Birdsong requires complex learned motor skills involving the coordination of respiratory, vocal organ and craniomandibular muscle groups. Recent studies have added to our understanding of how these vocal subsystems function and interact during song production. The respiratory rhythm determines the temporal pattern of song. Sound is produced during expiration and each syllable is typically followed by a small inspiration, except at the highest syllable repetition rates when a pattern of pulsatile expiration is used. Both expiration and inspiration are active processes. The oscine vocal organ, the syrinx, contains two separate sound sources at the cranial end of each bronchus, each with independent motor control. Dorsal syringeal muscles regulate the timing of phonation by adducting the sound-generating labia into the air stream. Ventral syringeal muscles have an important role in determining the fundamental frequency of the sound. Di¡erent species use the two sides of their vocal organ in di¡erent ways to achieve the particular acoustic properties of their song. Reversible paralysis of the vocal organ during song learning in young birds reveals that motor practice is particularly important in late plastic song around the time of song crystallization in order for normal adult song to develop. Even in adult crystallized song, expiratory muscles use sensory feedback to make compensatory adjustments to perturbations of respiratory pressure. The stereotyped beak movements that accompany song appear to have a role in suppressing harmonics, particularly at low frequencies.

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1. The role of syringeal muscles in song production, particularly in regulating airflow through the syrinx, was studied in singing brown thrashers (Toxostoma rufum). In nine individuals, muscle activity was recorded electromyographically together with bilateral syringeal airflow, subsyringeal air sac pressure, and vocal output. 2. Dorsal muscles, m. syringealis dorsalis (dS) and m. tracheolateral dorsalis (dTB), are consistently activated during ipsilateral closing of the syrinx or increasing syringeal resistance, suggesting that their main role is adduction. This interpretation is supported by the motor patterns accompanying syllables with rapid oscillations in the rate of airflow. Bursts of electrical activity (2-10 ms) in dorsal muscles are precisely synchronized with decreasing airflow. 3. Electrical activity in m. tracheobronchialis ventralis (vTB) and m. tracheolateralis (TL) is associated with active abduction. An important contribution of vTB is to open the syringeal lumen for short inspirations in between syllables. In syllables with oscillatory flow modulations, vTB bursts show variable alignment with the phase of increasing flow. From this and activity during other syllables, it appears that, during phonation, vTB activity fine tunes the syringeal configuration, which is set by action of the dorsal muscles into a partially constricted state. 4. Activity in the ventral portion of TL, an extrinsic muscle, is strikingly similar to that of vTB, an intrinsic muscle, suggesting that the two muscles have a similar functional role. This supports the notion that intrinsic syringeal muscles of songbirds evolved from extrinsic muscles of nonpasserines. 5. M. syringealis ventralis (vS) does not appear to contribute directly to gating of airflow. Its activity is not consistently correlated with active changes in syringeal resistance. 6. Activity in m. sternotrachealis (ST) is most prominent during rapid changes in the rate of airflow or when switching between expiratory and inspiratory flow, suggesting a role in stabilizing the syringeal framework.

Our current understanding of the soundgenerating mechanism in the songbird vocal organ, the syrinx, is based on indirect evidence and theoretical treatments. The classical avian model of sound production postulates that the medial tympaniform membranes (MTM) are the principal sound generators. We tested the role of the MTM in sound generation and studied the songbird syrinx more directly by filming it endoscopically. After we surgically incapacitated the MTM as a vibratory source, zebra finches and cardinals were not only able to vocalize, but sang nearly normal song. This result shows clearly that the MTM are not the principal sound source. The endoscopic images of the intact songbird syrinx during spontaneous and brain stimulation-induced vocalizations illustrate the dynamics of syringeal reconfiguration before phonation and suggest a different model for sound production. Phonation is initiated by rostrad movement and stretching of the syrinx. At the same time, the syrinx is closed through movement of two soft tissue masses, the medial and lateral labia, into the bronchial lumen. Sound production always is accompanied by vibratory motions of both labia, indicating that these vibrations may be the sound source. However, because of the low temporal resolution of the imaging system, the frequency and phase of labial vibrations could not be assessed in relation to that of the generated sound. Nevertheless, in contrast to the previous model, these observations show that both labia contribute to aperture control and strongly suggest that they play an important role as principal sound generators.The study of vocal communication in songbirds makes significant contributions to various biological disciplines (1, 2) and provides inspiration to related areas, such as linguistics (3, 4). However, unlike the case in human speech (5), the physical mechanism of phonation in birds is poorly understood (6 -7).Sound production in songbirds is commonly believed to involve a constriction of the bronchial lumen by the lateral labium (LL), which, when combined with high subsyringeal air sac pressure and increased air velocity, induces vibrations of the medial tympaniform membranes (MTM) by Bernoulli forces and pressure differences. Support for this interpretation was provided by direct observation of vibrations of the MTM in the excised syrinx during artificially induced sound production (8), indirect anatomical and physiological observations (8-17), acoustic analyses (10, 18), simple syringeal models (18), and theoretical accounts (19)(20)(21)(22)(23). Detailed analyses of the vocal mechanism in non-songbirds (24-29) also provided a framework for the development of this classical model of songbird phonation. However, unlike the case in nonsongbirds (25-27), the predictions of this model for songbirds never have been tested by manipulation of the presumed sound generators, the MTM.Here we report the results of experiments in which the MTM were surgically disabled. Furthermore, we studied the intact songbird syrinx in situ by...

1. The contribution of syringeal muscles to controlling the phonology of song was studied by recording bilateral airflow, subsyringeal air sac pressure, electromyograms (EMGs) of six syringeal muscles, and vocal output in spontaneously singing brown thrashers (Toxostoma rufum). 2. EMG activity in musculus syringealis ventralis (vS), the largest syringeal muscle, increases exponentially with the fundamental frequency of the ipsilaterally generated sound and closely parallels frequency modulation. 3. The EMG activity of other syringeal muscles is also positively correlated with sound frequency, but the amplitude of their EMGs changes only a small amount compared with variation in the amplitude of their EMGs correlated with changing syringeal resistance. The elevated activity in all syringeal muscles during high-frequency sounds may reflect an increased need for structural stability during the strong contractions of the largest syringeal muscle (vS). 4. Several syringeal mechanisms are used to generate amplitude modulation (AM). The most common of these involves modulating the rate of syringeal airflow, through activity by adductor (m. syringealis dorsalis and m. tracheobronchialis dorsalis) and abductor (m. tracheobronchialis ventralis) muscles, which change syringeal resistance, switch sound production from one side of the syrinx to the other, or produce rapid oscillatory flow changes. Variation in the phase relationship between AM and EMG bursts during oscillatory airflow suggests complex biomechanical interaction between antagonistic muscles. 5. AM can also arise from acoustic interactions of two independently generated sounds (beat notes) including cross talk signals between the two syringeal halves. In this latter mechanism, sound generated on one side radiates slightly out of phase with the source from the contralateral side, resulting in lateralized AM generation.

Song production in songbirds is considered a model system for studying learned vocal behavior. As in humans, bird phonation involves three main motor systems (respiration, vocal organ and vocal tract). The avian respiratory mechanism uses pressure regulation in air sacs to ventilate a rigid lung. In songbirds sound is generated with two independently controlled sound sources, which reside in a uniquely avian vocal organ, the syrinx. However, the physical sound generation mechanism in the syrinx shows strong analogies to that in the human larynx. Differences in the functional morphology of the sound producing system between birds and humans require specific motor control patterns. The songbird vocal apparatus is adapted for high speed, suggesting that temporal patterns and fast modulation of sound features are important in acoustic communication. Rapid respiratory patterns determine the coarse temporal structure of song and maintain gas exchange even during very long songs. The respiratory system also contributes to the fine control of airflow. Muscular control of the vocal organ regulates airflow and acoustic features. The upper vocal tract of birds plays a role in filtering the sounds generated in the syrinx, but source-filter interactions may also play a role. The unique morphology and biomechanical system for sound production in birds presents an interesting model for exploring parallels in control mechanisms that give rise to highly convergent physical patterns of sound generation. More comparative work should provide a rich source for our understanding of the evolution of complex sound producing systems.

Many birds perform visual signals during their learned songs, but little is known about the interrelationship between visual and vocal displays. We show here that male brown-headed cowbirds (Molothrus ater) synchronize the most elaborate wing movements of their display with atypically long silent periods in their song, potentially avoiding adverse biomechanical effects on sound production. Furthermore, expiratory effort for song is significantly reduced when cowbirds perform their wing display. These results show a close integration between vocal and visual displays and suggest that constraints and synergistic interactions between the motor patterns of multimodal signals influence the evolution of birdsong.

The beaks of Darwin's finches and other birds are among the best known examples of adaptive evolution. Beak morphology is usually interpreted in relation to its critical role in feeding. However, the beak also plays an important role in preening, which is the first line of defence against harmful ectoparasites such as feather lice, fleas, bugs, flies, ticks and feather mites. Here, we show a feature of the beak specifically adapted for ectoparasite control. Experimental trimming of the tiny (1-2 mm) maxillary overhang of rock pigeons (Columba livia) had no effect on feeding efficiency, yet triggered a dramatic increase in feather lice and the feather damage they cause. The overhang functions by generating a shearing force against the tip of the lower mandible, which moves forward remarkably quickly during preening, at up to 31 times per second. This force damages parasite exoskeletons, significantly enhancing the efficiency of preening for parasite control. Overhangs longer than the natural mean of 1.6 mm break significantly more often than short overhangs. Hence, stabilizing selection will favour overhangs of intermediate length. The adaptive radiation of beak morphology should be re-assessed with both feeding and preening in mind.