Variations in respiratory muscle activity during echolocation when stationary in three species of bat (Microchiroptera: Vespertilionidae).

Echolocating bats use respiratory muscles to power the production of biosonar vocalisations. The physical characteristics of these calls vary among species of bat, and variations also exist in the timing and patterns of respiratory muscle recruitment during echolocation. We recorded electromyograms from the respiratory muscles of three species of bat (Family Vespertilionidae) while the animals vocalised from stationary positions. Activity was recorded consistently from the lateral abdominal muscles (internal abdominal oblique and transversus abdominis) from all calling bats, but we found much variation within and among species. Bats in the family Vespertilionidae devoted longer periods of expiratory muscle activity to each call than did the mormoopid bat Pteronotus parnellii. These differences correlate negatively with the duration of calls. We suggest that morphological adaptations in some bats may facilitate the economic production of echolocation calls at rest.

Morphology of the abdominal wall in the bat, Pteronotus parnellii (Microchiroptera: Mormoopidae): implications for biosonar vocalization.

We investigated the structure of the abdominal wall of Pteronotus parnellii and made comparisons with eight other species of Microchiroptera and one megachiropteran. Similar to other mammals, the abdominal wall of bats consists of the three flank muscles laterally and the m. rectus abdominis ventrally. In Microchiroptera, flank muscles are mostly confined to dorsal portions of the wall. The mm. transversus abdominis and obliquus internus abdominis form the bulk of the wall; the m. obliquus externus is poorly developed. Ventrolaterally, a large portion of the wall is a dense, bilaminar aponeurosis, composed of collagen, elastin, and fibroblasts. The thicker, superficial lamina derives from the mm. obliquus internus and transversus abdominis. The deep lamina is a continuation of the transversalis fascia. Collagen fibers of the two fused laminae are oriented orthogonally, resulting in a resilient, composite fabric. Fascicles of the flank muscles are oriented along the margins of the aponeurosis so that their forces appear to be concentrated onto the aponeurosis. We suggest that this system is adapted for the regulation and generation of intra-abdominal pressure. The abdominal wall of Pteropus, the one megachiropteran examined, lacks the derived aponeurosis and is similar to other mammals. We consider the abdominal wall of Microchiroptera to be analogous to the diaphragma, in that it functions in the regulation of pressure within body cavities and facilitates biosonar vocalization.

Respiratory muscle activity in relation to vocalization in flying bats.

The structure of the thoracic and abdominal walls of Pteronotus parnellii (Microchiroptera: Mormoopidae) was described with respect to their function in respiration and vocalization. We monitored electromyographic activity of respiratory and flight muscles in relation to echolocative vocalization. In flight, signals were telemetered with a small FM transmitter modified to summate the low-frequency myopotentials with biosonar signals from a ceramic-crystal microphone. Recordings were also made from the same bats confined to a small cage. Vocalizations were used as the parameter by which all muscle activities were correlated. A discrete burst of activity in the lateral abdominal wall muscles accompanied each vocalization. Diaphragmatic myopotentials occurred between groups of calls and did not coincide with activity of the abdominal wall or with vocalizations. Flight muscles were not active in resting bats. During flight, vocalizations and the abdominal muscle activity that accompanied them coincided with myopotentials of the pectoralis and serratus ventralis muscles. We propose that contractions of the lateral abdominal wall provide the primary power for the production of intense biosonar vocalization in flying and in stationary bats. In flight, synchronization of vocalization with activity of the pectoralis and serratus ventralis jointly contribute to the pressurization of the thoraco-abdominal cavity. This utilization of pressure that is normally generated in flight facilitates respiration and allows for the production of intense vocalizations with little additional energetic expenditure.

Doppler-shift compensation by the mustached bat: quantitative data.

Quantitative data for Doppler-shift compensation by Pteronotus parnellii parnellii were obtained with a device which propelled the bats at constant velocities over a distance of 12 m. The bats compensated for Doppler shifts at all velocities tested (0.1-5.0 ms-1). The main findings were (1) that compensation was usually accomplished by a progressive lowering of the approximately 61 kHz second harmonic constant-frequency component of emitted sounds in small frequency steps (93 +/- 72 Hz); (2) that the time needed to reach a steady compensation level averaged 514 +/- 230 ms and the number of pulses required to reach full compensation averaged 10.78 +/- 5.16; (3) that the animals compensated to hold the echo (reference) frequency at a value that was slightly higher than the resting frequency and slightly lower than the cochlear resonance frequency; (4) that reference frequency varied as a function of velocity, the higher the velocity of the animal, the higher was the reference frequency (slope 55 Hz m-1s-2); and (5) that the mean reference frequency was always an undercompensation. The average amount of undercompensation was 15.8%. There was a significant difference (P < or = 0.005) in Doppler-shift compensation data collected at velocities that differed by 0.1 ms-1. A velocity difference of 0.1 ms-1 corresponds to a Doppler-shift difference of about 35 Hz in the approximately 61 kHz signals reaching the ear.

Ultrasonic vocalizations of flying bats monitored by radiotelemetry.

Ultrasonic vocalizations of flying bats were effectively monitored with radiotelemetry. We describe a device light enough to be carried by an 11 g bat for periods of up to 1 h. It transmitted signals adequate for fine frequency analysis within a range of approximately 3 m. Telemetry permitted the recording of constant-frequency pulses free from flight-induced Doppler shifts and without time delays. The difference in frequency between telemetered signals and the same signals detected by a remote microphone was used to calculate velocity and Doppler shifts. Pulse emission behavior of Pteronotus parnellii in flight was compared with simulated flight on a pendulum. The data showed significant differences in echo bandwidths, constant-frequency pulse durations and interpulse intervals. In flight, pulses and interpulse intervals tended to be shorter and bats maintained echo frequencies within a significantly narrower band. Phases of echolocation that characterized the approach to a target were clearly evident in flight, but not during pendulum swings. Differences in pulse durations and interpulse intervals may be correlated with the integration of wingbeat, respiration and vocalization. The absence of wing motion in simulated flight changes this integration.