottal space which allows removing its acoustic resonances. We performed excised larynx experiments using both anechoic and resonant subglottal spaces in order to analyze and compare, for the very first time, the corresponding subglottal pressures, electroglottographic and radiated acoustic waveforms. In contrast to the resonant conditions, the anechoic subglottal pressure waveforms showed negligible oscillations during the vocal fold contact phase, as expected. When inverted, these waveforms closely matched the inverse filtered radiated sound waveforms. Subglottal resonances modified also the radiated sound pressures (Level 1 interactions). Furthermore, they changed the fundamental frequency (fo) of the vocal fold oscillations and offset phonation threshold pressures (Level 2 interactions), even for subglottal resonance frequencies 4-10 times higher than fo. The obtained data offer the basis for better understanding the inherent vibratory properties of the vocal folds, for studying the impact of structure-acoustic interactions on voice, and for validation of computational models of voice production.

Hemi-laryngeal Setup for Studying Vocal Fold Vibration in Three Dimensions.

The voice of humans and most non-human mammals is generated in the larynx through self-sustaining oscillation of the vocal folds. Direct visual documentation of vocal fold vibration is challenging, particularly in non-human mammals. As an alternative, excised larynx experiments provide the opportunity to investigate vocal fold vibration under controlled physiological and physical conditions. However, the use of a full larynx merely provides a top view of the vocal folds, excluding crucial portions of the oscillating structures from observation during their interaction with aerodynamic forces. This limitation can be overcome by utilizing a hemi-larynx setup where one half of the larynx is mid-sagittally removed, providing both a superior and a lateral view of the remaining vocal fold during self-sustained oscillation. Here, a step-by-step guide for the anatomical preparation of hemi-laryngeal structures and their mounting on the laboratory bench is given. Exemplary phonation of the hemi-larynx preparation is documented with high-speed video data captured by two synchronized cameras (superior and lateral views), showing three-dimensional vocal fold motion and corresponding time-varying contact area. The documentation of the hemi-larynx setup in this publication will facilitate application and reliable repeatability in experimental research, providing voice scientists with the potential to better understand the biomechanics of voice production.

Relationship Between the Electroglottographic Signal and Vocal Fold Contact Area.

OBJECTIVE: Electroglottography (EGG) is a widely used noninvasive method that purports to measure changes in relative vocal fold contact area (VFCA) during phonation. Despite its broad application, the putative direct relation between the EGG waveform and VFCA has to date only been formally tested in a single study, suggesting an approximately linear relationship. However, in that study, flow-induced vocal fold (VF) vibration was not investigated. A rigorous empirical evaluation of EGG as a measure of VFCA under proper physiological conditions is therefore still needed. METHODS/DESIGN: Three red deer larynges were phonated in an excised hemilarynx preparation using a conducting glass plate. The time-varying contact between the VF and the glass plate was assessed by high-speed video recordings at 6000 fps, synchronized to the EGG signal. RESULTS: The average differences between the normalized [0, 1] VFCA and EGG waveforms for the three larynges were 0.180 (±0.156), 0.075 (±0.115), and 0.168 (±0.184) in the contacting phase and 0.159 (±0.112), -0.003 (±0.029), and 0.004 (±0.032) in the decontacting phase. DISCUSSIONS AND CONCLUSIONS: Overall, there was a better agreement between VFCA and the EGG waveform in the decontacting phase than in the contacting phase. Disagreements may be caused by nonuniform tissue conductance properties, electrode placement, and electroglottograph hardware circuitry. Pending further research, the EGG waveform may be a reasonable first approximation to change in medial contact area between the VFs during phonation. However, any quantitative and statistical data derived from EGG should be interpreted cautiously, allowing for potential deviations from true VFCA.

Vocal Fold Adjustment Caused by Phonation Into a Tube: A Double-Case Study Using Computed Tomography.

OBJECTIVES: Phonation into a tube is a widely used method for vocal training and therapy. Previous studies and practical experience show that the phonation becomes easier and louder after such an exercise. The purpose of this study was to find out whether there are systematic changes in the vocal fold adjustment after the exercise. METHODS: Two volunteer subjects (1 male and 1 female) without voice disorders were examined with computed tomography (CT). Both produced a sustained vowel [a:] at comfortable pitch and loudness before and after the tube phonation and a vowel-like phonation into the tube. Computed tomography (CT) scans were obtained before, during, and after the exercise, twice for each condition. The gathered CT images were used for measurements of vertical vocal fold thickness, bulkiness, length, and glottal width. RESULTS: No prominent trends common to both subjects were found in vocal fold adjustment during and after the phonation into the tube. Variability observed under the same conditions was usually of the same magnitude as the changes before and after the tube phonation. CONCLUSIONS: Changes in vocal tract configuration observed after the resonance tube exercises in previous related studies were more prominent than the changes in vocal fold configuration observed here.