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• W2752673541 abstract "•The tracheophone syrinx is the first documented vocal organ with three sources•A tracheal membrane pair interacts with the main dual bronchial sound source•Interactions of the three sources generate various spectral and amplitude features•This morphological novelty facilitated the evolution of diverse vocalizations The evolution of complex behavior is driven by the interplay of morphological specializations and neuromuscular control mechanisms [1Galis F. The application of functional morphology to evolutionary studies.Trends Ecol. Evol. 1996; 11: 124-129Abstract Full Text PDF PubMed Scopus (50) Google Scholar, 2Fuxjager M.J. Schlinger B.A. Perspectives on the evolution of animal dancing: a case study of manakins.Curr. Opin. Behav. Sci. 2015; 6: 7-12Crossref Scopus (26) Google Scholar, 3Dickinson M.H. Farley C.T. Full R.J. Koehl M.A.R. Kram R. Lehman S. How animals move: an integrative view.Science. 2000; 288: 100-106Crossref PubMed Scopus (1152) Google Scholar], and it is often difficult to tease apart their respective contributions. Avian vocal learning and associated neural adaptations are thought to have played a major role in bird diversification [4Jarvis E.D. Mirarab S. Aberer A.J. Li B. Houde P. Li C. Ho S.Y.W. Faircloth B.C. Nabholz B. Howard J.T. et al.Whole-genome analyses resolve early branches in the tree of life of modern birds.Science. 2014; 346: 1320-1331Crossref PubMed Scopus (1177) Google Scholar, 5Whitney O. Pfenning A.R. Howard J.T. Blatti C.A. Liu F. Ward J.M. Wang R. Audet J.N. Kellis M. Mukherjee S. et al.Core and region-enriched networks of behaviorally regulated genes and the singing genome.Science. 2014; 346: 1256780Crossref PubMed Scopus (77) Google Scholar, 6Pfenning A.R. Hara E. Whitney O. Rivas M.V. Wang R. Roulhac P.L. Howard J.T. Wirthlin M. Lovell P.V. Ganapathy G. et al.Convergent transcriptional specializations in the brains of humans and song-learning birds.Science. 2014; 346: 1256846Crossref PubMed Scopus (283) Google Scholar, 7Zhang G. Li C. Li Q. Li B. Larkin D.M. Lee C. Storz J.F. Antunes A. Greenwold M.J. Meredith R.W. et al.Avian Genome ConsortiumComparative genomics reveals insights into avian genome evolution and adaptation.Science. 2014; 346: 1311-1320Crossref PubMed Scopus (616) Google Scholar, 8Verzijden M.N. ten Cate C. Servedio M.R. Kozak G.M. Boughman J.W. Svensson E.I. The impact of learning on sexual selection and speciation.Trends Ecol. Evol. 2012; 27: 511-519Abstract Full Text Full Text PDF PubMed Scopus (252) Google Scholar], whereas functional significance of substantial morphological diversity of the vocal organ remains largely unexplored. Within the most species-rich order, Passeriformes, “tracheophones” are a suboscine group that, unlike their oscine sister taxon, does not exhibit vocal learning [9Touchton J.M. Seddon N. Tobias J.A. Captive rearing experiments confirm song development without learning in a tracheophone suboscine bird.PLoS ONE. 2014; 9: e95746Crossref PubMed Scopus (44) Google Scholar] and is thought to phonate with tracheal membranes [10Rüppell W. Physiologie und Akustik der Vogelstimme.J. Ornithol. 1933; 81: 433-542Crossref Scopus (48) Google Scholar, 11Müller J. Über die bisher unbekannten typischen Verschiedenheiten der Stimmorgane der Passerinen. Abhandlungen der Königlichen Akademie der Wissenschaften, 1847: 321-391Google Scholar] instead of the two independent sources found in other passerines [12King A.S. Functional anatomy of the syrinx.in: King A.S. McLelland J. Form and Function in Birds. Academic Press, 1989: 105-192Google Scholar, 13Raikow R.J. Bledsoe A.H. Phylogeny and evolution of the passerine birds: independent methods of phylo-genetic analysis have produced a well-supported hypothesis of passerine phylogeny, one that has proved particularly useful in ecological and evolutionary studies.Bioscience. 2000; 50: 487-499Crossref Scopus (30) Google Scholar, 14Ames P.L. The Morphology of the Syrinx in Passerine Birds. Volume 37. Peabody Museum of Natural History, Yale University, 1971Google Scholar]. Here we show tracheophones possess three sound sources, two oscine-like labial pairs and the unique tracheal membranes, which collectively represent the largest described number of sound sources for a vocal organ. Birds with experimentally disabled tracheal membranes were still able to phonate. Instead of the main sound source, the tracheal membranes constitute a morphological specialization, which, through interaction with bronchial labia, contributes to different acoustic features such as spectral complexity, amplitude modulation, and enhanced sound amplitude. In contrast, these same features arise in oscines from neuromuscular control of two labial sources [15Suthers R.A. Zollinger S.A. Mechanisms of song production in songbirds.in: Zeigler H.P. Marler P. Neuroscience of Birdsong. Cambridge University Press, 2008: 78-98Google Scholar, 16Goller F. Riede T. Integrative physiology of fundamental frequency control in birds.J. Physiol. Paris. 2013; 107: 230-242Crossref PubMed Scopus (51) Google Scholar, 17Beckers G.J.L. Bird speech perception and vocal production: a comparison with humans.Hum. Biol. 2011; 83: 191-212Crossref PubMed Scopus (16) Google Scholar]. These findings are supported by a modeling approach and provide a clear example for how a morphological adaptation of the tracheophone vocal organ can generate specific, complex sound features. Morphological specialization therefore constitutes an alternative path in the evolution of acoustic diversity to that of oscine vocal learning and complex neural control. The evolution of complex behavior is driven by the interplay of morphological specializations and neuromuscular control mechanisms [1Galis F. The application of functional morphology to evolutionary studies.Trends Ecol. Evol. 1996; 11: 124-129Abstract Full Text PDF PubMed Scopus (50) Google Scholar, 2Fuxjager M.J. Schlinger B.A. Perspectives on the evolution of animal dancing: a case study of manakins.Curr. Opin. Behav. Sci. 2015; 6: 7-12Crossref Scopus (26) Google Scholar, 3Dickinson M.H. Farley C.T. Full R.J. Koehl M.A.R. Kram R. Lehman S. How animals move: an integrative view.Science. 2000; 288: 100-106Crossref PubMed Scopus (1152) Google Scholar], and it is often difficult to tease apart their respective contributions. Avian vocal learning and associated neural adaptations are thought to have played a major role in bird diversification [4Jarvis E.D. Mirarab S. Aberer A.J. Li B. Houde P. Li C. Ho S.Y.W. Faircloth B.C. Nabholz B. Howard J.T. et al.Whole-genome analyses resolve early branches in the tree of life of modern birds.Science. 2014; 346: 1320-1331Crossref PubMed Scopus (1177) Google Scholar, 5Whitney O. Pfenning A.R. Howard J.T. Blatti C.A. Liu F. Ward J.M. Wang R. Audet J.N. Kellis M. Mukherjee S. et al.Core and region-enriched networks of behaviorally regulated genes and the singing genome.Science. 2014; 346: 1256780Crossref PubMed Scopus (77) Google Scholar, 6Pfenning A.R. Hara E. Whitney O. Rivas M.V. Wang R. Roulhac P.L. Howard J.T. Wirthlin M. Lovell P.V. Ganapathy G. et al.Convergent transcriptional specializations in the brains of humans and song-learning birds.Science. 2014; 346: 1256846Crossref PubMed Scopus (283) Google Scholar, 7Zhang G. Li C. Li Q. Li B. Larkin D.M. Lee C. Storz J.F. Antunes A. Greenwold M.J. Meredith R.W. et al.Avian Genome ConsortiumComparative genomics reveals insights into avian genome evolution and adaptation.Science. 2014; 346: 1311-1320Crossref PubMed Scopus (616) Google Scholar, 8Verzijden M.N. ten Cate C. Servedio M.R. Kozak G.M. Boughman J.W. Svensson E.I. The impact of learning on sexual selection and speciation.Trends Ecol. Evol. 2012; 27: 511-519Abstract Full Text Full Text PDF PubMed Scopus (252) Google Scholar], whereas functional significance of substantial morphological diversity of the vocal organ remains largely unexplored. Within the most species-rich order, Passeriformes, “tracheophones” are a suboscine group that, unlike their oscine sister taxon, does not exhibit vocal learning [9Touchton J.M. Seddon N. Tobias J.A. Captive rearing experiments confirm song development without learning in a tracheophone suboscine bird.PLoS ONE. 2014; 9: e95746Crossref PubMed Scopus (44) Google Scholar] and is thought to phonate with tracheal membranes [10Rüppell W. Physiologie und Akustik der Vogelstimme.J. Ornithol. 1933; 81: 433-542Crossref Scopus (48) Google Scholar, 11Müller J. Über die bisher unbekannten typischen Verschiedenheiten der Stimmorgane der Passerinen. Abhandlungen der Königlichen Akademie der Wissenschaften, 1847: 321-391Google Scholar] instead of the two independent sources found in other passerines [12King A.S. Functional anatomy of the syrinx.in: King A.S. McLelland J. Form and Function in Birds. Academic Press, 1989: 105-192Google Scholar, 13Raikow R.J. Bledsoe A.H. Phylogeny and evolution of the passerine birds: independent methods of phylo-genetic analysis have produced a well-supported hypothesis of passerine phylogeny, one that has proved particularly useful in ecological and evolutionary studies.Bioscience. 2000; 50: 487-499Crossref Scopus (30) Google Scholar, 14Ames P.L. The Morphology of the Syrinx in Passerine Birds. Volume 37. Peabody Museum of Natural History, Yale University, 1971Google Scholar]. Here we show tracheophones possess three sound sources, two oscine-like labial pairs and the unique tracheal membranes, which collectively represent the largest described number of sound sources for a vocal organ. Birds with experimentally disabled tracheal membranes were still able to phonate. Instead of the main sound source, the tracheal membranes constitute a morphological specialization, which, through interaction with bronchial labia, contributes to different acoustic features such as spectral complexity, amplitude modulation, and enhanced sound amplitude. In contrast, these same features arise in oscines from neuromuscular control of two labial sources [15Suthers R.A. Zollinger S.A. Mechanisms of song production in songbirds.in: Zeigler H.P. Marler P. Neuroscience of Birdsong. Cambridge University Press, 2008: 78-98Google Scholar, 16Goller F. Riede T. Integrative physiology of fundamental frequency control in birds.J. Physiol. Paris. 2013; 107: 230-242Crossref PubMed Scopus (51) Google Scholar, 17Beckers G.J.L. Bird speech perception and vocal production: a comparison with humans.Hum. Biol. 2011; 83: 191-212Crossref PubMed Scopus (16) Google Scholar]. These findings are supported by a modeling approach and provide a clear example for how a morphological adaptation of the tracheophone vocal organ can generate specific, complex sound features. Morphological specialization therefore constitutes an alternative path in the evolution of acoustic diversity to that of oscine vocal learning and complex neural control. The avian vocal organ, the syrinx, shows remarkable morphological diversity across taxa [12King A.S. Functional anatomy of the syrinx.in: King A.S. McLelland J. Form and Function in Birds. Academic Press, 1989: 105-192Google Scholar], yet its significance for acoustic behavior is poorly understood. Notably, the syringes of oscines are quite similar [13Raikow R.J. Bledsoe A.H. Phylogeny and evolution of the passerine birds: independent methods of phylo-genetic analysis have produced a well-supported hypothesis of passerine phylogeny, one that has proved particularly useful in ecological and evolutionary studies.Bioscience. 2000; 50: 487-499Crossref Scopus (30) Google Scholar], yet oscines are considered to have the broadest range of acoustic features in their vocal repertoires. This broad range has been attributed in major part to vocal learning and complex neural control [4Jarvis E.D. Mirarab S. Aberer A.J. Li B. Houde P. Li C. Ho S.Y.W. Faircloth B.C. Nabholz B. Howard J.T. et al.Whole-genome analyses resolve early branches in the tree of life of modern birds.Science. 2014; 346: 1320-1331Crossref PubMed Scopus (1177) Google Scholar, 5Whitney O. Pfenning A.R. Howard J.T. Blatti C.A. Liu F. Ward J.M. Wang R. Audet J.N. Kellis M. Mukherjee S. et al.Core and region-enriched networks of behaviorally regulated genes and the singing genome.Science. 2014; 346: 1256780Crossref PubMed Scopus (77) Google Scholar, 6Pfenning A.R. Hara E. Whitney O. Rivas M.V. Wang R. Roulhac P.L. Howard J.T. Wirthlin M. Lovell P.V. Ganapathy G. et al.Convergent transcriptional specializations in the brains of humans and song-learning birds.Science. 2014; 346: 1256846Crossref PubMed Scopus (283) Google Scholar]. In contrast, the syringes of their sister group, suboscines, show remarkable morphological diversity [14Ames P.L. The Morphology of the Syrinx in Passerine Birds. Volume 37. Peabody Museum of Natural History, Yale University, 1971Google Scholar]. In particular, tracheophones have a unique set of membranes on the dorsal and ventral surfaces of the trachea directly above the tracheal bifurcation, the membranae tracheales (MT) [11Müller J. Über die bisher unbekannten typischen Verschiedenheiten der Stimmorgane der Passerinen. Abhandlungen der Königlichen Akademie der Wissenschaften, 1847: 321-391Google Scholar, 14Ames P.L. The Morphology of the Syrinx in Passerine Birds. Volume 37. Peabody Museum of Natural History, Yale University, 1971Google Scholar]. These tracheal membranes, in addition to the unique skeletal elements (the processi vocales) found on their sides, constitute a synapomorphy for the tracheophone clade [14Ames P.L. The Morphology of the Syrinx in Passerine Birds. Volume 37. Peabody Museum of Natural History, Yale University, 1971Google Scholar]. Based solely on anatomical inference, these membranes have been considered the main sound source of the tracheophone syrinx, although the presence of bronchial sources has been recognized [10Rüppell W. Physiologie und Akustik der Vogelstimme.J. Ornithol. 1933; 81: 433-542Crossref Scopus (48) Google Scholar, 14Ames P.L. The Morphology of the Syrinx in Passerine Birds. Volume 37. Peabody Museum of Natural History, Yale University, 1971Google Scholar]. However, here we show that sound generation in tracheophones relies on more complex morphology than commonly assumed. The presence of three vibratory sources gives rise to diverse sound features and may have played an important role in the diversification of this clade. In order to investigate potential sound sources, we used a fiberscope inserted into the trachea (Figure 1) to visualize the syrinx while we induced phonation by injecting air into the air sac system [18Goller F. Larsen O.N. In situ biomechanics of the syrinx and sound generation in pigeons.J. Exp. Biol. 1997; 200: 2165-2176PubMed Google Scholar]. In all six species investigated (Thamnophilus caerulescens, variable antshrike; Cercomacroides tyrannina, dusky antbird; Furnarius rufus, rufous ovenbird; Syndactyla rufosuperciliata, buff-browed foliage-gleaner; Lepidocolaptes angustirostris, narrow-billed woodcreeper; Certhiaxis cinnamomeus, yellow-chinned spinetail; Figure S1), the MTs were drawn into the trachea upon pressurization and were induced to oscillate under certain flow conditions (Movies S1 and S2). In addition, all species possess a labial sound source in each bronchus, which seem to be the main sound generators for vocalizations in most tracheophones. We used two approaches to verify this conclusion. First, we visualized the syrinx, and once the MTs began to oscillate, we pushed the fiberscope deeper so as to block their vibrations and see past them. After pushing past the MTs, we were able to see the bronchial labia, and they continued to oscillate and produce sound (Figure 1; Movie S1). Second, we prevented oscillations of the MTs by either rupturing the ventral MT or reducing elasticity by applying tissue adhesive to the ventral MT (these manipulative data are unavailable for C. cinnamomeus, and so this species is excluded from quantitative results). Despite these treatments, all species except the dusky antbird were still able to phonate spontaneously. These treatments resulted in specific acoustic changes, which indicate that in the intact tracheophone syrinx, all three sound sources can interact to produce complex acoustic features (Figures 2 and 3).Figure 2Manipulation of the Ventral MT Reveals Role in Sound Production and Interactions with Labial Sound Sources in F. rufusShow full caption(A–C) Examples (oscillographic and spectrographic representation) of spontaneous distress calls (A), induced phonation (B), and mathematically modeled sound (C) before (MT intact) and after (MT disabled) manipulation of the MT in F. rufus. Note the absence of the prominent, low-frequency amplitude modulations of spontaneous distress calls and the absence of the pulsations in induced sound after membrane manipulation, as well as an increase in fundamental frequency (see also Figures 3 and S3). Mathematical modeling of the MTs and a labial sound source interacting to produce sound (C) makes two clear predictions: (1) when the MTs oscillate at a frequency disparate from that of the bronchial sound sources, amplitude modulation will occur; and (2) when the natural frequency of the MTs is similar to that of the fundamental frequency of the bronchial sound sources, they lock and oscillate at an intermediate frequency.View Large Image Figure ViewerDownload Hi-res image Download (PPT)Figure 3Sound Frequency Shifts after Manipulation of MT in TracheophonesShow full caption(A–E) Examples of phonations (spectrograms) and frequency occurrence normalized to most common frequency before (red) and after (blue) manipulation are displayed for (A) F. rufus, spontaneous distress call (n = 3); (B) S. rufosuperciliata, spontaneous distress call (n = 1); (C) L. angustirostris, induced phonation (n = 1); (D) T. caerulescens, spontaneous distress call (n = 1); and (E) C. tyrannina, spontaneous distress call (n = 1) (see also Figure S1). All distributions after the manipulation are significantly different from those before (Kolmogorov-Smirnov, p < 0.001). Manipulation caused immobilization of the ventral MT (vMT) and was accomplished by either rupture or glue application. Sounds after manipulation are characterized by a shift to higher frequencies, either through absence of very low frequencies indicating loss of amplitude modulation as seen in F. rufus (see also Figure S3) or through smaller shifts indicating loss of locked vibration as seen in L. angustirostris and T. caerulescens (see also Figure S2). C. tyrannina was unable to produce sound after manipulation. The dotted portion of the x axis indicates the frequency range of the vocal repertoire of each species measured from recordings found at xeno-canto.org. Images downloaded from various sources. See also Movie S1.View Large Image Figure ViewerDownload Hi-res image Download (PPT) (A–C) Examples (oscillographic and spectrographic representation) of spontaneous distress calls (A), induced phonation (B), and mathematically modeled sound (C) before (MT intact) and after (MT disabled) manipulation of the MT in F. rufus. Note the absence of the prominent, low-frequency amplitude modulations of spontaneous distress calls and the absence of the pulsations in induced sound after membrane manipulation, as well as an increase in fundamental frequency (see also Figures 3 and S3). Mathematical modeling of the MTs and a labial sound source interacting to produce sound (C) makes two clear predictions: (1) when the MTs oscillate at a frequency disparate from that of the bronchial sound sources, amplitude modulation will occur; and (2) when the natural frequency of the MTs is similar to that of the fundamental frequency of the bronchial sound sources, they lock and oscillate at an intermediate frequency. (A–E) Examples of phonations (spectrograms) and frequency occurrence normalized to most common frequency before (red) and after (blue) manipulation are displayed for (A) F. rufus, spontaneous distress call (n = 3); (B) S. rufosuperciliata, spontaneous distress call (n = 1); (C) L. angustirostris, induced phonation (n = 1); (D) T. caerulescens, spontaneous distress call (n = 1); and (E) C. tyrannina, spontaneous distress call (n = 1) (see also Figure S1). All distributions after the manipulation are significantly different from those before (Kolmogorov-Smirnov, p < 0.001). Manipulation caused immobilization of the ventral MT (vMT) and was accomplished by either rupture or glue application. Sounds after manipulation are characterized by a shift to higher frequencies, either through absence of very low frequencies indicating loss of amplitude modulation as seen in F. rufus (see also Figure S3) or through smaller shifts indicating loss of locked vibration as seen in L. angustirostris and T. caerulescens (see also Figure S2). C. tyrannina was unable to produce sound after manipulation. The dotted portion of the x axis indicates the frequency range of the vocal repertoire of each species measured from recordings found at xeno-canto.org. Images downloaded from various sources. See also Movie S1. Consistent across all species, the intact syrinx produced low-frequency sounds, characterized by vibrations with pulse-like time waveforms, giving rise to complex harmonic content of sounds. Disabling of the MTs resulted in three consistent changes. First, pulse-like quality decreased as indicated by the increased duty cycle (Figure 4). This decrease in pulse-like quality was found in all four species for which data after manipulation of the membrane are available. Second, sound amplitude significantly decreased (Figure 4D). Lastly, sound frequency showed a consistent increase after membrane disabling (Figures 2, 3, and S2), though the degree of increase varies between species. Strikingly, the prominent amplitude modulations in spontaneously generated distress calls of the rufous ovenbird (F. rufus) were absent after ventral MT rupture (Figure 2). The low-frequency component representing the vibration of the MTs was strongly reduced after the membrane was disabled with various approaches (membrane rupture, membrane gluing, and fibroscopic prevention of vibration) for both spontaneous distress calls (Figures 2A and 3A) and induced phonation (Figure 3B). This strongly suggests that the MTs produce the low-frequency vibrations that drive the change in amplitude, which is superimposed on the higher-frequency labial oscillations. This proposed mechanism for the generation of amplitude modulation involves interaction of the three sound sources. To confirm that the amplitude modulations are not produced by neuromuscular control, we denervated the syringeal muscles by transection of the tracheosyringeal nerves, and amplitude modulation clearly persisted (Figure S3). To better understand these interactions, we used a mathematical model of the syrinx [19Arneodo E.M. Perl Y.S. Mindlin G.B. Acoustic signatures of sound source-tract coupling.Phys. Rev. E Stat. Nonlin. Soft Matter Phys. 2011; 83: 041920Crossref PubMed Scopus (7) Google Scholar] to generate sound with two different oscillators representing one set of labia and the MTs. If the frequencies of the two sources are sufficiently different, the lower frequency of the MTs modulates the higher frequency of the labial sound source (Figure 2C, “pulsations”) as seen in the pronounced amplitude modulation in the spontaneous calls of the rufous ovenbird (Figure 2A). If the frequencies are more similar to each other, the two sources become “locked,” and the resulting vibrations are of lower frequency than if the labial source were to vibrate on its own (Figure 2C, “locking”). This is consistent with the observed shift to higher frequencies after membrane disabling across species (Figure 3), as well as reduced pulse-like quality of the vibrations (Figure 4) [20Jensen K.K. Cooper B.G. Larsen O.N. Goller F. Songbirds use pulse tone register in two voices to generate low-frequency sound.Proc. Biol. Sci. 2007; 274: 2703-2710Crossref PubMed Scopus (29) Google Scholar, 21Sitt J.D. Amador A. Goller F. Mindlin G.B. Dynamical origin of spectrally rich vocalizations in birdsong.Phys. Rev. E Stat. Nonlin. Soft Matter Phys. 2008; 78: 011905Crossref PubMed Scopus (42) Google Scholar]. The pulsatile oscillations arise from the nonlinear interaction of two oscillators, the MTs and the labia. We assume that the labial oscillators can be represented by a harmonic oscillation of higher frequency. For this reason, when the MT is disabled, the remaining labial sources are free to oscillate at their natural, higher frequency and with more harmonic character [22Strogatz S.H. Nonlinear Dynamics and Chaos: With Applications to Physics, Biology, Chemistry, and Engineering. Westview press, 2014Google Scholar]. In the other species, frequency also significantly increases, but the increase is smaller than that found in F. rufus. This further illustrates the diversity by which the MTs affect sound features in different tracheophones. In other words, the MTs and bronchial labia of F. rufus likely vibrate at significantly disparate frequencies (as supported by data from the model and the presence of amplitude modulation), which therefore results in a large shift to higher frequencies after MT manipulation, while the variable antshrike shows a much smaller shift to higher frequencies, indicating the MTs and bronchial labia likely vibrate at more similar frequencies. In our simulations, we found that sounds generated with locked sound sources tend to be of higher amplitude than those generated with only one sound source. Consistent with that observation, the amplitude of spontaneous distress calls of individuals, for which data are available, decreased markedly after the membrane was disabled (Figure 4D). To quantify sound amplitude before and after membrane disabling in spontaneously generated distress calls, we expressed amplitude as signal amplitude (volts) relative to background noise (volts) to account for different recording settings. This reduction in sound amplitude was consistent for all manipulations (note that the L. angustirostris individual did not produce spontaneous distress calls after manipulation, and so amplitude data are unavailable) and can therefore not have resulted from altered pressure differentials across the syrinx after membrane rupture. Lastly, simulations of the model indicate that the transition between pulsations and locking (Figure 2C) occurs with certain changes in pressure. This may explain in part the diversity with which tracheophones seem to use the MTs (Figures 3 and 4) since, depending on the natural pressure produced by the bird, they may or may not produce sounds with both pulsations or locking and may instead produce one or the other. The MTs and their interactions with the bronchial sound sources in the tracheophone syrinx constitute a morphological solution to generating loud sounds, amplitude modulations, pulse-like quality, and low-frequency sounds. Ultimately, these acoustic features could be attributed to the increased resistance across the tracheophone syrinx when the MTs are engaged as they cause a narrowing of the trachea. While oscines achieve some of these acoustic features through neuromuscular control of their two sound sources [20Jensen K.K. Cooper B.G. Larsen O.N. Goller F. Songbirds use pulse tone register in two voices to generate low-frequency sound.Proc. Biol. Sci. 2007; 274: 2703-2710Crossref PubMed Scopus (29) Google Scholar, 21Sitt J.D. Amador A. Goller F. Mindlin G.B. Dynamical origin of spectrally rich vocalizations in birdsong.Phys. Rev. E Stat. Nonlin. Soft Matter Phys. 2008; 78: 011905Crossref PubMed Scopus (42) Google Scholar, 23Suthers R.A. Contributions to birdsong from the left and right sides of the intact syrinx.Nature. 1990; 347: 473-477Crossref Scopus (121) Google Scholar, 24Suthers R.A. Goller F. Hartley R.S. Motor dynamics of song production by mimic thrushes.J. Neurobiol. 1994; 25: 917-936Crossref PubMed Scopus (71) Google Scholar, 25Goller F. Suthers R.A. Role of syringeal muscles in controlling the phonology of bird song.J. Neurophysiol. 1996; 76: 287-300Crossref PubMed Scopus (147) Google Scholar, 26Goller F. Suthers R.A. Role of syringeal muscles in gating airflow and sound production in singing brown thrashers.J. Neurophysiol. 1996; 75: 867-876Crossref PubMed Scopus (137) Google Scholar, 27Srivastava K.H. Elemans C.P.H. Sober S.J. Multifunctional and context-dependent control of vocal acoustics by individual muscles.J. Neurosci. 2015; 35: 14183-14194Crossref PubMed Scopus (22) Google Scholar], suboscines do not have the same level of neuromuscular control [12King A.S. Functional anatomy of the syrinx.in: King A.S. McLelland J. Form and Function in Birds. Academic Press, 1989: 105-192Google Scholar, 28Amador A. Goller F. Mindlin G.B. Frequency modulation during song in a suboscine does not require vocal muscles.J. Neurophysiol. 2008; 99: 2383-2389Crossref PubMed Scopus (46) Google Scholar] and instead likely rely on intrinsic, morphological qualities such as the MTs to achieve these acoustic features. Our data support this and consistently indicate that the intact MTs enable the generation of higher amplitudes, increased pulse-like quality, and lower frequencies. In addition, different tracheophone species seem to use the MTs to different effect, as not all species in this study produce prominent amplitude modulations. Of those that do (F. rufus" @default.
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• W2752673541 title "Evolution of Vocal Diversity through Morphological Adaptation without Vocal Learning or Complex Neural Control" @default.
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