In human speech, the sound generated by the larynx is modified by articulatory movements of the upper vocal tract, which acts as a variable resonant filter concentrating energy near particular frequencies, or formants, essential in speech recognition. Despite its potential importance in vocal communication, little is known about the presence of tunable vocal tract filters in other vertebrates. The tonal quality of much birdsong, in which upper harmonics have relatively little energy, depends on filtering of the vocal source, but the nature of this filter is controversial. Current hypotheses treat the songbird vocal tract as a rigid tube with a resonance that is modulated by the end-correction of a variable beak opening. Through x-ray cinematography of singing birds, we show that birdsong is accompanied by cyclical movements of the hyoid skeleton and changes in the diameter of the cranial end of the esophagus that maintain an inverse relationship between the volume of the oropharyngeal cavity and esophagus and the song's fundamental frequency. A computational acoustic model indicates that this song-related motor pattern tunes the major resonance of the oropharyngeal-esophageal cavity to actively track the song's fundamental frequency.bioacoustics ͉ hyoid motor pattern ͉ larynx ͉ beak gape ͉ vocal tract filter

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.

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.

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.

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Carollia perspicillata (Phyllostomidae) is a frugivorous bat that emits low-intensity, broadband, frequency-modulated echolocation pulses through nostrils surrounded by a noseleaf. The emission pattern of this bat is of interest because the ratio between the nostril spacing and the emitted wavelength varies during the pulse, causing complex interference patterns in the horizontal dimension. Sound pressures around the bat were measured using a movable microphone and were referenced to those at a stationary microphone positioned directly in front of the animal. Interference between the nostrils was confirmed by blocking one nostril, which eliminated sidelobes and minima in the emission pattern, and by comparison of real emission patterns with simple computer models. The positions of minima in the patterns indicate effective nostril spacings of over a half-wavelength. Displacement of the dorsal lancet of the noseleaf demonstrated that this structure directs sound in the vertical dimension.

Several groups of mammals such as bats, dolphins and whales are known to produce ultrasonic signals which are used for navigation and hunting by means of echolocation, as well as for communication. In contrast, frogs and birds produce sounds during night- and day-time hours that are audible to humans; their sounds are so pervasive that together with those of insects, they are considered the primary sounds of nature. Here we show that an Old World frog (Amolops tormotus) and an oscine songbird (Abroscopus albogularis) living near noisy streams reliably produce acoustic signals that contain prominent ultrasonic harmonics. Our findings provide the first evidence that anurans and passerines are capable of generating tonal ultrasonic call components and should stimulate the quest for additional ultrasonic species.