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.

Acoustical Theory of Vowel Modification Strategies in Belting.

Various authors have argued that belting is to be produced by "speech-like" sounds, with the first and second supraglottic vocal tract resonances (fR1 and fR2) at frequencies of the vowels determined by the lyrics to be sung. Acoustically, the hallmark of belting has been identified as a dominant second harmonic, possibly enhanced by first resonance tuning (fR1≈2fo). It is not clear how both these concepts - (a) phonating with "speech-like," unmodified vowels; and (b) producing a belting sound with a dominant second harmonic, typically enhanced by fR1 - can be upheld when singing across a singer's entire musical pitch range. For instance, anecdotal reports from pedagogues suggest that vowels with a low fR1, such as [i] or [u], might have to be modified considerably (by raising fR1) in order to phonate at higher pitches. These issues were systematically addressed in silico with respect to treble singing, using a linear source-filter voice production model. The dominant harmonic of the radiated spectrum was assessed in 12987 simulations, covering a parameter space of 37 fundamental frequencies (fo) across the musical pitch range from C3 to C6; 27 voice source spectral slope settings from -4 to -30 dB/octave; computed for 13 different IPA vowels. The results suggest that, for most unmodified vowels, the stereotypical belting sound characteristics with a dominant second harmonic can only be produced over a pitch range of about a musical fifth, centered at fo≈0.5fR1. In the [ɔ] and [ɑ] vowels, that range is extended to an octave, supported by a low second resonance. Data aggregation - considering the relative prevalence of vowels in American English - suggests that, historically, belting with fR1≈2fo was derived from speech, and that songs with an extended musical pitch range likely demand considerable vowel modification. We thus argue that - on acoustical grounds - the pedagogical commandment for belting with unmodified, "speech-like" vowels can not always be fulfilled.

Vocal Vibrato Characteristics in Historical and Contemporary Opera, Operetta, and Schlager.

OBJECTIVES/HYPOTHESIS: Vibrato is a core aesthetic element in singing. It varies considerably by both genre and era. Though studied extensively in Western classical singing over the years, there is a dearth of studies on vibrato in contemporary commercial music. In addressing this research gap, the objective of this study was to find and investigate common crossover song material from the opera, operetta, and Schlager singing styles from the historical early 20th to the contemporary 21st century epochs. STUDY DESIGN/METHODS: A total of 51 commercial recordings of two songs, "Es muss was Wunderbares sein" by Ralph Benatzky, and "Die ganze Welt ist himmelblau" by Robert Stolz, from "The White Horse Inn" ("Im weißen Rößl") were collected from opera, operetta, and Schlager singers. Each sample was annotated using Praat and analyzed in a custom Matlab- and Python-based algorithmic approach of singing voice separation and sine wave fitting novel to vibrato research. RESULTS: With respect to vibrato rate and extent, the three most notable findings were that (1) fo and vibrato were inherently connected; (2) Schlager, as a historical aesthetic category, has unique vibrato characteristics, with higher overall rate and lower overall extent; and (3) fo and vibrato extent varied over time based on the historical or contemporary recording year for each genre. CONCLUSIONS: Though these results should be interpreted with caution due to the limited sample size, conducting such acoustical analysis is relevant for voice pedagogy. This study sheds light on the complexity of vocal vibrato production physiology and acoustics while providing insight into various aesthetic choices when performing music of different genres and stylistic time periods. In the age of crossover singing training and commercially available recordings, this investigation reveals important distinctions regarding vocal vibrato across genres and eras that bear beneficial implications for singers and teachers of singing.

Voices in the ocean.

Toothed whales evolved a third way of making sounds similar to that of land mammals and birds.

Dynamic System Coupling in Voice Production.

Voice is a major means of communication for humans, non-human mammals and many other vertebrates like birds and anurans. The physical and physiological principles of voice production are described by two theories: the MyoElastic-AeroDynamic (MEAD) theory and the Source-Filter Theory (SFT). While MEAD employs a multiphysics approach to understand the motor control and dynamics of self-sustained vibration of vocal folds or analogous tissues, SFT predominantly uses acoustics to understand spectral changes of the source via linear propagation through the vocal tract. Because the two theories focus on different aspects of voice production, they are often applied distinctly in specific areas of science and engineering. Here, we argue that the MEAD and the SFT are linked integral aspects of a holistic theory of voice production, describing a dynamically coupled system. The aim of this manuscript is to provide a comprehensive review of both the MEAD and the source-filter theory with its nonlinear extension, the latter of which suggests a number of conceptual similarities to sound production in brass instruments. We discuss the application of both theories to voice production of humans as well as of animals. An appraisal of voice production in the light of non-linear dynamics supports the notion that voice production can best be described with a systems view, considering coupled systems rather than isolated contributions of individual sub-systems.

Time-Synchronized MRI-Assessment of Respiratory Apparatus Subsystems-A Feasibility Study.

The thorax (TH), the thoracic diaphragm (TD), and the abdominal wall (AW) are three sub-systems of the respiratory apparatus whose displacement motion has been well studied with the use of magnetic resonance imaging (MRI). Another sub-system, which has however received less research attention with respect to breathing, is the pelvic floor (PF). In particular, there is no study that has investigated the displacement of all four sub-systems simultaneously. Addressing this issue, it was the purpose of this feasibility study to establish a data acquisition paradigm for time-synchronous quantitative analysis of dynamic MRI data from these four major contributors to respiration and phonation (TH, TD, AW, and PF). Three healthy females were asked to breathe in and out forcefully while being recorded in a 1.5-Tesla whole body MR-scanner. Spanning a sequence of 15.12 seconds, 40 MRI data frames were acquired. Each data frame contained two slices, simultaneously documenting the mid-sagittal (TH, TD, PF) and transversal (AW) planes. The displacement motion of the four anatomical structures of interest was documented using kymographic analysis, resulting in time-varying calibrated structure displacement data. After computing the fundamental frequency of the cyclical breathing motion, the phase offsets of the TH, PF, and AW with respect to the TD were computed. Data analysis revealed three fundamentally different displacement patterns. Total structure displacement was in the range of 0.94 cm (TH) to 4.27 cm (TD). Phase delays of up to 90∘ (i.e., a quarter of a breathing cycle) between different structures were found. Motion offsets in the range of -28.30∘ to 14.90∘ were computed for the PF with respect to the TD. The diversity of results in only three investigated participants suggests a variety of possible breathing strategies, warranting further research.

Computer simulation of vocal tract resonance tuning strategies with respect to fundamental frequency and voice source spectral slope in singing.

A well-known concept of singing voice pedagogy is "formant tuning," where the lowest two vocal tract resonances ( fR1, fR2) are systematically tuned to harmonics of the laryngeal voice source to maximize the level of radiated sound. A comprehensive evaluation of this resonance tuning concept is still needed. Here, the effect of fR1, fR2 variation was systematically evaluated in silico across the entire fundamental frequency range of classical singing for three voice source characteristics with spectral slopes of -6, -12, and -18 dB/octave. Respective vocal tract transfer functions were generated with a previously introduced low-dimensional computational model, and resultant radiated sound levels were expressed in dB(A). Two distinct strategies for optimized sound output emerged for low vs high voices. At low pitches, spectral slope was the predominant factor for sound level increase, and resonance tuning only had a marginal effect. In contrast, resonance tuning strategies became more prevalent and voice source strength played an increasingly marginal role as fundamental frequency increased to the upper limits of the soprano range. This suggests that different voice classes (e.g., low male vs high female) likely have fundamentally different strategies for optimizing sound output, which has fundamental implications for pedagogical practice.

Evolutionary loss of complexity in human vocal anatomy as an adaptation for speech.

Human speech production obeys the same acoustic principles as vocal production in other animals but has distinctive features: A stable vocal source is filtered by rapidly changing formant frequencies. To understand speech evolution, we examined a wide range of primates, combining observations of phonation with mathematical modeling. We found that source stability relies upon simplifications in laryngeal anatomy, specifically the loss of air sacs and vocal membranes. We conclude that the evolutionary loss of vocal membranes allows human speech to mostly avoid the spontaneous nonlinear phenomena and acoustic chaos common in other primate vocalizations. This loss allows our larynx to produce stable, harmonic-rich phonation, ideally highlighting formant changes that convey most phonetic information. Paradoxically, the increased complexity of human spoken language thus followed simplification of our laryngeal anatomy.

In-Person or Virtual? - Assessing the Impact of COVID-19 on the Teaching Habits of Voice Pedagogues.

The social distancing measures implemented world-wide in the wake of the novel Coronavirus (COVID-19) crisis have forced voice pedagogues to alter their teaching habits, likely shifting from customary in-person teaching to virtual teaching. An online survey, distributed world-wide in April/May 2020, investigated how singing voice pedagogues were impacted by the COVID-19 crisis. The collected responses from 387 survey participants suggest that, overall, voice teachers were only moderately satisfied with having to teach virtually, indicating that virtual voice teaching is not a sufficient replacement for in-person teaching. The participants indicated that during virtual teaching the singing voice can be assessed relatively well through features which provide both acoustic and visual clues. In contrast, depending on utilized technology, it may be harder to judge those aspects of the singing voice that are solely defined acoustically, such as dynamic range and spectral composition. This may be explained by limitations imposed by "out of the box" technology for online communication, which is typically optimized for speech instead of singing. This calls for better information on technological solutions for virtual voice teaching.

Global Inventory and Similarity Rating of Singing Voice Assessment Terms Used at English Speaking Academic Institutions.

The choice of terms to describe and assess the singing voice is an essential part of vocal pedagogy. However, previous work suggested that singing terminology used in academia may be somewhat ambiguous. To address this issue, the authors a) compiled a comprehensive inventory of singing voice assessment terms used by English-speaking academic institutions worldwide and b) with the help of 22 highly experienced singing voice teachers, grouped the most prevalent terms based on their conceptual similarity. Only about a fifth of all targeted institutions provided materials and information online. Overall, a total of 292 different terms were found in the 64 available sources. This surprisingly large number of terms could be reduced by approximately 61% through lexical grouping. In the resulting data set, only 24 of the 114 terms occurred in at least 20% of the online sources, suggesting a rather low current density of information as well as little to no systematic and coordinated use of terms across institutions. The singing voice expert's similarity rating of the 24 most prevalent terms revealed a non-uniform distribution, suggesting that only some of these terms can be used interchangeably. Overall, these findings hint at the underlying complexity of voice assessment on a descriptive and qualitative level, highlighting the need for further research in this area.

Are source-filter interactions detectable in classical singing during vowel glides?

In recent studies, it has been assumed that vocal tract formants (Fn) and the voice source could interact. However, there are only few studies analyzing this assumption in vivo. Here, the vowel transition /i/-/a/-/u/-/i/ of 12 professional classical singers (6 females, 6 males) when phonating on the pitch D4 [fundamental frequency (ƒo) ca. 294 Hz] were analyzed using transnasal high speed videoendoscopy (20.000 fps), electroglottography (EGG), and audio recordings. Fn data were calculated using a cepstral method. Source-filter interaction candidates (SFICs) were determined by (a) algorithmic detection of major intersections of Fn/nƒo and (b) perceptual assessment of the EGG signal. Although the open quotient showed some increase for the /i-a/ and /u-i/ transitions, there were no clear effects at the expected Fn/nƒo intersections. In contrast, ƒo adjustments and changes in the phonovibrogram occurred at perceptually derived SFICs, suggesting level-two interactions. In some cases, these were constituted by intersections between higher nƒo and Fn. The presented data partially corroborates that vowel transitions may result in level-two interactions also in professional singers. However, the lack of systematically detectable effects suggests either the absence of a strong interaction or existence of confounding factors, which may potentially counterbalance the level-two-interactions.

Effect of Ventricular Folds on Vocalization Fundamental Frequency in Domestic Pigs (Sus scrofa domesticus).

This study investigates the effect of the ventricular folds on fundamental frequency (fo) in the voice production of domestic pigs (Sus scrofa domesticus). The excised larynges of six subadult pigs were phonated in two preparation stages, with the ventricular folds present (PS1) and removed (PS2). Vocal fold resonances were tested with a laser vibrometer, and a four-mass computational model was created. Highly significant fo differences were found between PS1 and PS2 (means at 93.7 and 409.3 Hz, respectively). Two tissue resonances were found at 115 Hz and 250-290 Hz. The computational model had unique solutions for abducted and adducted ventricular folds at about 150 and 400 Hz, roughly matching the fo measured ex vivo for PS1 and PS2. The differing fo encountered across preparation stages PS1 and PS2 is explained by distinct activation of either a high or a low eigenfrequency mode, depending on the engagement of the ventricular folds. The inability of the investigated larynges to vibrate at frequencies below 250 Hz in PS2 suggests that in vivo low-frequency calls of domestic pigs (pre-eminently grunts) are likely produced with engaged ventricular folds. Allometric comparison suggests that the special, mechanically coupled "double oscillator" has evolved to prevent signaling disadvantages. Given these traits, the porcine larynx might - apart from special applications relating to the involvement of ventricular folds - not be an ideal candidate for emulating human voice production in excised larynx experimentation.

Performance Evaluation of Subharmonic-to-Harmonic Ratio (SHR) Computation.

Subharmonics are an important class of voice signals, relevant for speech, pathological voice, singing, and animal bioacoustics. They arise from special cases of amplitude (AM) or frequency modulation (FM) of the time-domain signal. Surprisingly, to date there is only one open source subharmonics detector available to the scientific community: Sun's subharmonic-to-harmonic ratio (SHR). Here, this algorithm was subjected to a formal evaluation with two data sets of synthesized and empirical speech samples. Both data sets consisted of electroglottographic (EGG) signals, ie, a physiological correlate of vocal fold oscillation that bypasses vocal tract acoustics. Data Set I contained 2560 synthesized EGG signals with varying degrees of AM and FM, fundamental frequency (fo), periodicity, and signal-to-noise ratio (SNR). Data Set II was made up of 25 EGG samples extracted from the CMU Arctic speech data base. For a "ground truth" of subharmonicity, these samples were manually annotated by a group of five external experts. Analysis of the synthesized data suggested that the SHR metric is relatively robust as long as the subharmonic modulation extent is below 0.35 and 0.7 for the FM and AM scenarios, respectively. In the CMU Arctic speech data samples, the SHR analysis reached a maximum sensitivity of about 87% at a specificity of over 90%, but only for adaptive algorithm parameter settings. In contrast, the algorithm's default parameter settings could only successfully classify about 9% of all subharmonic instances. The SHR is a useful metric for assessing the degree of subharmonics contained in voice signals, but only at adaptive parameter settings. In particular, the frequency ceiling should be set to five times the highest fo, and the frame length to at least five times the largest fundamental period of the analyzed signal. For subharmonic classification a threshold of SHR  ≥  0.01 is recommended.

Electroglottography - An Update.

Electroglottography (EGG) is a low-cost, noninvasive technology for measuring changes of relative vocal fold contact area during laryngeal voice production. EGG was introduced about 60 years ago and has gone through a "golden era" of increased scientific attention in the late 1980s and early 90s. During that period, four eminent review papers were written. Here, an update to these reviews is given, recapitulating some earlier landmark contributions and documenting noteworthy developments during the past 25 years. After presenting an algorithmic bibliographic analysis, some methodological aspects pertaining to measurement technology, qualitative and quantitative analysis, and respective interpretation are discussed. In particular, the interpretation of landmarks in the (first derivative of the) EGG waveform is critically examined. It is argued that because of inferior-superior and anterior-posterior phase differences of vocal fold vibration, vocal fold (de)contacting does not occur instantaneously, but over an interval of time. For this reason, instants of vocal fold closing and opening cannot be resolved exactly from the EGG signal. Consequently, any quantitative analysis parameter relying on the determination of (de)contacting events (such as the EGG contact quotient) should be interpreted with care. Finally, recent developments are reviewed for the various fields of application of EGG, including basic voice science and voice production physiology, speech signal processing and classification, clinical practice including swallowing, phonetics, hearing sciences, psychology, singing, trumpet playing, and mammalian and avian bioacoustics. Overall, EGG has over the past six decades developed into a mature technology with a wide range of applications. However, due to current limitations, the full potential of the methodology has as yet not been fully exploited. Future development may occur on three levels: (a) rigorous validation of existent measurement approaches; (b) introduction and rigorous validation of novel quantitative and interpretative approaches; and (c) advancement of the measurement technology itself.

Fundamental Frequency Estimation of Low-quality Electroglottographic Signals.

Fundamental frequency (fo) is often estimated based on electroglottographic (EGG) signals. Because of the nature of the method, the quality of EGG signals may be impaired by certain features like amplitude or baseline drifts, mains hum, or noise. The potential adverse effects of these factors on fo estimation have to date not been investigated. Here, the performance of 13 algorithms for estimating fo was tested, based on 147 synthesized EGG signals with varying degrees of signal quality deterioration. Algorithm performance was assessed through the standard deviation σfo of the difference between known and estimated fo data, expressed in octaves. With very few exceptions, simulated mains hum, and amplitude and baseline drifts did not influence fo results, even though some algorithms consistently outperformed others. When increasing either cycle-to-cycle fo variation or the degree of subharmonics, the SIGMA algorithm had the best performance (max. σfo = 0.04). That algorithm was, however, more easily disturbed by typical EGG equipment noise, whereas the NDF and Praat's auto-correlation algorithms performed best in this category (σfo = 0.01). These results suggest that the algorithm for fo estimation of EGG signals needs to be selected specifically for each particular data set. Overall, estimated fo data should be interpreted with care.

Quantifying syringeal dynamics in vitro using electroglottography.

The complex and elaborate vocalizations uttered by many of the 10,000 extant bird species are considered a major driver in their evolutionary success, warranting study of the underlying mechanisms of vocal production. Additionally, birdsong has developed into a highly productive model system for vocal imitation learning and motor control, where, in contrast to humans, we have experimental access to the entire neuromechanical control loop. In human voice production, complex laryngeal geometry, vocal fold tissue properties, airflow and laryngeal musculature all interact to ultimately control vocal fold kinematics. Quantifying vocal fold kinematics is thus critical to understanding neuromechanical control of voiced sound production, but in vivo imaging of vocal fold kinematics in birds is experimentally challenging. Here, we adapted and tested electroglottography (EGG) as a novel tool for examining vocal fold kinematics in the avian vocal organ, the syrinx. We furthermore imaged and quantified syringeal kinematics in the pigeon (Columba livia) syrinx with unprecedented detail. Our results show that EGG signals predict (1) the relative amount of contact between the avian equivalent of vocal folds and (2) essential parameters describing vibratory kinematics, such as fundamental frequency, and timing of syringeal opening and closing events. As such, EGG provides novel opportunities for measuring syringeal vibratory kinematic parameters in vivo Furthermore, the opportunity for imaging syringeal vibratory kinematics from multiple planar views (horizontal and coronal) simultaneously promotes birds as an excellent model system for studying kinematics and control of voiced sound production in general, including in humans and other mammals.

Japanese macaque phonatory physiology.

Although the call repertoire and its communicative function are relatively well explored in Japanese macaques (Macaca fuscata), little empirical data are available on the physics and the physiology of this species' vocal production mechanism. Here, a 6 year old female Japanese macaque was trained to phonate under an operant conditioning paradigm. The resulting 'coo' calls and spontaneously uttered 'growl' and 'chirp' calls were recorded with sound pressure level (SPL) calibrated microphones and electroglottography (EGG), a non-invasive method for assessing the dynamics of phonation. A total of 448 calls were recorded, complemented by ex vivo recordings on an excised Japanese macaque larynx. In this novel multidimensional investigative paradigm, in vivo and ex vivo data were matched via comparable EGG waveforms. Subsequent analysis suggests that the vocal range (range of fundamental frequency and SPL) of the macaque was comparable to that of a 7-10 year old human, with the exception of low intensity chirps, the production of which may be facilitated by the species' vocal membranes. In coo calls, redundant control of fundamental frequency in relation to SPL was also comparable to that in humans. EGG data revealed that growls, coos and chirps were produced by distinct laryngeal vibratory mechanisms. EGG further suggested changes in the degree of vocal fold adduction in vivo, resulting in spectral variation within the emitted coo calls, ranging from 'breathy' (including aerodynamic noise components) to 'non-breathy'. This is again analogous to humans, corroborating the notion that phonation in humans and non-human primates is based on universal physical and physiological principles.

Publisher Correction: Acoustic allometry revisited: morphological determinants of fundamental frequency in primate vocal production.

A correction to this article has been published and is linked from the HTML and PDF versions of this paper. The error has been fixed in the paper.

Publisher Correction: Acoustic allometry revisited: morphological determinants of fundamental frequency in primate vocal production.

A correction to this article has been published and is linked from the HTML version of this paper. The error has not been fixed in the paper.

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.