SUMMARY Bird vocalizations resonate as they propagate through a relatively long trachea and radiate out from the oral cavity. Several studies have described the dynamics with which birds actively vary beak gape while singing and it has been hypothesized that birds vary beak gape as a mechanism for varying vocal tract resonances. Nevertheless, few studies have attempted to quantify the effects of beak gape on vocal tract resonances. We replaced eastern towhee, Pipilo erythrophthalmus L. , syringes with a small speaker and obtained recordings of frequency sweeps while rotating each subject in a horizontal plane aligned with either the maxilla or mandible. We describe vocal tract resonances as well as how sound radiates as a function of beak gape. Results are inconsistent with the hypothesis that songbirds vary beak gape as a mechanism for `tracking' fundamental frequencies in vocalizations. Instead, decreases in beak gape seem to attenuate resonances that occur between ∼4 and 7.5 kHz. We propose that songbirds vary beak gape as a mechanism for excluding and/or concentrating energy within at least two distinct sound frequency channels.
SUMMARY Ring doves vocalize with their beaks and nostrils closed, exhaling into inflatable chambers in the head and neck region. The source sound produced at the syrinx contains a fundamental frequency with prominent second and third harmonic overtones, but these harmonics are filtered out of the emitted signal. We show by cineradiography that the upper esophagus, oral and nasal cavities collect the expired air during vocalization and that the inflated esophagus becomes part of the suprasyringeal vocal tract. The level of the second and third harmonics, relative to the fundamental frequency( f 0 ), is reduced in the esophagus and emitted vocalization compared with in the trachea, although these harmonics are still considerably higher in the esophagus than in the emitted signal. When the esophagus is prevented from fully inflating, there is a pronounced increase in the level of higher harmonics in the emitted vocalization. Our data suggest that the trachea and esophagus act in series as acoustically separate compartments attenuating harmonics by different mechanisms. We hypothesize that the trachea behaves as a tube closed at the syringeal end and with a variable, restricted opening at the glottal end that lowers the tracheal first resonance to match the f 0 of the coo. The inflated esophagus may function as a Helmholtz resonator in which the elastic walls form the vibrating mass. Such a resonator could support the f 0 over a range of inflated volumes.
SUMMARY Birdsong assumes its complex and specific forms by the modulation of phonation in frequency and time domains. The organization of control mechanisms and intrinsic properties causing such modulation have been studied in songbirds but much less so in non-songbirds, the songs of which are often regarded as relatively simple. We examined mechanisms of frequency and amplitude modulation of phonation in ring doves Streptopelia risoria ,which are non-songbirds. Spontaneous coo vocalizations were recorded together with concurrent pressure patterns in two different air sacs and air flow rate in the trachea. The results show that amplitude modulation is the result of the cyclic opening and closure of a valve instead of fluctuations in driving pressure, as is the current explanation. Frequency modulation is more complex than previously recognized and consists of gradual, continuous time–frequency patterns, punctuated by instantaneous frequency jumps. Gradual frequency modulation patterns correspond to pressure variation in the interclavicular air sac but not to pressure variation in the cranial thoracic air sac or air flow rate variation in the trachea. The cause of abrupt jumps in frequency has not been identified but can be explained on the basis of intrinsic properties of the vocal organ. Air sac pressure variation as a mechanism for frequency modulation contrasts with the specialized syringeal musculature of songbirds and may explain why the fundamental frequency in non-songbird vocalizations is generally modulated within a limited frequency range.