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      Predicting Achievable Fundamental Frequency Ranges in Vocalization Across Species

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          Abstract

          Vocal folds are used as sound sources in various species, but it is unknown how vocal fold morphologies are optimized for different acoustic objectives. Here we identify two main variables affecting range of vocal fold vibration frequency, namely vocal fold elongation and tissue fiber stress. A simple vibrating string model is used to predict fundamental frequency ranges across species of different vocal fold sizes. While average fundamental frequency is predominantly determined by vocal fold length (larynx size), range of fundamental frequency is facilitated by (1) laryngeal muscles that control elongation and by (2) nonlinearity in tissue fiber tension. One adaptation that would increase fundamental frequency range is greater freedom in joint rotation or gliding of two cartilages (thyroid and cricoid), so that vocal fold length change is maximized. Alternatively, tissue layers can develop to bear a disproportionate fiber tension (i.e., a ligament with high density collagen fibers), increasing the fundamental frequency range and thereby vocal versatility. The range of fundamental frequency across species is thus not simply one-dimensional, but can be conceptualized as the dependent variable in a multi-dimensional morphospace. In humans, this could allow for variations that could be clinically important for voice therapy and vocal fold repair. Alternative solutions could also have importance in vocal training for singing and other highly-skilled vocalizations.

          Author Summary

          Mammals, birds, and reptiles vocalize (make sounds with vocal cords). Various species, and individuals within the species, are identified by pitch, loudness, and other acoustic features they can build into a repertoire of rhythmic and melodic patterns. Range of pitch, or more precisely fundamental frequency, is required to produce a variety of patterns. Whereas the average fundamental frequency is predictable by body size, the range of fundamental frequency depends on two different factors. The first is a freedom of movement factor–how much vocal cord length change can be produced by muscles that rotate or glide cartilages in the larynx. The second is a molecular composition factor–how much collagen density can be produced in the vocal cord ligament. Development and evolution has not been uniform with regard to these factors, suggesting that alternative choices are available for growth, training, and repair.

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          Most cited references49

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          The Role of Body Size, Phylogeny, and Ambient Noise in the Evolution of Bird Song

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            Functional Versus Morphological Diversity in Macroevolution

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              Synthesis of Voiced Sounds From a Two-Mass Model of the Vocal Cords

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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS Comput Biol
                PLoS Comput. Biol
                plos
                ploscomp
                PLoS Computational Biology
                Public Library of Science (San Francisco, CA USA )
                1553-734X
                1553-7358
                16 June 2016
                June 2016
                : 12
                : 6
                : e1004907
                Affiliations
                [1 ]National Center for Voice and Speech, University of Utah, Salt Lake City, Utah, United States of America
                [2 ]Department of Physiology, Midwestern University, Glendale, Arizona, United States of America
                [3 ]Department of Otolaryngology-Head and Neck Surgery, University of Texas Southwestern Medical Center, Dallas, Texas, United States of America
                University of California at Berkeley, UNITED STATES
                Author notes

                The authors have declared that no competing interests exist.

                Conceived and designed the experiments: IT TR TM. Performed the experiments: IT TR. Analyzed the data: IT TR TM. Wrote the paper: IT TR TM.

                Article
                PCOMPBIOL-D-15-01662
                10.1371/journal.pcbi.1004907
                4911068
                27309543
                b73c293f-391d-40eb-8eea-ce8e0ce10b14
                © 2016 Titze et al

                This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 1 October 2015
                : 8 April 2016
                Page count
                Figures: 5, Tables: 1, Pages: 13
                Funding
                Funded by: funder-id http://dx.doi.org/10.13039/100000055, National Institute on Deafness and Other Communication Disorders;
                Award ID: R01 DC013573-01
                Funding for this research was provided by the National Institute on Deafness and Other Communication Disorders ( www.nidcd.gov) R01 DC013573-01. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Research Article
                Physical Sciences
                Physics
                Classical Mechanics
                Vibration
                Biology and Life Sciences
                Anatomy
                Biological Tissue
                Connective Tissue
                Ligaments
                Medicine and Health Sciences
                Anatomy
                Biological Tissue
                Connective Tissue
                Ligaments
                Biology and Life Sciences
                Anatomy
                Neck
                Throat
                Larynx
                Medicine and Health Sciences
                Anatomy
                Neck
                Throat
                Larynx
                Biology and Life Sciences
                Anatomy
                Biological Tissue
                Connective Tissue
                Cartilage
                Medicine and Health Sciences
                Anatomy
                Biological Tissue
                Connective Tissue
                Cartilage
                Biology and Life Sciences
                Biochemistry
                Proteins
                Collagens
                Physical Sciences
                Physics
                Acoustics
                Biology and Life Sciences
                Organisms
                Animals
                Vertebrates
                Amniotes
                Birds
                Biology and Life Sciences
                Physiology
                Physiological Parameters
                Medicine and Health Sciences
                Physiology
                Physiological Parameters
                Custom metadata
                All relevant data is within the paper itself. Inquiries may be directed to the National Center for Voice and Speech website ( www.ncvs.org).

                Quantitative & Systems biology
                Quantitative & Systems biology

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