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Adapted to Roar: Functional Morphology of Tiger and
Lion Vocal Folds
1 2,3 4 1,3
Sarah A. Klemuk , Tobias Riede *, Edward J. Walsh , Ingo R. Titze
1 Department of Communication Sciences and Disorders, The University of Iowa, Iowa City, Iowa, United States of America, 2 Department of Biology, The University of
Utah, Salt Lake City, Utah, United States of America, 3 National Center for Voice and Speech, The University of Utah, Salt Lake City, Utah, United States of America, 4 Boys
Town National Research Hospital, Omaha, Nebraska, United States of America
Abstract
Vocal production requires active control of the respiratory system, larynx and vocal tract. Vocal sounds in mammals are
produced by flow-induced vocal fold oscillation, which requires vocal fold tissue that can sustain the mechanical stress
during phonation. Our understanding of the relationship between morphology and vocal function of vocal folds is very
limited. Here we tested the hypothesis that vocal fold morphology and viscoelastic properties allow a prediction of
fundamental frequency range of sounds that can be produced, and minimal lung pressure necessary to initiate phonation.
We tested the hypothesis in lions and tigers who are well-known for producing low frequency and very loud roaring sounds
that expose vocal folds to large stresses. In histological sections, we found that the Panthera vocal fold lamina propria
consists of a lateral region with adipocytes embedded in a network of collagen and elastin fibers and hyaluronan. There is
also a medial region that contains only fibrous proteins and hyaluronan but no fat cells. Young’s moduli range between 10
and 2000 kPa for strains up to 60%. Shear moduli ranged between
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