Coding of Small Sinusoidal Frequency and Amplitude Modulations in the Inferior Colliculus of 'CF-FM' Bat, Rhinolophus Ferrumequinum

Single neurons in the inferior colliculus of the Greater Horseshoe bat, Rhinolophus ferrumequinum, showed two broad categories of response patterns to sinusoidally frequency (SFM) or amplitude (SAM) modulated stimuli. Tonic responding cells (best excitatory frequency (BEF) between 10 and 90 kHz) showed a rough sinusoidal modulation of the discharge pattern to SFM. Transient responding neurons, generally showing on- or off-responses to pure tones, (BEF between 65 and 88 kHz), displayed highly synchronized discharge patterns to SFM-cycles (Fig. 1). Modulation rates between 20 and 100 Hz were most effective and some neurons encoded modulation rates up to 350 Hz (Figs. 2 and 3). The SFM responses were best synchronized to the modulation envelope for center frequencies in the upper portion of the tuning curve (Figs. 4 and 5). Sharply tuned neurons with BEF around 80 kHz had the lowest threshold for modulation depth (± 10 Hz or 0.025%) (Fig. 6). In general, SAMs evoked the same type of response patterns and were encoded down to modulation index of 3% (Fig. 7). The fine frequency and amplitude discriminations for periodical modulations by collicular neurons is discussed as related to the detection and discrimination performance of bats, when preying on flying insects in clustered surroundings

Natural ultrasonic echoes from wing beating insects are encoded by collicular neurons in the CF-FM bat, Rhinolophus f errumequinum

1. Acoustic reflections from a wing beating moth to an 80 kHz ultrasonic signal were recorded from six different incident angles and analyzed in spectral and time domains. The recorded echoes as well as independent components of amplitude and frequency modulations of the echoes were employed as acoustic stimuli during single unit studies. 2. The responses of single inferior colliculus neurons to these stimuli were recorded from four horseshoe bats,Rhinolophus ferrumequinum, a species which uses a long constant frequency (CF) sound with a final frequency modulated (FM) sweep during echolocation. All neurons responding to wing beat echoes reliably encoded the fundamental wing beat frequency as well as the more refined frequency and amplitude modulations. 3. These neurons may provide the bat a neural mechanism to detect periodically moving targets against a cluttered background and also to discriminate various insect species on the basis of their wing beat patterns

Neural control of vocalization in bats: mapping of brainstem areas with electrical microstimulation eliciting species-specific echolocation calls in the rufous horseshoe bat

1. The functional role of brainstem structures in the emission of echolocation calls was investigated in the rufous horseshoe bat, Rhinolophus rouxi, with electrical low-current microstimulation procedures. 2. Vocalizations without temporal and/or spectral distortions could be consistently elicited at low threshold currents (typically below 10 A) within three clearly circumscribed brainstem areas, namely, the deep layers and ventral parts of the intermediate layers of the superior colliculus (SC), the deep mesencephalic nucleus (NMP) in the dorsal and lateral midbrain reticular formation and in a distinct area medial to the rostral parts of the dorsal nucleus of the lateral lemniscus. The mean latencies in the three vocal areas between the start of the electrical stimulus and the elicited vocalizations were 47 msec, 38 msec and 31 msec, respectively. 3. In pontine regions and the cuneiform nucleus adjacent to these three vocal areas, thresholds for eliciting vocalizations were also low, but the vocalizations showed temporal and/or spectral distortions and were often accompanied or followed by arousal of the animal. 4. Stimulus intensity systematically influenced vocalization parameters at only a few brain sites. In the caudo-ventra1 portions of the deep superior colliculus the sound pressure level of the vocalizations systematically increased with stimulus intensity. Bursts of multiple vocalizations were induced at locations ventral to the rostral parts of the cuneiform nucleus. No stimulus-intensity dependent frequency changes of the emitted vocalizations were observed. 5. The respiratory cycle was synchronized to the electrical stimuli in all regions where vocalizations could be elicited as well as in more ventrally and medially adjacent areas not yielding vocalizations on stimulation. 6. The possible functional involvement of the vocal structures in the audio-vocal feedback system of the Dopplercompensating horseshoe bat is discussed

Spectral and temporal gating mechanisms enhance the clutter rejection in the echolocating bat, Rhinolophus rouxi

Doppler shift compensation behaviour in horseshoe bats, Rhinolophus rouxi, was used to test the interference of pure tones and narrow band noise with compensation performance. The distortions in Doppler shift compensation to sinusoidally frequency shifted echoes (modulation frequency: 0.1 Hz, maximum frequency shift: 3 kHz) consisted of a reduced compensation amplitude and/or a shift of the emitted frequency to lower frequencies (Fig. 1). Pure tones at frequencies between 200 and 900 Hz above the bat's resting frequency (RF) disturbed the Doppler shift compensation, with a maximum of intererence between 400 and 550 Hz (Fig. 2). Minimum duration of pure tones for interference was 20 ms and durations above 40 ms were most effective (Fig. 3). Interfering pure tones arriving later than about 10 ms after the onset of the echolocation call showed markedly reduced interference (Fig. 4). Doppler shift compensation was affected by pure tones at the optimum interfering frequency with sound pressure levels down to –48 dB rel the intensity level of the emitted call (Figs. 5, 6). Narrow bandwidth noise (bandwidth from ± 100 Hz to ± 800 Hz) disturbed Doppler shift compensation at carrier frequencies between –250 Hz below and 800 Hz above RF with a maximum of interference between 250 and 500 Hz above resting frequency (Fig. 7). The duration and delay of the noise had similar influences on interference with Doppler shift compensation as did pure tones (Figs. 8, 9). Intensity dependence for noise interference was more variable than for pure tones (-32 dB to -45 dB rel emitted sound pressure level, Fig. 10). The temporal and spectral gating in Doppler shift compensation behaviour is discussed as an effective mechanism for clutter rejection by improving the processing of frequency and amplitude transients in the echoes of horseshoe bats

Storage of Doppler-Shift Information in the Echolocation System of the "CF-FM"-Bat, Rhinolophus ferrumequinum

The greater horseshoe bat (Rhinolophus ferrumequinum) emits echolocation sounds consisting of a long constant-frequency (CF) component preceeded and followed by a short frequency-modulated (FM) component. When an echo returns with an upward Doppler-shift, the bat compensates for the frequency-shift by lowering the emitted frequency in the subsequent orientation sounds and stabilizes the echo image. The bat can accurately store frequency-shift information during silent periods of at least several minutes. The stored frequency-shift information is not affected by tone bursts delivered during silent periods without an overlap with an emitted orientation sound. The system for storage of Doppler-shift information has properties similar to a sample and hold circuit with sampling at vocalization time and with a rather flat slewing rate for the stored frequency information

Laryngeal Nerve Activity During Pulse Emission in the CF-FM Bat, Rhinolophus ferrumequinum. II. The Recurrent Laryngeal Nerve

The activity of the recurrent laryngeal nerve (RLN) was recorded in the greater horseshoe bat,Rhinolophus ferrumequinum. Respiration, vocalization and nerve discharges were monitored while vocalizations were elicted by stimulation of the central gray matter. This stimulation evoked either expiration or expiration plus vocalization depending on the stimulus strength. When vocalization occurred it always took place during expiration. Recordings from the RLN during respiration showed activity during the inspiration phase, but when vocalization occurred there was activity during inspiration and expiration. These results are consistent with the view that the RLN innervates muscles which control the opening and closing of the glottis. During vocalization the vocal folds are closely approximated and the discharge patterns of the nerve suggests that it controls the muscles which start and end each pulse