Air Sac Pressure Research Articles

Song-like activation of syringeal and respiratory muscles during sleep in canaries.

Sleep replay activity involves the reactivation of brain structures with patterns similar to those observed during waking behavior. In this study, we demonstrate that adult male canaries exhibit spontaneous, song-like peripheral reactivation during night sleep. Our findings include: (1) the presence of activity in respiratory muscles, leading to song-like air sac pressure patterns of low amplitude, (2) the simultaneous occurrence of respiratory replay events and reactivation of syringeal muscles, and (3) the reactivation of syringeal muscles without concurrent respiratory system activity. This song-like reactivation of peripheral motor systems enables the identification of specific motor patterns, with replay events preserving individual morphological and temporal properties. The activation of peripheral motor systems in songbirds and the differences in activation patterns between species give unique insights into the fictive behavioral output of activation of a complex learned motor behavior during sleep, shedding light on the neural control mechanisms and potential functions.

A neural circuit for vocal production responds to viscerosensory input in the songbird.

Motor performance is monitored continuously by specialized brain circuits and used adaptively to modify behavior on a moment-to-moment basis and over longer time periods. During vocal behaviors, such as singing in songbirds, internal evaluation of motor performance relies on sensory input from the auditory and vocal-respiratory systems. Sensory input from the auditory system to the motor system, often referred to as auditory feedback, has been well studied in singing zebra finches (Taeniopygia guttata), but little is known about how and where nonauditory sensory feedback is evaluated. Here we show that brief perturbations in air sac pressure cause short-latency neural responses in the higher-order song control nucleus HVC (used as proper name), an area necessary for song learning and song production. Air sacs were briefly pressurized through a cannula in anesthetized or sedated adult male zebra finches, and neural responses were recorded in both nucleus parambigualis (PAm), a brainstem inspiratory center, and HVC, a cortical premotor nucleus. These findings show that song control nuclei in the avian song system are sensitive to perturbations directly targeted to vocal-respiratory, or viscerosensory, afferents and support a role for multimodal sensory feedback integration in modifying and controlling vocal control circuits.NEW & NOTEWORTHY This study presents the first evidence of sensory input from the vocal-respiratory periphery directly activating neurons in a motor circuit for vocal production in songbirds. It was previously thought that this circuit relies exclusively on sensory input from the auditory system, but we provide groundbreaking evidence for nonauditory sensory input reaching the higher-order premotor nucleus HVC, expanding our understanding of what sensory feedback may be available for vocal control.

System for Monitoring Avian Cardiac Output and Breathing Patterns Using Transmission-Type Microwaves

We report the development of a non-contact monitoring device for avian cardiac output and breathing patterns based on the anterior thoracic air sac pressure that uses transmission-type microwaves (2,400−2,500 MHz, continuous wave). Since the phase waveform represents the dielectric constant change, the phase reflects −j/ωc and the dielectric constant change is related to blood flow. The magnitude waveform is reflected from the electronic resistance of tissues due to the expansion of the anterior thoracic air sac, which mainly consists of the thoracic wall. To confirm these waveforms, pigeons and chickens were used for testing. To validate the output waveforms of the developed transmission-type microwave device, data from esophageal catheters and pressure sensors in the anterior thoracic air sac, abdominal air sac, and intraoral cavity were obtained. The waveform for the esophageal catheter, where electrocardiogram electrodes and an angular velocity sensor were installed, correlates with cardiac output. A heart sound microphone was used to confirm the closing sound of the arterial and mitral valves. The experimental results confirm that a linear waveform synchronized with the cardiac blood flow and the anterior thoracic air sac pressure of birds was obtained using transmission-type microwaves. The proposed device, which can monitor cardiac output and respiratory patterns, may enable the early screening of cytokine storms caused by avian influenza viruses. Existing devices use Doppler radar in the 10 to 77 GHz band; these high frequencies are reflected by the chest wall and do not reach deep into body, making it impossible to monitor the blood flow inside the body.

Dynamics behind rough sounds in the song of the Pitangus sulphuratus.

The complex vocalizations found in different bird species emerge from the interplay between morphological specializations and neuromuscular control mechanisms. In this work we study the dynamical mechanisms used by a nonlearner bird from the Americas, the suboscine Pitangus sulphuratus, in order to achieve a characteristic timbre of some of its vocalizations. By measuring syringeal muscle activity, air sac pressure, and sound as the bird sings, we are able to show that the birds of this species manage to lock the frequency difference between two sound sources. This provides a precise control of sound amplitude modulations, which gives rise to a distinct timbral property.

Dynamical model for the neural activity of singing Serinus canaria.

Vocal production in songbirds is a key topic regarding the motor control of a complex, learned behavior. Birdsong is the result of the interaction between the activity of an intricate set of neural nuclei specifically dedicated to song production and learning (known as the "song system"), the respiratory system and the vocal organ. These systems interact and give rise to precise biomechanical motor gestures which result in song production. Telencephalic neural nuclei play a key role in the production of motor commands that drive the periphery, and while several attempts have been made to understand their coding strategy, difficulties arise when trying to understand neural activity in the frame of the song system as a whole. In this work, we report neural additive models embedded in an architecture compatible with the song system to provide a tool to reduce the dimensionality of the problem by considering the global activity of the units in each neural nucleus. This model is capable of generating outputs compatible with measurements of air sac pressure during song production in canaries (Serinus canaria). In this work, we show that the activity in a telencephalic nucleus required by the model to reproduce the observed respiratory gestures is compatible with electrophysiological recordings of single neuron activity in freely behaving animals.

2335 the Application of Air SAC Hemostasis Device in Laproscopic Ovarian Cystectomy to Protect Ovarian Function

Video Objective To evaluate the efficacy of air pressure device in laparoscopic ovarian endometrioma removal for ovarian wound hemostasis and the significance of ovarian function protection. Setting A prospective study in university hospital. Interventions We developed the air sac device and patented it. After the cyst removed, we placed the air sac pressure device into the pelvic cavity through the right trocar. And then we pumped the air into the sac through a three port valve. When the sac flowed up, the air in sac created air pressure to the wound surface of the ovary. The pressure was kept for 6-10 minutes to stop the bleeding. In 5 cases, the bleeding on ovarian wound surface was inhibited effectively, and the electric coagulation was reduced greatly. Conclusion The application of air sac hemostasis device in laparoscopic cystectomy could stop the bleeding effectively and then reduce the electric coagulation greatly. This may avoid the excessive heat damage to ovarian tissue.

An integrated model for motor control of song in Serinus canaria

Birdsong is a learned motor behavior controlled by an interconnected structure of neural nuclei. This pathway is bilaterally organized, with anatomically indistinguishable structures in each brain hemisphere. In this work, we present a computational model whose variables are the average activities of different neural nuclei of the song system of oscine birds. Two of the variables are linked to the air sac pressure and the tension of the labia during canary song production. We show that these time dependent gestures are capable of driving a model of the vocal organ to synthesize realistic canary like songs.

Motor control of sound frequency in birdsong involves the interaction between air sac pressure and labial tension.

Frequency modulation is a salient acoustic feature of birdsong. Its control is usually attributed to the activity of syringeal muscles, which affect the tension of the labia responsible for sound production. We use experimental and theoretical tools to test the hypothesis that for birds producing tonal sounds such as domestic canaries (Serinus canaria), frequency modulation is determined by both the syringeal tension and the air sac pressure. For different models, we describe the structure of the isofrequency curves, which are sets of parameters leading to sounds presenting the same fundamental frequencies. We show how their shapes determine the relative roles of syringeal tension and air sac pressure in frequency modulation. Finally, we report experiments that allow us to unveil the features of the isofrequency curves.

A Mechanism for Frequency Modulation in Songbirds Shared with Humans

In most animals that vocalize, control of fundamental frequency is a key element for effective communication. In humans, subglottal pressure controls vocal intensity but also influences fundamental frequency during phonation. Given the underlying similarities in the biomechanical mechanisms of vocalization in humans and songbirds, songbirds offer an attractive opportunity to study frequency modulation by pressure. Here, we present a novel technique for dynamic control of subsyringeal pressure in zebra finches. By regulating the opening of a custom-built fast valve connected to the air sac system, we achieved partial or total silencing of specific syllables, and could modify syllabic acoustics through more complex manipulations of air sac pressure. We also observed that more nuanced pressure variations over a limited interval during production of a syllable concomitantly affected the frequency of that syllable segment. These results can be explained in terms of a mathematical model for phonation that incorporates a nonlinear description for the vocal source capable of generating the observed frequency modulations induced by pressure variations. We conclude that the observed interaction between pressure and frequency was a feature of the source, not a result of feedback control. Our results indicate that, beyond regulating phonation or its absence, regulation of pressure is important for control of fundamental frequencies of vocalizations. Thus, although there are separate brainstem pathways for syringeal and respiratory control of song production, both can affect airflow and frequency. We hypothesize that the control of pressure and frequency is combined holistically at higher levels of the vocalization pathways.

Characterization of respiratory neurons in the rostral ventrolateral medulla, an area critical for vocal production in songbirds

Much is known about the neuronal cell types and circuitry of the mammalian respiratory brainstem and its role in normal, quiet breathing. Our understanding of the role of respiration in the context of vocal production, however, is very limited. Songbirds contain a well-defined neural circuit, known as the song system, which is necessary for song production and is strongly coupled to the respiratory system. A major target of this system is nucleus parambigualis (PAm) in the ventrolateral medulla, a structure that controls inspiration by way of its bulbospinal projections but is also an integral part of the song-pattern generation circuit by way of its "thalamocortical" projections to song-control nuclei in the telencephalon. We have mapped out PAm to characterize the cell types and its functional organization. Extracellular single units were obtained in anesthetized adult male zebra finches while measuring air sac pressure to monitor respiration. Single units were characterized by their discharge patterns and the phase of the activity in the respiratory cycle. Several classes of neurons were identified and were analogous to those reported for mammalian medullary respiratory neurons. The majority of the neurons in PAm was classified as inspiratory augmenting or preinspiratory, although other basic discharge patterns were observed as well. The well-characterized connectivity of PAm within the vocal motor circuit and the similarity of its neural firing patterns to the rostral ventral respiratory group and pre-Bötzinger complex of mammals make it an ideal system for investigating the integration of breathing and vocalization.

Control of fundamental frequency in zebra finch vocal behavior

Event Abstract Back to Event Control of fundamental frequency in zebra finch vocal behavior Franz Goller1* and Tobias Riede1 1 University of Utah, Biology, United States In the avian syrinx generates sound by flow-induced tissue (labia or membrane) oscillations. The fundamental frequency of sound (F0) arises from a complex interplay of biomechanical and aerodynamic properties as well as neural control of syringeal movements and respiration. Each variable affects labial position and tension either directly or indirectly, and synchronously or independently. In the zebra finch (Taeniopygia guttata), song syllables with low F0 are pulse-tone like and are generated by both sound sources, whereas high-frequency syllables are tonal and are generated by the right side of the syrinx. It has been suggested that these different vibratory modes arise from different dynamical regimes (Sitt et al., Physical Rev E 78,011905, 2008). No major asymmetries exist in the extracellular matrix or size of the left and right labia (Riede et al., PLoS One 5, e11368), suggesting that difference in F0 of the two sides involves aerodynamic and neural mechanisms. Here we investigate the respective roles of all contributing variables in setting F0 of different zebra finch sounds. Ventral syringeal muscle activity increases with increasing F0 for the pulse-tone syllables. For high-frequency syllables, activation is maximal, but is not correlated with differences in F0. The activity of other syringeal muscles (dorsal syringeal, ventral tracheobronchial and dorsal tracheobronchial) shows a similar relationship with F0, indicating that synergistic activation patterns cannot explain F0-control for high-frequency syllables. The possibility that active closing of the left side generates a biomechanical gearing of the effect of right-side tension control by muscles can also be eliminated. Denervation of the left syringeal muscles does not consistently change F0 of right-side generated syllables. The effect of air sac pressure on F0 was investigated by bilateral denervation of the syringeal muscles. F0 increases slightly with increasing air sac pressure (approx. 20-45 Hz per kPa) if neural control of labial tension and position is eliminated. We then asked whether zebra finches compensate for the increase in F0 at higher air sac pressures through muscle control when they vocalize with different sound amplitude. Changes in song amplitude arise from differences in air sac pressure and can be induced by the Lombard effect or by experimentally reducing expiratory volume of air. In both manipulations the air sac pressure changes are accompanied by corresponding changes in F0. The changes in F0 are larger in high-frequency syllables than in low-frequency harmonic stacks. These results suggest that zebra finches do not adjust muscular control of labial tension to compensate for this effect of air sac pressure on F0. These results illustrate how aerodynamic effects and neural control of labial tension interact in setting F0 of song syllables. However, tension control for the high-frequency modal sounds is still not fully understood. Although we can eliminate some hypotheses (gearing, muscle synergies), the precise control mechanism is still unknown. Acknowledgements: supported by NIDCD 06876. Keywords: birdsong, frequency control Conference: Tenth International Congress of Neuroethology, College Park. Maryland USA, United States, 5 Aug - 10 Aug, 2012. Presentation Type: Poster Presentation (see alternatives below as well) Topic: Sensorimotor Integration Citation: Goller F and Riede T (2012). Control of fundamental frequency in zebra finch vocal behavior. Conference Abstract: Tenth International Congress of Neuroethology. doi: 10.3389/conf.fnbeh.2012.27.00367 Copyright: The abstracts in this collection have not been subject to any Frontiers peer review or checks, and are not endorsed by Frontiers. They are made available through the Frontiers publishing platform as a service to conference organizers and presenters. The copyright in the individual abstracts is owned by the author of each abstract or his/her employer unless otherwise stated. Each abstract, as well as the collection of abstracts, are published under a Creative Commons CC-BY 4.0 (attribution) licence (https://creativecommons.org/licenses/by/4.0/) and may thus be reproduced, translated, adapted and be the subject of derivative works provided the authors and Frontiers are attributed. For Frontiers’ terms and conditions please see https://www.frontiersin.org/legal/terms-and-conditions. Received: 01 May 2012; Published Online: 07 Jul 2012. * Correspondence: Dr. Franz Goller, University of Utah, Biology, Salt Lake City, Utah, 84112, United States, goller@biology.utah.edu Login Required This action requires you to be registered with Frontiers and logged in. To register or login click here. Abstract Info Abstract The Authors in Frontiers Franz Goller Tobias Riede Google Franz Goller Tobias Riede Google Scholar Franz Goller Tobias Riede PubMed Franz Goller Tobias Riede Related Article in Frontiers Google Scholar PubMed Abstract Close Back to top Javascript is disabled. Please enable Javascript in your browser settings in order to see all the content on this page.

Reconstruction of physiological instructions from Zebra finch song

We reconstruct the physiological parameters that control an avian vocal organ during birdsong production using recorded song. The procedure involves fitting the time dependent parameters of an avian vocal organ model. Computationally, the model is implemented as a dynamical system ruling the behavior of the oscillating labia that modulate the air flow during sound production, together with the equations describing the dynamics of pressure fluctuations in the vocal tract. We tested our procedure for Zebra finch song with, simultaneously recorded physiological parameters: air sac pressure and the electromyographic activity of the left and right ventral syringeal muscles. A comparison of the reconstructed instructions with measured physiological parameters during song shows a high degree of correlation. Integrating the model with reconstructed parameters leads to the synthesis of highly realistic songs.

Control of Vocal and Respiratory Patterns in Birdsong: Dissection of Forebrain and Brainstem Mechanisms Using Temperature

Learned motor behaviors require descending forebrain control to be coordinated with midbrain and brainstem motor systems. In songbirds, such as the zebra finch, regular breathing is controlled by brainstem centers, but when the adult songbird begins to sing, its breathing becomes tightly coordinated with forebrain-controlled vocalizations. The periods of silence (gaps) between song syllables are typically filled with brief breaths, allowing the bird to sing uninterrupted for many seconds. While substantial progress has been made in identifying the brain areas and pathways involved in vocal and respiratory control, it is not understood how respiratory and vocal control is coordinated by forebrain motor circuits. Here we combine a recently developed technique for localized brain cooling, together with recordings of thoracic air sac pressure, to examine the role of cortical premotor nucleus HVC (proper name) in respiratory-vocal coordination. We found that HVC cooling, in addition to slowing all song timescales as previously reported, also increased the duration of expiratory pulses (EPs) and inspiratory pulses (IPs). Expiratory pulses, like song syllables, were stretched uniformly by HVC cooling, but most inspiratory pulses exhibited non-uniform stretch of pressure waveform such that the majority of stretch occurred late in the IP. Indeed, some IPs appeared to change duration by the earlier or later truncation of an underlying inspiratory event. These findings are consistent with the idea that during singing the temporal structure of EPs is under the direct control of forebrain circuits, whereas that of IPs can be strongly influenced by circuits downstream of HVC, likely in the brainstem. An analysis of the temporal jitter of respiratory and vocal structure suggests that IPs may be initiated by HVC at the end of each syllable and terminated by HVC immediately before the onset of the next syllable.

Metabolic and respiratory costs of increasing song amplitude in zebra finches.

Bird song is a widely used model in the study of animal communication and sexual selection, and several song features have been shown to reflect the quality of the singer. Recent studies have demonstrated that song amplitude may be an honest signal of current condition in males and that females prefer high amplitude songs. In addition, birds raise the amplitude of their songs to communicate in noisy environments. Although it is generally assumed that louder song should be more costly to produce, there has been little empirical evidence to support this assumption. We tested the assumption by measuring oxygen consumption and respiratory patterns in adult male zebra finches (Taeniopygia guttata) singing at different amplitudes in different background noise conditions. As background noise levels increased, birds significantly increased the sound pressure level of their songs. We found that louder songs required significantly greater subsyringeal air sac pressure than quieter songs. Though increased pressure is probably achieved by increasing respiratory muscle activity, these increases did not correlate with measurable increases in oxygen consumption. In addition, we found that oxygen consumption increased in higher background noise, independent of singing behaviour. This observation supports previous research in mammals showing that high levels of environmental noise can induce physiological stress responses. While our study did not find that increasing vocal amplitude increased metabolic costs, further research is needed to determine whether there are other non-metabolic costs of singing louder or costs associated with chronic noise exposure.

Learning to breathe and sing: development of respiratory-vocal coordination in young songbirds

How do animals with learned vocalizations coordinate vocal production with respiration? Songbirds such as the zebra finch learn their songs, beginning with highly variable babbling vocalizations known as subsong. After several weeks of practice, zebra finches are able to produce a precisely timed pattern of syllables and silences, precisely coordinated with expiratory and inspiratory pulses (Franz M, Goller F. J Neurobiol 51: 129-141, 2002). While respiration in adult song is well described, relatively little is known about respiratory patterns in subsong or about the processes by which respiratory and vocal patterns become coordinated. To address these questions, we recorded thoracic air sac pressure in juvenile zebra finches prior to the appearance of any consistent temporal or acoustic structure in their songs. We found that subsong contains brief inspiratory pulses (50 ms) alternating with longer pulses of sustained expiratory pressure (50-500 ms). In striking contrast to adult song, expiratory pulses often contained multiple (0-8) variably timed syllables separated by expiratory gaps and were only partially vocalized. During development, expiratory pulses became shorter and more stereotyped in duration with shorter and fewer nonvocalized parts. These developmental changes eventually resulted in the production of a single syllable per expiratory pulse and a single inspiratory pulse filling each gap, forming a coordinated sequence similar to that of adult song. To examine the role of forebrain song-control nuclei in the development of respiratory patterns, we performed pressure recordings before and after lesions of nucleus HVC (proper name) and found that this manipulation reverses the developmental trends in measures of the respiratory pattern.

Disrupting vagal feedback affects birdsong motor control.

Coordination of different motor systems for sound production involves the use of feedback mechanisms. Song production in oscines is a well-established animal model for studying learned vocal behavior. Whereas the online use of auditory feedback has been studied in the songbird model, very little is known about the role of other feedback mechanisms. Auditory feedback is required for the maintenance of stereotyped adult song. In addition, the use of somatosensory feedback to maintain pressure during song has been demonstrated with experimentally induced fluctuations in air sac pressure. Feedback information mediating this response is thought to be routed to the central nervous system via afferent fibers of the vagus nerve. Here, we tested the effects of unilateral vagotomy on the peripheral motor patterns of song production and the acoustic features. Unilateral vagotomy caused a variety of disruptions and alterations to the respiratory pattern of song, some of which affected the acoustic structure of vocalizations. These changes were most pronounced a few days after nerve resection and varied between individuals. In the most extreme cases, the motor gestures of respiration were so severely disrupted that individual song syllables or the song motif were atypically terminated. Acoustic changes also suggest altered use of the two sound generators and upper vocal tract filtering, indicating that the disruption of vagal feedback caused changes to the motor program of all motor systems involved in song production and modification. This evidence for the use of vagal feedback by the song system with disruption of song during the first days after nerve cut provides a contrast to the longer-term effects of auditory feedback disruption. It suggests a significant role for somatosensory feedback that differs from that of auditory feedback.

Hormonal acceleration of song development illuminates motor control mechanism in canaries

In songbirds, the ontogeny of singing behavior shows strong parallels with human speech learning. As in humans, development of learned vocal behavior requires exposure to an acoustic model of species-typical vocalizations, and, subsequently, a sensorimotor practice period after which the vocalization is produced in a stereotyped manner. This requires mastering motor instructions driving the vocal organ and the respiratory system. Recently, it was shown that, in the case of canaries (Serinus canaria), the diverse syllables, constituting the song, are generated with air sac pressure patterns with characteristic shapes, remarkably, those belonging to a very specific mathematical family. Here, we treated juvenile canaries with testosterone at the onset of the sensorimotor practice period. This hormone exposure accelerated the development of song into stereotyped adultlike song. After 20 days of testosterone treatment, subsyringeal air sac pressure patterns of song resembled those produced by adults, while those of untreated control birds of the same age did not. Detailed temporal structure and modulation patterns emerged rapidly with testosterone treatment, and all previously identified categories of adult song were observed. This research shows that the known effect of testosterone on the neural circuits gives rise to the stereotyped categories of respiratory motor gestures. Extensive practice of these motor patterns during the sensorimotor phase is not required for their expression.

Dynamical origin of complex motor patterns

Behavior emerges as the nervous system generates motor patterns in charge of driving a peripheral biomechanical device. For several cases in the animal kingdom, it has been identified that the motor patterns used in order to accomplish a diversity of tasks are the different solutions of a simple, low dimensional nonlinear dynamical system. Yet, motor patterns emerge from the interaction of an enormous number of individual dynamical units. In this work, we study the dynamics of the average activity of a large set of coupled excitable units which are periodically forced. We show that low dimensional, yet non trivial dynamics emerges. As a case study, we analyze the air sac pressure patterns used by domestic canaries during song, which consists of a succession of repetitions of different syllable types. We show that the pressure patterns used to generate different syllables can be approximated by the solutions of the investigated model. In this way, we are capable of integrating different description scales of our problem.

Low-dimensional dynamical model for the diversity of pressure patterns used in canary song

During song production, oscine birds produce large air sac pressure pulses. During those pulses, energy is transferred to labia located at the juncture between the bronchii and the trachea, inducing the high frequency labial oscillations which are responsible for airflow modulations, i.e., the uttered sound. In order to generate diverse syllables, canaries (Serinus canaria) use a set of air sac pressure patterns with characteristic shapes. In this work we show that these different shapes can be approximated by the subharmonic solutions of a forced normal form. This simple model is built from identifying dynamical elements which allow to reproduce the shape of the pressure pattern corresponding to one syllable type. Remarkably, integrating that simple model for other parameters allows to recover the other pressure patterns used during song. Interpreting the diversity of these physiological gestures as subharmonic solutions of a simple nonlinear system allows us to account simultaneously for their morphological features as well as for the syllabic timing and suggests a strategy for the generation of complex motor patterns.

Two-voice complexity from a single side of the syrinx in northern mockingbirdMimus polyglottosvocalizations

The diverse vocal signals of songbirds are produced by highly coordinated motor patterns of syringeal and respiratory muscles. These muscles control separate sound generators on the right and left side of the duplex vocal organ, the syrinx. Whereas most song is under active neural control, there has been a growing interest in a different class of nonlinear vocalizations consisting of frequency jumps, subharmonics, biphonation and deterministic chaos that are also present in the vocal repertoires of many vertebrates, including many birds. These nonlinear phenomena may not require active neural control, depending instead on the intrinsic nonlinear dynamics of the oscillators housed within each side of the syrinx. This study investigates the occurrence of these phenomena in the vocalizations of intact northern mockingbirds Mimus polyglottos. By monitoring respiratory pressure and airflow on each side of the syrinx, we provide the first analysis of the contribution made by each side of the syrinx to the production of nonlinear phenomena and are able to reliably discriminate two-voice vocalizations from potentially similar appearing, unilaterally produced, nonlinear events. We present the first evidence of syringeal lateralization of nonlinear dynamics during bilaterally produced chaotic calls. The occurrence of unilateral nonlinear events was not consistently correlated with fluctuations in air sac pressure or the rate of syringeal airflow. Our data support previous hypotheses for mechanical and acoustic coupling between the two sides of the syrinx. These results help lay a foundation upon which to understand the communicative functions of nonlinear phenomena.