Universal mechanisms of sound production and control in birds and mammals.

As animals vocalize, their vocal organ transforms motor commands into vocalizations for social communication. In birds, the physical mechanisms by which vocalizations are produced and controlled remain unresolved because of the extreme difficulty in obtaining in vivo measurements. Here, we introduce an ex vivo preparation of the avian vocal organ that allows simultaneous high-speed imaging, muscle stimulation and kinematic and acoustic analyses to reveal the mechanisms of vocal production in birds across a wide range of taxa. Remarkably, we show that all species tested employ the myoelastic-aerodynamic (MEAD) mechanism, the same mechanism used to produce human speech. Furthermore, we show substantial redundancy in the control of key vocal parameters ex vivo, suggesting that in vivo vocalizations may also not be specified by unique motor commands. We propose that such motor redundancy can aid vocal learning and is common to MEAD sound production across birds and mammals, including humans.

Resonance properties of the vocal folds: in vivo laryngoscopic investigation of the externally excited laryngeal vibrations.

The study presents the first attempt to investigate resonance properties of the living vocal folds by means of laryngoscopy. Laryngeal vibrations were excited via a shaker placed on the neck of a male subject and observed by means of videostroboscopy and videokymography (VKG). When the vocal folds were tuned to the phonation frequency of 110 Hz and sinusoidal vibration with sweeping frequency (in the range 50-400 Hz) was delivered to the larynx, three clearly pronounced resonance peaks at frequencies around 110, 170, and 240 Hz were identified in the vocal fold tissues. Different modes of vibration of the vocal folds, observed as distinct lateral-medial oscillations with one, two, and three half-wavelengths along the glottal length, respectively, were associated with these resonance frequencies. At the external excitation frequencies below 100 Hz, vibrations of the ventricular folds, aryepiglottic folds and arytenoid cartilages were dominant in the larynx.

On pitch jumps between chest and falsetto registers in voice: data from living and excised human larynges.

The paper offers a new concept of studying abrupt chest-falsetto register transitions (jumps) based on the theory of nonlinear dynamics. The jumps were studied in an excised human larynx and in three living subjects (one female and two male). Data from the excised larynx revealed that a small and gradual change in tension of the vocal folds can cause an abrupt change of register and pitch. This gives evidence that the register jumps are manifestations of bifurcations in the vocal-fold vibratory mechanism. A hysteresis was observed; the upward register jump occurred at higher pitches and tensions than the downward jump. Due to the hysteresis, the chest and falsetto registers can be produced with practically identical laryngeal adjustments within a certain range of longitudinal tensions. The magnitude of the frequency jump was measured as the "leap ratio" F0F:F0C (fundamental frequency of the falsetto related to that of the chest register) and alternatively expressed as a corresponding musical interval, termed the "leap interval." Ranges of this leap interval were found to be different for the three living subjects (0-5 semitones for the female, 5-10 and 10-17 for the two males, respectively). These differences are considered to reflect different biomechanical properties of the vocal folds of the examined subjects. A small magnitude of the leap interval was associated with a smooth chest-falsetto transition in the female subject.

First results of clinical application of videokymography.

OBJECTIVES: Stroboscopy is based on the assumption that the vibration of the vocal folds is stable and regular. Irregular vibrations, which are common in voice pathology, cannot easily be studied and described in a reliable way. Videokymography overcomes most of these drawbacks. DESIGN: The use of the recently invented videokymography for studying vocal fold vibrations in patients is introduced. METHOD: Videokymography, using a modified CCD-video camera, works in two modes: standard and high speed. In standard mode the vocal folds are displayed on a video monitor in the usual way, providing 50 images per second (or 60 in the National Television Standards Committee (NTSC) system). This is used for routine laryngoscopic and stroboscopic examination of the larynx. In high-speed mode (nearly 8000 images per second) only one line from the whole image is selected and displayed on the x-axis of the monitor; the y-axis represents the time dimension. RESULTS: All kinds of vocal fold vibrations, including those leading to pathological rough, breathy, hoarse, or diplophonic voice productions can be observed. Videokymography visualizes small left-right asymmetries, open quotient differences along the glottis, lateral propagation of mucosal waves, and movements of the upper margin and, sometimes in the closing phase, the lower margin of the vocal folds. CONCLUSION: Videokymography is advantageous for a more accurate diagnosis of voice disorders. Videokymography provides a simple way to study irregular vibrations of the vocal folds. Information is directly available for further processing and allows a first-time quantification of vibrations registered.

Videokymography: high-speed line scanning of vocal fold vibration.

A digital technique for high-speed visualization of vibration, called videokymography, was developed and applied to the vocal folds. The system uses a modified video camera able to work in two modes: high-speed (nearly 8,000 images/s) and standard (50 images/s in CCIR norm). In the high-speed mode, the camera selects one active horizontal line (transverse to the glottis) from the whole laryngeal image. The successive line images are presented in real time on a commercial TV monitor, filling each video frame from top to bottom. The system makes it possible to observe left-right asymmetries, open quotient, propagation of mucosal waves, movement of the upper and, in the closing phase, the lower margins of the vocal folds, etc. The technique is suitable for further processing and quantification of recorded vibration.

A subharmonic vibratory pattern in normal vocal folds.

This study observes in detail an F0/2 (sounding an octave below an original tone) subharmonic vibratory pattern produced in a normal larynx. Simultaneous electroglottographic and photoglottographic measurements reveal two different open phases within a subharmonic cycle- the first shorter with a simple shape, the second longer with a shape containing a "ripple." Such parameters as the large open quotient (ca. 0.8) and the high airflow values (ca. 1000 cm3/s) distinguish this phonation from the vocal fry (pulse) register. Using an electronic divider to track the subharmonic frequency, a method has been developed to observe the subharmonic vibration of the vocal folds stroboscopically. The stroboscopic visualization reveals an unusual mucosal movement during the "ripple," characterized by an opening movement of the upper margins, which interrupts the closing movement of the vocal folds. An explanation is offered that this vibratory pattern arises as a consequence of detuning of the usually identical frequencies of the dominant modes of the vocal folds, with 3:2 entrainment replacing the normal 1:1 pattern.

Measurement of formant frequencies and bandwidths in singing.

That singers under certain circumstances adjust the articulation of the vocal tract (formant tuning) to enhance acoustic output is both apparent from measurements and understood in theory. The precise effect of a formant on an approaching (retreating) harmonic as the latter varies in frequency during actual singing, however, is difficult to isolate. In this study variations in amplitude of radiated sound components as well as supraglottal and subglottal (esophageal) pressures accompanying the vibrato-related sweep of voice harmonics were used as a basis for estimating the effective center frequencies and bandwidths of the first and second formants.