Discovery of a low frequency sound source in Mysticeti (baleen whales): anatomical establishment of a vocal fold homolog.

The mechanism of mysticete (baleen whale) vocalization has remained a mystery. Vocal folds (true vocal "cords"), the structures responsible for sound production in terrestrial mammals, were thought to be absent in whales. This study tests the hypothesis that the mysticete larynx possesses structures homologous to vocal folds and that they are capable of sound generation. Laryngeal anatomy was examined in 37 specimens representing 6 mysticete species. Results indicate the presence of a U-shaped fold (U-fold) in the lumen of the larynx. The U-fold is supported by arytenoid cartilages, controlled by skeletal muscles innervated by the recurrent laryngeal nerve, is adjacent to a diverticulum (laryngeal sac) covered with mucosa innervated by the superior laryngeal nerve, and contains a ligament-conditions that also define the vocal folds of terrestrial mammals and, therefore, supports homology. Unlike the vocal folds of terrestrial mammals, which are perpendicular to airflow, the mysticete U-fold is oriented parallel to airflow. U-fold adduction/abduction and elevation/depression may control airflow, and vibration of its edges may generate sounds. The walls of the laryngeal sac can expand and contract, may serve as a resonant space, and may also propagate vibrations generated by movements of the supporting arytenoid cartilages. The extensive musculature surrounding the laryngeal sac may enable rapid and forceful expulsion of air from the lumen of the sac into other respiratory spaces, or maintain a constant sac volume despite the effects of ambient pressure (e.g., changes during diving or ascent). The size and complexity of the mysticete larynx indicates an organ with multiple functions, including protection during breathing/swallowing, regulation of airflow and pressures in the respiratory spaces, and sound generation. The presence of a vocal fold homolog offers a new insight into both the mechanism of sound generation by mysticetes and the divergent evolution of odontocete and mysticete cetaceans.