What's special about human speech? A student exercise for comparing speech production between humans and chimpanzees.

Modern humans and chimpanzees share a common ancestor on the phylogenetic tree, yet chimpanzees do not spontaneously produce speech or speech sounds. The lab exercise presented in this paper was developed for undergraduate students in a course entitled "What's Special About Human Speech?" The exercise is based on acoustic analyses of the words "cup" and "papa" as spoken by Viki, a home-raised, speech-trained chimpanzee, as well as the words spoken by a human. The analyses allow students to relate differences in articulation and vocal abilities between Viki and humans to the known anatomical differences in their vocal systems. Anatomical and articulation differences between humans and Viki include (1) potential tongue movements, (2) presence or absence of laryngeal air sacs, (3) presence or absence of vocal membranes, and (4) exhalation vs inhalation during production.

The domestication of the larynx: The neural crest connection.

Wolves howl and dogs bark, both are able to produce variants of either vocalization, but we see a distinct difference in usage between wild and domesticate. Other domesticates also show distinct changes to their vocal output: domestic cats retain meows, a distinctly subadult trait in wildcats. Such differences in acoustic output are well-known, but the causal mechanisms remain little-studied. Potential links between domestication and vocal output are intriguing for multiple reasons, and offer a unique opportunity to explore a prominent hypothesis in domestication research: the neural crest/domestication syndrome hypothesis. This hypothesis suggests that in the early stages of domestication, selection for tame individuals decreased neural crest cell (NCCs) proliferation and migration, which led to a downregulation of the sympathetic arousal system, and hence reduced fear and reactive aggression. NCCs are a transitory stem cell population crucial during embryonic development that tie to diverse tissue types and organ systems. One of these neural-crest derived systems is the larynx, the main vocal source in mammals. We argue that this connection between NCCs and the larynx provides a powerful test of the predictions of the neural crest/domestication syndrome hypothesis, discriminating its predictions from those of other current hypotheses concerning domestication.

The partial upward migration of the laryngeal motor cortex: A window to the human brain evolution.

The pioneer cortical electrical stimulation studies of the last century did not explicitly mark the location of the human laryngeal motor cortex (LMC), but only the "vocalization area" in the lower half of the lateral motor cortex. In the final years of 2010́s, neuroimaging studies did demonstrate two human cortical laryngeal representations, located at the opposing ends of the orofacial motor zone, therefore termed dorsal (LMCd) and ventral laryngeal motor cortex (LMCv). Since then, there has been a continuing debate regarding the origin, function and evolutionary significance of these areas. The "local duplication model" posits that the LMCd evolved by a duplication of an adjacent region of the motor cortex. The "duplication and migration model" assumes that the dorsal LMCd arose by a duplication of motor regions related to vocalization, such as the ancestry LMC, followed by a migration into the orofacial region of the motor cortex. This paper reviews the basic arguments of these viewpoints and suggests a new explanation, declaring that the LMCd in man is rather induced through the division of the unitary LMC in nonhuman primates, upward shift and relocation of its motor part due to the disproportional growth of the head, face, mouth, lips, and tongue motor areas in the ventral part of the human motor homunculus. This explanation may be called "expansion-division and relocation model".

CT and MR anatomy of the larynx and hypopharynx.

Imaging of the larynx and hypopharynx is frequently requested to assess the extent of neoplasms beyond the field of view of endoscopic evaluation. The combination of optical and cross-sectional imaging allows tumors to be classified according to AJCC/UICC guidelines. A thorough understanding of laryngeal and hypopharyngeal anatomy is crucial to guide the radiological eye along the possible pathways of the spread of diseases and to guide differential diagnoses. Computed tomography (CT) has been the first cross-sectional imaging technique used to evaluate the larynx and hypopharynx; its spatial resolution combined with volumetric capability and the use of injectable contrast medium made CT the working horse in the assessment of neoplastic and inflammatory diseases. In the last two decades, magnetic resonance (MR) supported CT in the most challenging cases, when the optimal contrast resolution due to the multisequence portfolio is needed to assess the neoplastic involvement of laryngeal cartilages, paraglottic space(s), and extra laryngeal spread. The aim of this paper is to give a comprehensive radiological overview of larynx and hypopharynx complex anatomy, combining in vivo images, anatomical sections, and images of ex vivo specimens.

Striking a (vocal) chord: musical instruments as mnemonic devices when teaching the functional anatomy of the larynx.

Mnemonic devices are memory aids that make it easier to recall information and are widely used by students studying anatomy and physiology. Simple musical instruments and toys can serve as mnemonic devices for students learning the functional anatomy of the larynx: balloons can help learners understand and recall how sound is produced; tuning pegs can help learners understand how tension affects vocal pitch; fingers on a fretboard can help learners understand how pitch is further modulated; and a common coach's whistle can demonstrate how vocal volume is controlled. Using instruments and toys engages adult learners and helps them connect complex laryngeal anatomy with previous experiences.NEW & NOTEWORTHY Musical instruments and toys can be used as mnemonic devices to help students recall and understand the functional anatomy of voice production. The mnemonics can be implemented in a variety of classrooms and are flexible and engaging.

Homology and the evolution of vocal folds in the novel avian voice box.

The origin of novel traits, those that are not direct modifications of a pre-existing ancestral structure, remains a fundamental problem in evolutionary biology. For example, little is known about the evolutionary and developmental origins of the novel avian vocal organ, the syrinx. Located at the tracheobronchial junction, the syrinx is responsible for avian vocalization, but it is unclear whether avian vocal folds are homologous to the laryngeal vocal folds in other tetrapods or convergently evolved. Here, we identify a core developmental program involved in avian vocal fold formation and infer the morphology of the syrinx of the ancestor of modern birds. We find that this ancestral syrinx had paired sound sources induced by a conserved developmental pathway and show that shifts in these signals correlate with syringeal diversification. We show that, despite being derived from different developmental tissues, vocal folds in the syrinx and larynx have similar tissue composition and are established through a strikingly similar developmental program, indicating that co-option of an ancestral developmental program facilitated the origin of vocal folds in the avian syrinx.

An open-source toolbox for measuring vocal tract shape from real-time magnetic resonance images.

Real-time magnetic resonance imaging (rtMRI) is a technique that provides high-contrast videographic data of human anatomy in motion. Applied to the vocal tract, it is a powerful method for capturing the dynamics of speech and other vocal behaviours by imaging structures internal to the mouth and throat. These images provide a means of studying the physiological basis for speech, singing, expressions of emotion, and swallowing that are otherwise not accessible for external observation. However, taking quantitative measurements from these images is notoriously difficult. We introduce a signal processing pipeline that produces outlines of the vocal tract from the lips to the larynx as a quantification of the dynamic morphology of the vocal tract. Our approach performs simple tissue classification, but constrained to a researcher-specified region of interest. This combination facilitates feature extraction while retaining the domain-specific expertise of a human analyst. We demonstrate that this pipeline generalises well across datasets covering behaviours such as speech, vocal size exaggeration, laughter, and whistling, as well as producing reliable outcomes across analysts, particularly among users with domain-specific expertise. With this article, we make this pipeline available for immediate use by the research community, and further suggest that it may contribute to the continued development of fully automated methods based on deep learning algorithms.

Morphology and computed tomography of the larynx in the white-eared opossum (Didelphis albiventris).

The white-eared opossum (Didelphis albiventris) is a Neotropical marsupial that occurs in the Brazilian territory. The larynx is an important organ of vocalization in mammals, although, other laryngeal functions are more fundamental for survival of mammals than phonation. The anatomical knowledge of respiratory structures is pivotal for a better understanding of the species. Thus, this study aimed to examine the larynx of the white-eared opossum by gross anatomy, computed tomography and histological description. For this, 10 adult (six females and four males) white-eared opossums (D. albiventris) were used. The white-eared opossum larynx was formed by epiglottis, thyroid, cricoids and arytenoid cartilages and a corniculate process. There is a similarity between the larynx of this marsupial and those reported in other wild mammals, regarding the number of cartilages and their location. Histologically, the epiglottis consisted of elastic cartilage and thyroid, cricoid and arytenoid cartilages were composed of hyaline cartilage. The epiglottis protruded rostrally into the nasal part of the pharynx, above the soft palate, and this is probably a marsupial characteristic as the fact that the thyroid and cricoid cartilages were ventrally fused. The hyoid apparatus was similar to that of other animals, with the same bony constitution, but with a greatly reduced stylohyoid bone. Histologically, the larynx was similar to those of other species such as koala, armadillo, crab-eating foxes and giant anteaters. The knowledge of the larynx morphology is important for the anatomical features of the species and clinical and surgical procedures, such as endotracheal intubation.

Morphology of the syrinx of three species of birds from Brazilian cerrado (Psittacara leucophthalmus, Rhynchotus rufescens and Cariama cristata): Gross anatomy and light microscopy study.

The aim of this study was to describe the morphology of the trachea and syrinx at macroscopic and light microscopy levels of three species of birds from different orders that inhabit the Brazilian cerrado. For that, five adult specimens (three males and two females of each species) of white-eyed parakeet (Psittacara leucophthalmus), red-winged tinamou (Rhynchotus rufescens) and red-legged seriema (Cariama cristata) were used. The trachea and syrinx of the birds were collected and destined for anatomical and histological studies. The trachea of the studied birds presented an elongated path and originated in the larynx and extended caudally to the syrinx. No sexual dimorphism was observed in the syrinx of the studied species, probably because it is associated with their song, which is very similar between males and females of these species. The findings of this study allowed us to classify the syrinx as tracheal in the white-eyed parakeet and tracheobronchial in the red-winged tinamou and red-legged seriema. In general, the morphological features of the trachea and syrinx were similar to those described for other species of birds, such as the presence of intrinsic and extrinsic syringeal muscles, and the lateral and medial tympaniform membranes, which would represent important anatomical structures in sound production through vibration during expiration and eventual inspiration. The morphological structure of the syrinx in the three avian species of the Brazilian cerrado is consistent with the ability of these avian species to perform a potential vocalization, especially the red-legged seriema that emits characteristic sounds very loud and can carry several kilometres.

Amount and spatial arrangement of muscle fibers in the human laryngeal Musculus ventricularis.

Textbooks and atlases of human macroscopic and microscopic anatomy of the larynx generally provide, if at all, only sparse information on the laryngeal Musculus ventricularis. However, several studies indicate that this muscle takes over the function of vestibular (ventricular) fold phonation after denervation of the Musculus vocalis. In the present study, 29 laryngeal specimens were coronally dissected at different levels, i.e. the anterior (L1), middle (L2), and posterior third of the vestibular fold (L3), and they underwent histological analysis. In all specimens the vestibular folds of both hemi-larynxes contained striated muscle bundles in variable amounts, representing a ventricularis muscle. These muscle bundles obviously originated from the lateral (external) and thyroepiglottic part of the thyroarytenoid muscle and the aryepiglottic part of the oblique arytenoid muscle, as has been described by other authors. The areas of vestibular folds and their amounts of ventricularis muscle bundles were measured using image analysis software (imageJ) by manual tracing. The mean area of the vestibular folds of both hemi-larynxes was 27.9 mm2 (SD [standard deviation] ± 9.17), and the area occupied by fibers of the ventricularis muscle was 1.5 mm2 (SD ± 1.78). Statistical analysis comparing the areas of both hemi-larynxes and levels resulted in no significant differences, except for the levels 2 and 3. In level 2, significantly more muscle fibers (2.0 mm2 ; SD ± 2.21) were detectable within the vestibular fold than in level 3 (0.9 mm2 ; SD ± 1.43). Level 1 also contained more muscle fibers (1.1 mm2 ; SD ± 1.06) than level 3, however, without significance. In conclusion, the laryngeal ventricularis muscle is present in the majority of reported cases. Since the muscle is of clinical relevance, it should be included in anatomical textbooks by default.

Laryngeal Framework Surgical Anatomy: A Radiological Study.

PURPOSE: Awareness of variations in laryngeal anatomy among different age and gender groups is crucial during laryngeal framework surgery. The aim of this study is to demonstrate the relationship between gender and laryngeal radiologic morphometrics among different age groups and the applicability of important anatomical landmarks of laryngeal surgery. METHODS: Laryngeal images of 180 adult patients older than 18 years of age were obtained by computed tomography and assessed. A total of 11 measurements of important laryngeal landmarks were taken from the patients' computed tomography images. Results were subgrouped according to gender and age, and these groups were compared for each measurement. RESULTS: The majority of laryngeal measurements obtained in the study were higher in males than females, with the exception of the interlaminar angle. The mean interlaminar angle value was 88.27°± 14.99 for males and 103.04°± 14.81 for females (P <0.005). The distance from the anterior commissure to the inferior border of the thyroid cartilage was 10.46 ± 2.5 mm for males and 7.72 ± 1.9 mm for females. The anterior commissure locates slightly higher than the midpoint of the distance from the thyroid notch to the thyroid inferior border. The shortest distance between the muscular process of the arytenoid cartilage and the thyroid cartilage was found to be 9.60 ± 3.47 mm for males and 7.72 ± 2.33 mm for females (P <0.001). CONCLUSION: Observation of obvious diversities in the size and distance of the important laryngeal structures between the gender groups is an important factor to be considered for successful laryngeal framework surgery. Also, using the midpoint of the thyroid cartilage as a landmark for anterior commissure is a practical method during surgery, especially for thyroplasty.

Anatomy of the Larynx and Cervical Trachea.

The larynx serves as the gateway between the upper and lower respiratory tracts and is involved in the tasks of phonation, deglutition, and airway protection. Familiarity with the complex anatomy of the larynx is critical for detecting and characterizing disease in the region, especially in cancer staging. In this article, we review the anatomy of the larynx and cervical trachea, including an overview of their cartilages, supporting tissues, muscles, mucosal spaces, neurovascular supply, and lymphatics, followed by correlation to the clinically relevant anatomic sites of the larynx. Imaging techniques for evaluating the larynx and trachea will also be discussed briefly.

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.

Anatomy of the respiratory system in the, "Portrait of a Musician" by Leonardo da Vinci.

The object of the research is the painting, Portrait of a Musician''. Upon closer examination of the painting, we discovered three details. Two have the shape of trachea and bronchi and the third a larynx. By moving the details through the program Paint X we got an image of the larynx, trachea and bronchi. The larynx is presented by thyroid and cricoid cartilage. The characteristic cartilaginous rings are distinguished on the trachea. The right and left main bronchi are also seen. In our opinion, Leonardo da Vinci reflected the elements of the respiratory system in the, Portrait of a Musician'', thus emphasizing the fact that the bronchi, trachea and larynx are involved in the production of sound.

Endoscopic and Computed Tomography Evaluation of Posterior Glottic Closure During Phonation.

OBJECTIVES/HYPOTHESIS: The anatomy of the posterior glottis, specifically the states of the posterior glottis during phonation, has not been thoroughly explored in laryngology. Conventional wisdom about the posterior glottis indicates that it tends to be completely closed in men but may be open in women. Furthermore, professional singers are expected to have a completely closed posterior glottis. The aim of this study was to investigate whether these generalizations are true by comparing rigid videolaryngostroboscopy results with high-resolution computed tomography (HRCT) and three-dimensional (3D) reconstruction findings. STUDY DESIGN: Prospective study. METHODS: Of the 90 volunteers (58 women, 32 men) examined, 48 were female professional singers, 10 were female nonsingers, 22 were male professional singers, and 10 were male nonsingers. Rigid videolaryngostroboscopy as well as HRCT scans were performed during singing at the average singing fundamental frequency. HRCT images of the larynx and air-column were 3D visualized using the software MIMICS®. The states of the posterior glottis were assessed in both examinations and compared among participants. RESULTS: The sensitivity of endoscopy was 67.5%. Complete closure of the posterior glottis was observed in 62.5% men and 52% women (P = .33). Complete closure of the posterior glottis was observed in 35% nonsingers and 61% professional singers (P = .036). CONCLUSIONS: The closure of the posterior glottis seen on videolaryngostroboscopy does not always correlate with actual closure. There seems to be no link between sex and complete closure of the glottis. However, there is strong evidence that posterior glottis closure can be influenced, to some degree, by vocal training. LEVEL OF EVIDENCE: 4 Laryngoscope, 132:124-129, 2022.

Effects of Laryngeal Vibratory Asymmetry and Neuromuscular Compensation on Voice Quality.

INTRODUCTION: Vibratory asymmetry and neuromuscular compensation are often seen in laryngeal neuromuscular pathology. However, the ramifications of these findings on voice quality are unclear. This study investigated the effects of varying levels of vibratory asymmetry and neuromuscular compensation on cepstral peak prominence (CPP), an analog of voice quality. STUDY DESIGN: In vivo canine phonation model. METHODS: Varying degrees of vocal fold vibratory asymmetry were achieved by stimulating one recurrent laryngeal nerve (RLN) over 11 levels from threshold to maximal muscle activation. For each of these levels, phonation was induced at systematically varied combinations of neuromuscular compensation: three levels each of contralateral RLN stimulation (80%, 90%, and 100% of maximal), superior laryngeal nerve (SLN) activation (0%, 50%, and 100% of maximal), and airflow levels (500, 700, and 900 mL/s). Vocal fold symmetry was determined by assessing the opening phase of the vibratory cycle in high-speed video recordings. Voice quality was estimated acoustically by calculating CPP for each voice sample. RESULTS: Eight hundred twenty-two phonatory conditions with varying degrees of vibratory asymmetry were evaluated. CPP was highest at vibratory symmetry. Increasing levels of asymmetry resulted in significant decreases in CPP. CPP increased significantly with increasing contralateral RLN activation. CPP was significantly higher at 50% SLN activation than 0% or 100% SLN activation. CONCLUSION: Voice quality, as approximated by CPP, is best at vibratory symmetry and deteriorates with increasing degrees of asymmetry. Voice quality may be improved with neuromuscular compensation by increased adduction of the contralateral vocal fold or increased vocal fold tension at mid-levels of SLN activation. LEVEL OF EVIDENCE: NA, Basic Science Laryngoscope, 132:130-134, 2022.

Implementation of a Low-Cost 3D-Printed Feline Larynx Model for Veterinary Students.

Endotracheal intubation (EI) in domestic cats is an important skill that veterinary students learn in order to perform anesthesia safely in this species. Implementing a 3D-printed larynx model (LaryngoCUBE) during the instruction process may improve student's learning of EI in felines. Twenty-two third-year students performed EI in cats with standard training (ST), and 16 students trained with the model (MT) the day before the laboratory. It was evaluated whether training with the model decreases the time and number of EI attempts, students' perceived difficulty performing EI using a visual analog score (VAS; 0 cm = very easy, 10 cm = extremely difficult; median [minimum-maximum]), and the incidence of failure to perform EI. The EI time on ST (58 [18-160] seconds) was longer, but not statistically different from MT (29 [13-120] seconds; p = .101). The number of EI attempts on ST (2 [1-3]) was higher than MT (1 [1-3]; p = .005). The VAS on the ST and MT were 4.5 (0.0-10.0) cm and 3.0 (0.2-10.0) cm, respectively (p = .029). The failure rate was 27% on the ST and 25% on the MT (p = 1.000). Students who practiced with a larynx model took fewer attempts to perform EI, tended to be faster, and found that EI was easier. However, the EI success rate in MT was not improved.

The hyoid as a sound conducting apparatus in laryngeally echolocating bats.

The morphology of the stylohyal-tympanic bone articulation found in laryngeally echolocating bats is highly indicative of a function associated with signal production. One untested hypothesis is that this morphology allows the transfer of a sound signal from the larynx to the tympanic bones (auditory bulla) via the hyoid apparatus during signal production by the larynx. We used µCT data and finite element analysis to model the propagation of sound through the hyoid chain into the tympanic bones to test this hypothesis. We modeled sound pressure (dB) wave propagation from the basihyal to the tympanic bones, vibratory behavior (m) of the stylohyal-tympanic bone unit, and the stylohyal and tympanic bones when the stylohyal bone is allowed to pivot on the tympanic bone. Sound pressure wave propagation was modeled using the harmonic acoustics solver in ANSYS and vibratory behavior was modeled using coupled modal and harmonic response analyses in ANSYS. For both analyses (harmonic acoustics and harmonic response), the input excitation on the basihyal and thyrohyals was modeled as the estimated pressure (Pa) imposed by the collision of the vibrating thyroid cartilage of the larynx against these bones during signal production. Our models support the hypothesis that this stereotypical hyoid morphology found in laryngeally echolocating bats can transfer sound to the auditory bullae at an amplitude that is likely heard for the species Artibeus jamaicensis and Rhinolophus pusillus.

Just Breathe: Tips and Highlights for Managing Pediatric Respiratory Distress and Failure.

Anatomically, the airway is ever changing in size, anteroposterior alignment, and point of most narrow dimension. Special considerations regarding obesity, chronic and acute illness, underlying developmental abnormalities, and age can all affect preparation and intervention toward securing a definitive airway. Mechanical ventilation strategies should focus on limiting peak inspiratory pressures and optimizing lung protective tidal volumes. Emergency physicians should work toward minimizing risk of peri-intubation hypoxemia and arrest. With review of anatomic and physiologic principles in the setting of a practical approach toward evaluating and managing distress and failure, emergency physicians can successfully manage critical pediatric airway encounters.

Echolocation in soft-furred tree mice.

Echolocation is the use of reflected sound to sense features of the environment. Here, we show that soft-furred tree mice (Typhlomys) echolocate based on multiple independent lines of evidence. Behavioral experiments show that these mice can locate and avoid obstacles in darkness using hearing and ultrasonic pulses. The proximal portion of their stylohyal bone fuses with the tympanic bone, a form previously only seen in laryngeally echolocating bats. Further, we found convergence of hearing-related genes across the genome and of the echolocation-related gene prestin between soft-furred tree mice and echolocating mammals. Together, our findings suggest that soft-furred tree mice are capable of echolocation, and thus are a new lineage of echolocating mammals.