The evolution of the syrinx: An acoustic theory.

The unique avian vocal organ, the syrinx, is located at the caudal end of the trachea. Although a larynx is also present at the opposite end, birds phonate only with the syrinx. Why only birds evolved a novel sound source at this location remains unknown, and hypotheses about its origin are largely untested. Here, we test the hypothesis that the syrinx constitutes a biomechanical advantage for sound production over the larynx with combined theoretical and experimental approaches. We investigated whether the position of a sound source within the respiratory tract affects acoustic features of the vocal output, including fundamental frequency and efficiency of conversion from aerodynamic energy to sound. Theoretical data and measurements in three bird species suggest that sound frequency is influenced by the interaction between sound source and vocal tract. A physical model and a computational simulation also indicate that a sound source in a syringeal position produces sound with greater efficiency. Interestingly, the interactions between sound source and vocal tract differed between species, suggesting that the syringeal sound source is optimized for its position in the respiratory tract. These results provide compelling evidence that strong selective pressures for high vocal efficiency may have been a major driving force in the evolution of the syrinx. The longer trachea of birds compared to other tetrapods made them likely predisposed for the evolution of a syrinx. A long vocal tract downstream from the sound source improves efficiency by facilitating the tuning between fundamental frequency and the first vocal tract resonance.

Vocalization with semi-occluded airways is favorable for optimizing sound production.

Vocalization in mammals, birds, reptiles, and amphibians occurs with airways that have wide openings to free-space for efficient sound radiation, but sound is also produced with occluded or semi-occluded airways that have small openings to free-space. It is hypothesized that pressures produced inside the airway with semi-occluded vocalizations have an overall widening effect on the airway. This overall widening then provides more opportunity to produce wide-narrow contrasts along the airway for variation in sound quality and loudness. For human vocalization described here, special emphasis is placed on the epilaryngeal airway, which can be adjusted for optimal aerodynamic power transfer and for optimal acoustic source-airway interaction. The methodology is three-fold, (1) geometric measurement of airway dimensions from CT scans, (2) aerodynamic and acoustic impedance calculation of the airways, and (3) simulation of acoustic signals with a self-oscillating computational model of the sound source and wave propagation.

Regulation of laryngeal resistance and maximum power transfer with semi-occluded airway vocalization.

Steady airflow resistances in semi-occluded airways as well as acoustic impedances in vocalization are quantified from the lungs to the lips. For clinical and voice training applications, the primary focus is on two airway conditions, an oral semi-occlusion and a semi-occlusion above the vocal folds. Laryngeal airflow resistance is divided into glottal airflow resistance and epilaryngeal airway resistance. Maximum aerodynamic power is transferred to the vocal tract if the glottal airflow resistance is reduced while the epilaryngeal airway resistance is increased. A semi-occlusion at the lips helps to set up this condition. For the acoustic power transfer, the epilaryngeal airway also serves to match the impedance of the source to the impedance of the vocal tract.

Analysis of Glottal Inverse Filtering in the Presence of Source-Filter Interaction.

The validity of glottal inverse filtering (GIF) to obtain a glottal flow waveform from radiated pressure signal in the presence and absence of source-filter interaction was studied systematically. A driven vocal fold surface model of vocal fold vibration was used to generate source signals. A one-dimensional wave reflection algorithm was used to solve for acoustic pressures in the vocal tract. Several test signals were generated with and without source-filter interaction at various fundamental frequencies and vowels. Linear Predictive Coding (LPC), Quasi Closed Phase (QCP), and Quadratic Programming (QPR) based algorithms, along with supraglottal impulse response, were used to inverse filter the radiated pressure signals to obtain the glottal flow pulses. The accuracy of each algorithm was tested for its recovery of maximum flow declination rate (MFDR), peak glottal flow, open phase ripple factor, closed phase ripple factor, and mean squared error. The algorithms were also tested for their absolute relative errors of the Normalized Amplitude Quotient, the Quasi-Open Quotient, and the Harmonic Richness Factor. The results indicated that the mean squared error decreased with increase in source-filter interaction level suggesting that the inverse filtering algorithms perform better in the presence of source-filter interaction. All glottal inverse filtering algorithms predicted the open phase ripple factor better than the closed phase ripple factor of a glottal flow waveform, irrespective of the source-filter interaction level. Major prediction errors occurred in the estimation of the closed phase ripple factor, MFDR, peak glottal flow, normalized amplitude quotient, and Quasi-Open Quotient. Feedback-related nonlinearity (source-filter interaction) affected the recovered signal primarily when f o was well below the first formant frequency of a vowel. The prediction error increased when f o was close to the first formant frequency due to the difficulty of estimating the precise value of resonance frequencies, which was exacerbated by nonlinear kinetic losses in the vocal tract.

A computational study of depth of vibration into vocal fold tissues.

The effective depth of vocal fold vibration is self-regulated and generally not known a priori in vocalization. In this study, the effective depth was quantified systematically under various phonatory conditions using a fiber-gel finite element vocal fold model. The horizontal and vertical excursions of each finite element nodal point trajectory were recorded to compute trajectory areas. The extent of vibration was then studied based on the variation of trajectory radii as a function of depth in several coronal sections along the anterior-posterior direction. The results suggested that the vocal fold nodal trajectory excursions decrease systematically as a function of depth but are affected by the layered structure of the vocal folds. The effective depth of vibration was found to range between 15 and 55% of the total anatomical depth across all phonatory conditions. The nodal trajectories from the current study were compared qualitatively with the results from excised human hemi-larynx experiments published in Döllinger and Berry [(2006). J. Voice. 20(3), 401-413]. An estimate of the effective mass of a one-mass vocal fold model was also computed based on the effective depth of vibration observed in this study under various phonatory conditions.

Radiation efficiency for long-range vocal communication in mammals and birds.

Long-distance vocal communication by birds and mammals, including humans, is facilitated largely by radiation efficiency from the mouth or beak. Here, this efficiency is defined and quantified. It depends on frequency content of vocalization, mouth opening, head and upper body geometry, and directionality. Each of these factors is described mathematically with a piston-in-a-sphere model. While this model is considered a classic, never before has the high frequency solution been applied in detail to vocalization. Results indicate that frequency content in the 1-50 kHz range can be radiated with nearly 100% efficiency if a reactance peak in the radiation impedance is utilized with adjustments of head size, mouth opening, and beam direction. Without these adjustments, radiation efficiency is generally below 1%, especially in human speech where a high fundamental frequency is a disadvantage for intelligibility. Thus, two distinct modes of vocal communication are identified, (1) short range with optimized information transfer and (2) long range with maximum efficiency for release of acoustic power.

WHERE HAS ALL THE POWER GONE? ENERGY PRODUCTION AND LOSS IN VOCALIZATION.

Human voice production for speech is an inefficient process in terms of energy expended to produce acoustic output. A traditional measure of vocal efficiency relates acoustic power radiated from the mouth to aerodynamic power produced in the trachea. This efficiency ranges between 0.001 % and 1.0 % in speech-like vocalization. Simplified Navier-Stokes equations for non-steady compressible airflow from trachea to lips were used to calculate steady aerodynamic power, acoustic power, and combined total power at seven strategic locations along the airway. A portion of the airway was allowed to collapse to produce self-sustained oscillation for sound production. A conversion efficiency, defined as acoustic power generated in the glottis to aerodynamic power dissipated, was found to be on the order of 10%, but wall vibration, air viscosity, and kinetic pressure losses consumed almost all of that power. This sound, reflected back and forth in the airway, was dissipated at a level on the order of 99.9 %.

Acoustic factors affecting the dynamic range of a choir.

Based on the assumption that individual sound intensities of singers are incoherent and add linearly to produce a combined choir intensity, a model of a voice range profile of a choir is produced. It is shown that this model predicts six distinct levels of choir dynamics (pp p mp mf f ff) over two octaves of fundamental frequency in a choir section. The levels are 3-6 dB apart, depending on the individual voice range profiles of the singers. Overall choir size has no effect on dynamic range, unless the size is varied dynamically by not all singers singing all the time. For a non-homogeneousn group of singers, a few loud voices dominate ff if everyone sings, while pp is not achieved effectively without suppressing all voices that cannot sing soft. Furthermore, the dynamic range can be significantly limited when choral blend for loudness is imposed on a non-homogeneous choir.

Comparison of a fiber-gel finite element model of vocal fold vibration to a transversely isotropic stiffness model.

A fiber-gel vocal fold model is compared to a transversely isotropic stiffness model in terms of normal mode vibration. The fiber-gel finite element model (FG-FEM) consists of a series of gel slices, each with a two-dimensional finite element mesh, in a plane transverse to the tissue fibers. The gel slices are coupled with fibers under tension in the anterior-posterior dimension. No vibrational displacement in the fiber-length direction is allowed, resulting in a plane strain state. This is consistent with the assumption of transverse displacement of a simple string, offering a wide range of natural frequencies (well into the kHz region) with variable tension. For low frequencies, the results compare favorably with the natural frequencies of a transversely isotropic elastic stiffness model (TISM) in which the shear modulus in the longitudinal plane is used to approximate the effect of fiber tension. For high frequencies, however, the natural frequencies do not approach the string mode frequencies unless plane strain is imposed on the TISM model. The simplifying assumption of plane strain, as well as the use of analytical closed-form shape functions, allow for substantial savings in computational time, which is important in clinical and exploratory applications of the FG-FEM model.

Sensitivity of odd-harmonic amplitudes to open quotient and skewing quotient in glottal airflow.

It is well known that a half-sinusoid has no odd harmonics other than the fundamental. If glottal flow in phonation were to approximate this exact waveshape, which is generally unlikely, some misperception of pitch and loss of vowel intelligibility would occur. The sensitivity of the glottal waveshape to this special shape is explored by systematically varying two parameters, open quotient and skewing quotient. Mild asymmetry (open quotient below 0.45 or above 0.55 and/or skewing quotient greater than 2.0) equalizes the odd-even harmonic series. Singers and speakers avoid the exact symmetry by skewing the flow pulse with source-filter interaction.

Bi-stable vocal fold adduction: a mechanism of modal-falsetto register shifts and mixed registration.

The origin of vocal registers has generally been attributed to differential activation of cricothyroid and thyroarytenoid muscles in the larynx. Register shifts, however, have also been shown to be affected by glottal pressures exerted on vocal fold surfaces, which can change with loudness, pitch, and vowel. Here it is shown computationally and with empirical data that intraglottal pressures can change abruptly when glottal adductory geometry is changed relatively smoothly from convergent to divergent. An intermediate shape between large convergence and large divergence, namely, a nearly rectangular glottal shape with almost parallel vocal fold surfaces, is associated with mixed registration. It can be less stable than either of the highly angular shapes unless transglottal pressure is reduced and upper stiffness of vocal fold tissues is balanced with lower stiffness. This intermediate state of adduction is desirable because it leads to a low phonation threshold pressure with moderate vocal fold collision. Achieving mixed registration consistently across wide ranges of F0, lung pressure, and vocal tract shapes appears to be a balancing act of coordinating laryngeal muscle activation with vocal tract pressures. Surprisingly, a large transglottal pressure is not facilitative in this process, exacerbating the bi-stable condition and the associated register contrast.

The accuracy of a voice vote.

The accuracy of a voice vote was addressed by systematically varying group size, individual voter loudness, and words that are typically used to express agreement or disagreement. Five judges rated the loudness of two competing groups in A-B comparison tasks. Acoustic analysis was performed to determine the sound energy level of each word uttered by each group. Results showed that individual voter differences in energy level can grossly alter group loudness and bias the vote. Unless some control is imposed on the sound level of individual voters, it is difficult to establish even a two-thirds majority, much less a simple majority. There is no symmetry in the bias created by unequal sound production of individuals. Soft voices do not bias the group loudness much, but loud voices do. The phonetic balance of the two words chosen (e.g., "yea" and "nay" as opposed to "aye" and "no") seems to be less of an issue.

Benchmarks for time-domain simulation of sound propagation in soft-walled airways: steady configurations.

Time-domain computer simulation of sound production in airways is a widely used tool, both for research and synthetic speech production technology. Speed of computation is generally the rationale for one-dimensional approaches to sound propagation and radiation. Transmission line and wave-reflection (scattering) algorithms are used to produce formant frequencies and bandwidths for arbitrarily shaped airways. Some benchmark graphs and tables are provided for formant frequencies and bandwidth calculations based on specific mathematical terms in the one-dimensional Navier-Stokes equation. Some rules are provided here for temporal and spatial discretization in terms of desired accuracy and stability of the solution. Kinetic losses, which have been difficult to quantify in frequency-domain simulations, are quantified here on the basis of the measurements of Scherer, Torkaman, Kucinschi, and Afjeh [(2010). J. Acoust. Soc. Am. 128(2), 828-838].

Simulation of Multiple Source Vocalization in the Larynx: How True Folds, False Folds, and Aryepiglottic Folds May Interact.

PURPOSE: This study was a modest beginning to determine dominance and entrainment between three soft tissues in the larynx that can be set into flow-induced oscillation and act as sound sources. The hypothesis was that they interact as coupled oscillators with observable bifurcations as energy is exchanged between them. METHODOLOGY: The true vocal folds, the ventricular (false) folds, and the aryepiglottic folds were part of a soft-walled airway that produced airflow for sound production. The methodology was computational, based on a simplified Navier-Stokes solution of convective and compressible airflow in a variable-geometry airway. RESULTS: Three serially connected sources could all produce flow-induced self-oscillation with soft wall tissue and small cross-sectional area. When the glottal cross-sectional areas were similar, bifurcations such as subharmonics, delayed voice onset, and aphonia occurred in the coupled oscillations. CONCLUSIONS: Closely spaced sound sources in the larynx are highly interactive. They appear to entrain to the source that has the combined advantage of small cross-sectional glottal area and proximity to a downstream vocal tract that supports oscillation with acoustic inertance.

Biomechanics of sound production in high-pitched classical singing.

Voice production of humans and most mammals is governed by the MyoElastic-AeroDynamic (MEAD) principle, where an air stream is modulated by self-sustained vocal fold oscillation to generate audible air pressure fluctuations. An alternative mechanism is found in ultrasonic vocalizations of rodents, which are established by an aeroacoustic (AA) phenomenon without vibration of laryngeal tissue. Previously, some authors argued that high-pitched human vocalization is also produced by the AA principle. Here, we investigate the so-called "whistle register" voice production in nine professional female operatic sopranos singing a scale from C6 (≈ 1047 Hz) to G6 (≈ 1568 Hz). Super-high-speed videolaryngoscopy revealed vocal fold collision in all participants, with closed quotients from 30 to 73%. Computational modeling showed that the biomechanical requirements to produce such high-pitched voice would be an increased contraction of the cricothyroid muscle, vocal fold strain of about 50%, and high subglottal pressure. Our data suggest that high-pitched operatic soprano singing uses the MEAD mechanism. Consequently, the commonly used term "whistle register" does not reflect the physical principle of a whistle with regard to voice generation in high pitched classical singing.

Simulation of Vocal Loudness Regulation with Lung Pressure, Vocal Fold Adduction, and Source-Airway Interaction.

In speaking, shouting, and singing, vocal loudness is known to be regulated with lung pressure, but the degree to which vocal fold adduction and airway shape play a role in loudness control is less well known. When loudness is quantified in sones instead of sound pressure level (SPL), the regulatory mechanisms are even less obvious. Here it is shown computationally that loudness is insensitive to changes in SPL produced with variable adduction. A trade-off exists between a reduction in glottal flow amplitude and a flatter spectral slope. When the airway configuration is changed from a uniform tube to a "belt" or "call" shape, loudness can increase with a slight decrease in SPL. When the airway configuration is changed from a uniform tube to an operatic "ring" shape, loudness is increased with only a small increase in SPL.

An Introduction to the Science of Information Transfer, Uncertainty, and a Personal View of Organizational Intelligence.

An introduction to the concepts of entropy, information transfer, and the uncertainty in transfer of information is given. The specific target is information transfer in voice and speech. The entropies of a square wave, a sinusoid, and a sawtooth are calculated because these waveshapes approximate information carriers in vocalization. This is followed by a less-known concept that the increase of organizational intelligence gleaned from physical systems is proportional to the gradient of entropy. That leads to a multi-dimensional interpretation of diversity. Finally, a personal meta-physical extension is made to a universal reservoir of intelligence from which species can draw to advance civilizations.

Vocal Demands of Musical Theatre Rehearsals: A Dosimetry Study.

OBJECTIVE: To investigate singers' vocal load by documenting three types of vocal doses (time, cycle, and distance doses) and sound pressure levels during the four phases of rehearsal and how the vocal doses vary between singers across rehearsals in the musical Nine, written by Maury Yeston. METHODS/DESIGN: Five student-singers participating in the musical Nine gave informed consent to participate in the study. All five participants were assigned female at birth and female-identifying individuals. They attached a KayPENTAX APM 3300 dosimeter sensor to their lower neck and wore the accelerometer during four three-hour rehearsals throughout the rehearsal process (the music learning phase, the choreography learning phase, the blocking learning phase, and the dress rehearsal) of the musical. The dosimeter records neck vibrations at a rate of 20 samples per second. but it does not record linguistic content. RESULTS: A dosimetric analysis of five student singers identified variability in voice production throughout the rehearsal process. According to the dosimetry findings, singers employed extensive low-frequency voicing below the first passaggio, with belting and mixed vocal strategies as the predominant stylistic choices when performing in Nine. Additionally, the singers used an occasional head voice effect at specific moments. The roles of Carla, Saraghina, La Fleur, and Ensemble One and Two required specific vocal ranges due to the musical score. CONCLUSIONS: Researchers have yet to establish a safe baseline vocal dose for singers. The vocal dose is affected by many factors, such as duration of phonation, frequency range, SPL, and styles of vocalism required by the score. Louder and heavier vocalization produces larger distance doses, representing the cumulative load placed on vibrating tissue. The cycle dose, distance dose, and SPL reported in this study varied within and between singers. The phonation density graphs show this variability and the low tessitura required by the score. Time doses ranged from 4% to 7% of rehearsal time; this short dose suggests that the rehearsals provided healthy conditions for the successful rehearsal process with efficient attention to the vocalization of a score that requires heavy vocal styles, including belting. While the rehearsal pace was not alarming, the demands of the score alone may prove to be much greater than the vocal dose reported through the rehearsal. Further studies are needed to establish the overall dose of each Broadway role to serve as parameters for vocal pacing and voice care.

Integrative Insights into the Myoelastic-Aerodynamic Theory and Acoustics of Phonation. Scientific Tribute to Donald G. Miller.

In this tribute article to D.G. Miller, we review some historical and recent contributions to understanding the myoelastic-aerodynamic (MEAD) theory of phonation and the related acoustic phenomena in subglottal and vocal tract. At the time of the formulation of MEAD by van den Berg in late 1950s, it was assumed that vocal fold oscillations are self-sustained thanks to increased subglottal pressure pushing the glottis to open and decreased subglottal pressure allowing the glottis to close. In vivo measurements of subglottal pressures during phonation invalidated these assumptions, however, and showed that at low fundamental frequencies subglottal pressure rather tends to reach a maximum value at the beginning of glottal closure and then exhibits damped oscillations. These events can be interpreted as transient acoustic resonance phenomena in the subglottal tract that are triggered by glottal closure. They are analogous to the transient acoustic phenomena seen in the vocal tract. Rather than subglottal pressure oscillations, a more efficient mechanism of transfer of aerodynamic energy to the vocal fold vibrations has been identified in the vertical phase differences (mucosal waves) making the glottal shape more convergent during glottis opening than during glottis closing. Along with other discoveries, these findings form the basis of our current understanding of MEAD.

Adhesion of a monolayer of fibroblast cells to fibronectin under sonic vibrations in a bioreactor.

OBJECTIVES: We examined cell adhesion to a surface under vibrational forces approximating those of phonation. METHODS: A monolayer of human fibroblast cells was seeded on a fibronectin-coated glass coverslip, which was attached to either the rotating part or the stationary part of a rheometer-bioreactor. The temperature, humidity, carbon dioxide level, nutrients, and cell seeding density were controlled. The cell density was on the order of 1,000 to 5,000 cells per square millimeter. Target stresses above 1 kPa at an oscillatory frequency of 100 Hz were chosen to reflect conditions of vocal fold tissue vibration. RESULTS: Fibronectin coating provided enough adhesion to support at least 2 kPa of oscillating stress, but only about 0.1 kPa of steady rotational shear. For stresses exceeding those limits, the cells were not able to adhere to the thin film of fibronectin. CONCLUSIONS: Cells will adhere to a planar surface under stresses typical of phonation, which provide a more stringent test than adherence in a 3-dimensional matrix. The density of cell seeding on the coverslip played a role in cell-extracellular matrix adhesion, in that the cells adhered to each other more than to the fibronectin coating when the cells were nearly confluent.