Sound and vibration sensitivity of VIIIth nerve fibers in the grassfrog, Rana temporaria

We have studied the sound and vibration sensitivity of 164 amphibian papilla fibers in the VIIIth nerve of the grassfrog, Rana temporaria. The VIIIth nerve was exposed using a dorsal approach. The frogs were placed in a natural sitting posture and stimulated by free-field sound. Furthermore, the animals were stimulated with dorso-ventral vibrations, and the sound-induced vertical vibrations in the setup could be canceled by emitting vibrations in antiphase from the vibration exciter. All low-frequency fibers responded to both sound and vibration with sound thresholds from 23 dB SPL and vibration thresholds from 0.02 cm/s². The sound and vibration sensitivity was compared for each fiber using the offset between the rate-level curves for sound and vibration stimulation as a measure of relative vibration sensitivity. When measured in this way relative vibration sensitivity decreases with frequency from 42 dB at 100 Hz to 25 dB at 400 Hz. Since sound thresholds decrease from 72 dB SPL at 100 Hz to 50 dB SPL at 400 Hz the decrease in relative vibration sensitivity reflects an increase in sound sensitivity with frequency, probably due to enhanced tympanic sensitivity at higher frequencies. In contrast, absolute vibration sensitivity is constant in most of the frequency range studied. Only small effects result from the cancellation of sound-induced vibrations. The reason for this probably is that the maximal induced vibrations in the present setup are 6–10 dB below the fibers' vibration threshold at the threshold for sound. However, these results are only valid for the present physical configuration of the setup and the high vibration-sensitivities of the fibers warrant caution whenever the auditory fibers are stimulated with free-field sound. Thus, the experiments suggest that the low-frequency sound sensitivity is not caused by sound-induced vertical vibrations. Instead, the low-frequency sound sensitivity is either tympanic or mediated through bone conduction or sound-induced pulsations of the lungs.Abbreviations AP amphibian papilla - BF best frequency - PST peristimulus time     

Directional hearing in the barn owl (Tyto alba)

Summary The acoustical properties of the external ear of the barn owl (Tyto alba) were studied by measuring sound pressure in the ear canal and outer ear cavity. Under normal conditions, pressure amplification by the external ear reaches about 20 dB between 3–9 kHz but decreases sharply above 10 kHz. The acoustic gain curve of the outer ear cavity alone is close to that of a finite-length exponential horn between 1.2–13 kHz with maximum gain reaching 20 dB between 5–9 kHz. Pressure gain by the facial ruff produces a maximum of 12 dB between 5–8 kHz and decreases rapidly above 9 kHz.The directional sensitivity of the external ear was obtained from pressure measurements in the ear canal. Directivity of the major lobe is explained, to a first approximation, by the sound diffraction properties of a circular aperture. Aperture size is based on the average radius (30 mm) of the open face of the ruff. Above 5 kHz, the external ear becomes highly directional and there is a 26° disparity in elevation between the acoustic axis of the left and right ear. In azimuth, directivity patterns are relocated closer to the midline as frequency increases and the acoustic axis moves at a rate of 20°/octave between 2–13 kHz. Movement of the axis can be explained, to a first approximation, by the acoustical diffraction properties of an obliquely truncated horn, due to the asymmetrical shape of the outer ear cavity.The directional sensitivity of the barn owl ear was studied by recording cochlear microphonic (CM) potentials from the round window membrane. Between 3–9 kHz, CM directivity patterns are clearly different to the directivity patterns of the external ear; CM directionality is abruptly lost above 10 kHz. Above 5 kHz, CM directivity patterns are characterized by an elongated major lobe containing the CM axis, forming a tilted band of high amplitude but low directionality (CM axial plane), closely bordered by minima or nulls. The highest directionality is found in theCM directional plane, approximately perpendicular to the CM axial plane. The left and right ear axial planes are symmetrical about the interaural midline (tilted 12° to the right of the midline of the head) and inclined by an average of 60° to the left and right respectively. In azimuth, the CM axis moves towards the midline at a rate of 37°/octave as frequency increases from 2–9 kHz, crossing into contralateral space near 7 kHz. In the CM directional plane, the directivity of the major lobe suggests that a pressure gradient may occur at the TM. The region of frontal space mapped by movement of the CM axis in azimuth closely matches the angle of sound incidence which would be expected to produce the maximum driving pressure at the TM. It is suggested that acoustical interference at the TM results from sound transmission through the interaural canal and therefore the ear is inherently directional. It is proposed that ear directionality in the barn owl may be explained by the combined effect of sound diffraction by the outer ear cavity and a pressure gradient at the TM.Abbreviations CM cochlear microphonic - RMS root mean square - SPL sound pressure level - TM tympanic membrane     

Specialization for underwater hearing by the tympanic middle ear of the turtle, Trachemys scripta elegans

Turtles, like other amphibious animals, face a trade-off between terrestrial and aquatic hearing. We used laser vibrometry and auditory brainstem responses to measure their sensitivity to vibration stimuli and to airborne versus underwater sound. Turtles are most sensitive to sound underwater, and their sensitivity depends on the large middle ear, which has a compliant tympanic disc attached to the columella. Behind the disc, the middle ear is a large air-filled cavity with a volume of approximately 0.5 ml and a resonance frequency of approximately 500 Hz underwater. Laser vibrometry measurements underwater showed peak vibrations at 500-600 Hz with a maximum of 300 μm s(-1) Pa(-1), approximately 100 times more than the surrounding water. In air, the auditory brainstem response audiogram showed a best sensitivity to sound of 300-500 Hz. Audiograms before and after removing the skin covering reveal that the cartilaginous tympanic disc shows unchanged sensitivity, indicating that the tympanic disc, and not the overlying skin, is the key sound receiver. If air and water thresholds are compared in terms of sound intensity, thresholds in water are approximately 20-30 dB lower than in air. Therefore, this tympanic ear is specialized for underwater hearing, most probably because sound-induced pulsations of the air in the middle ear cavity drive the tympanic disc.     

The directionality of the ear of the pigeon (Columba livia)

Summary The directionality of cochlear microphonic potentials in the azimuthal plane was investigated in the pigeon (Columba livia), using acoustic free-field stimulation (pure tones of 0.25–6 kHz).At high frequencies in the pigeon's hearing range (4–6 kHz), changing azimuth resulted in a maximum change of the cochlear microphonic amplitude by about 20 dB (SPL). The directionality decreased clearly with decreasing frequency.Acoustic blocking of the contralateral ear canal could reduce the directional sensitivity of the ipsilateral ear by maximally 8 dB. This indicates a significant sound transmission through the bird's interaural pathways. However, the magnitude of these effects compared to those obtained by sound diffraction (maximum > 15 dB) suggests that pressure gradients at the tympanic membrane are only of subordinate importance for the generation of directional cues.The comparison of interaural intensity differences with previous behavioral results confirms the hypothesis that interaural intensity difference is the primary directional cue of azimuthal sound localization in the high-frequency range (2–6 kHz).Abbreviations CM cochlear microphonic potential - IID interaural intensity difference - IID-MRA minimum resolvable angle calculated from interaural intensity difference - MRA minimum resolvable angle - OTD interaural ongoing time difference - RMS root mean square - SPL sound pressure level     

Auditory brainstem responses to airborne sounds in the aquatic frog Xenopus laevis: correlation with middle ear characteristics

In this study we recorded auditory brainstem responses to airborne sounds to determine the hearing sensitivity of Xenopus laevis frogs and correlated their hearing profiles with middle ear characteristics. In newly metamorphosed frogs (body mass 0.5–0.76 gm, snout-vent length 17–20 mm) best hearing sensitivities were measured in the 2.4–2.8 kHz range, whereas optimal hearing sensitivity of older adults (body mass 18–90 gm; snout-vent length 57–100 mm) ranged from 1.0 to 1.2 kHz. Middle ear volumes reconstructed from serial sections showed approximate volume of 0.002 cc and 0.04–0.07 cc in newly metamorphosed and older frogs, respectively. This inverse frequency–volume relationship is consistent with the properties of an acoustic resonator indicating that differences in best hearing sensitivity are at least in part correlated to variation in middle ear volumes for airborne sounds. These results are consistent with peak frequency vibrational velocity profiles of Xenopus tympanic disk that have been shown to be dependent on underlying middle ear volumes and corroborate the occurrence of peak amplitudes of otoacoustic emissions in the 1.0–1.2 kHz region in adult Xenopus frogs.     

Cartilage Conduction Is Characterized by Vibrations of the Cartilaginous Portion of the Ear Canal

Cartilage conduction (CC) is a new form of sound transmission which is induced by a transducer being placed on the aural cartilage. Although the conventional forms of sound transmission to the cochlea are classified into air or bone conduction (AC or BC), previous study demonstrates that CC is not classified into AC or BC (Laryngoscope 124: 1214–1219). Next interesting issue is whether CC is a hybrid of AC and BC. Seven volunteers with normal hearing participated in this experiment. The threshold-shifts by water injection in the ear canal were measured. AC, BC, and CC thresholds at 0.5–4 kHz were measured in the 0%-, 40%-, and 80%-water injection conditions. In addition, CC thresholds were also measured for the 20%-, 60%-, 100%-, and overflowing-water injection conditions. The contributions of the vibrations of the cartilaginous portion were evaluated by the threshold-shifts. For AC and BC, the threshold-shifts by the water injection were 22.6–53.3 dB and within 14.9 dB at the frequency of 0.5–4 kHz, respectively. For CC, when the water was filled within the bony portion, the thresholds were elevated to the same degree as AC. When the water was additionally injected to reach the cartilaginous portion, the thresholds at 0.5 and 1 kHz dramatically decreased by 27.4 and 27.5 dB, respectively. In addition, despite blocking AC by the injected water, the CC thresholds in force level were remarkably lower than those for BC. The vibration of the cartilaginous portion contributes to the sound transmission, particularly in the low frequency range. Although the airborne sound is radiated into the ear canal in both BC and CC, the mechanism underlying its generation is different between them. CC generates airborne sound in the canal more efficiently than BC. The current findings suggest that CC is not a hybrid of AC and BC.     

Directionality of the tympanal vibrations in a cicada: a biophysical analysis

1. Laser vibrometry and acoustic measurements were used to study the biophysics of directional hearing in males and females of a cicada, in which most of the male tympanum is covered by thick, water filled tissue “pads”. 2. In females, the tympanal vibrations are very dependent on the direction of sound incidence in the entire frequency range 1–20 kHz, and especially at the main frequencies of the calling song (3–7 kHz). At frequencies up to 10 kHz, the directionality disappears if the contralateral tympanum, metathoracic spiracle, and folded membrane are blocked with Vaseline. This suggests some pressure-difference receiver properties in the ear. 3. In males, the tympanal vibrations depend on the direction of sound incidence only within narrow frequency bands (around 1.8 kHz and at 6–7 kHz). At frequencies above 10–12 kHz, the directionality appears to be determined by diffraction, and the ear seems to work as a pressure receiver. The peak in directionality at 6–7 kHz disappears when the contralateral timbal, but not the tympanum, is covered. Covering the thin ventral abdominal wall causes the peak around 1.8 kHz to disappear. 4. Most observed tympanal directionalities, except around 1.8 kHz in males, are well predicted from measured transmissions of sound through the body and measured values of sound amplitude and phase at the ears at various directions of sound incidence. Accepted: 18 October 1996     

The influence of muscles activation on the dynamical behaviour of the tympano-ossicular system of the middle ear

The human ear is a complex biomechanical system and is divided into three parts: outer, middle and inner ear. The middle ear is formed by ossicles (malleus, incus and stapes), ligaments, muscles and tendons, which transfers sound vibrations from the eardrum to the inner ear, linking with mastoid and Eustachian tube. In this work, a finite element modelling of the tympano-ossicular system of the middle ear was developed. A dynamic study based on a structural response to harmonic vibrations, for a sound pressure level (SPL) of 110, 120 and 130 dB SPL applied in the eardrum, is presented. The connection between the ossicles is made using a contact formulation. The model includes the different ligaments considering its hyperelastic behaviour. The activation of the muscles is based on the constitutive model proposed by previous work. The harmonic responses of displacement and pressure obtained on the stapes footplate, for a frequency range between 100 Hz and 10 kHz, are obtained simulating the muscle activation. The results are compared considering the passive and active states. The results are discussed and they are in accordance with audiological data published with reference to the effects of the middle ear muscles contraction.     

Stridulation and hearing in the noctuid mothThecophora fovea (Tr.)

Summary MaleThecophora fovea (Tr.) (Noctuidae) sing continuously for several minutes by rubbing the 1. tarsal segment of the metathoracic leg against a stridulatory swelling on the hindwing. In Northern Yugoslavia (Slovenia) the males emerge in late October and start stridulating about a week later when the females emerge.The sounds are pulse trains consisting of 10–12 ms long sound pulses with main energy around 32 kHz and a PRR of 20 pulses/s. The mechanics of the sound producing apparatus was studied by activating the stridulatory swelling with short sound impulses. The impulse response of the swelling was recorded by laser vibrometry and amplitude spectra of the vibrations showed maximum velocities between 25 and 35 kHz. Hence, it seems likely that the stridulatory swelling is driven as a mechanical oscillator with a resonance frequency which determines the carrier frequency of the sounds.Audiograms of both males and females showed peak sensitivities at 25–30 kHz. The median threshold at the BF was 36 dB SPL. The peak intensity of the sound pulses was 83 dB SPL at 1 m, which should enable the moths to hear each other at distances of around 30 m. Therefore sound production inT. fovea might function in long distance calling. It is argued thatT. fovea can survive making such a noise in spite of being palatable to bats because it flies so late in the year that it is temporally isolated from bats.Abbreviations PRR pulse repetition rate - SPL sound pressure level - BF best frequency     

Sound radiation in a cicada: the role of different structures

Sound radiation was studied in males of Tympanistalna gastrica St»l during a spontaneous song with the characteristics of the conspecific calling song, which was elicited as an after effect of brain stimulation. The song contains two different kinds of sound pulses: 1) loud clicks and 2) soft pulses, presenting different spectra.The timbals, abdomen, tympana, folded membranes and opercula were tested as potential radiators of the song. The experiments included: 1) probe microphone measurements of the spectra of loud clicks and soft pulses in several positions around the animal and close to the body surface; 2) measurements of the spectra before and after covering with vaseline different structures that might be relevant to the radiation of the song, and manipulations of the size and shape of the abdominal and thoracic portions of the tracheal air sac; 3) laser vibrometry measurements in different parts of the body, both during singing and external sound stimulation.The data obtained demonstrate that several structures contribute differently to the radiation of clicks and soft pulses: 1) The timbals are the main radiators at frequencies around the dominant spectral peak, 10–11 kHz in clicks and 12–13 kHz in soft pulses; 2) The tympana are important in radiation of frequencies below and above the timbal peak, especially during the generation of soft pulses; 3) The abdomen is more activated during the generation of clicks, and is more important in the radiation of low frequencies around 5 kHz.Manipulations of the body cavities showed that neither the thoracic nor the abdominal portions of the air sac are critical for the song tuning. The large abdominal cavity do not seem to work as a Helmholtz resonator. We found no evidence that resonances inside this cavity should play an important role in enhancing sound radiation in T. gastrica.     

Physiology and tonotopic organization of auditory receptors in the cricketGryllus bimaculatus DeGeer

Summary Physiological recordings were obtained from identified receptors in the tympanal organ ofGryllus bimaculatus. By immersing the prothoracic leg in Ringer solution and removing the anterior tympanic membrane the auditory receptors were exposed without significantly altering the frequency response of the auditory organ (Fig. 1). Each receptor was tuned to a specific sound frequency. For sound frequencies below this characteristic frequency the roll-off in sensitivity decreased from 20–30 dB/octave to 10–15 dB/octave as the characteristic frequency of receptors increased from 3–11 kHz (Fig. 4A). For each individual receptor the slope, dynamic range and maximum spike response were similar for different sound frequencies (Fig. 9A). The receptors were tonotopically organized with the characteristic frequency of the receptors increasing from the proximal to the distal end of the array (Figs. 5, 6). Several receptors had characteristic frequencies of 5 kHz. These receptors were divided into two groups on the basis of their maximum spike response produced in response to pure tones of increasing intensity (Fig. 7). Independent of the tuning of the receptor no two-tone inhibition was observed in the periphery, thus confirming that such interactions are a property of central integration.     

Middle Ear Cavity Morphology Is Consistent with an Aquatic Origin for Testudines

The position of testudines in vertebrate phylogeny is being re-evaluated. At present, testudine morphological and molecular data conflict when reconstructing phylogenetic relationships. Complicating matters, the ecological niche of stem testudines is ambiguous. To understand how turtles have evolved to hear in different environments, we examined middle ear morphology and scaling in most extant families, as well as some extinct species, using 3-dimensional reconstructions from micro magnetic resonance (MR) and submillimeter computed tomography (CT) scans. All families of testudines exhibited a similar shape of the bony structure of the middle ear cavity, with the tympanic disk located on the rostrolateral edge of the cavity. Sea Turtles have additional soft tissue that fills the middle ear cavity to varying degrees. When the middle ear cavity is modeled as an air-filled sphere of the same volume resonating in an underwater sound field, the calculated resonances for the volumes of the middle ear cavities largely fell within testudine hearing ranges. Although there were some differences in morphology, there were no statistically significant differences in the scaling of the volume of the bony middle ear cavity with head size among groups when categorized by phylogeny and ecology. Because the cavity is predicted to resonate underwater within the testudine hearing range, the data support the hypothesis of an aquatic origin for testudines, and function of the middle ear cavity in underwater sound detection.     

Evoked potentials of the auditory cortex of the porpoise,Phocoena phocoena

Summary Evoked potential (EP) recordings in the auditory cortex of the porpoise,Phocoena phocoena, were used to obtain data characterizing the auditory perception of this dolphin. The frequency threshold curves showed that the lowest EP thresholds were within 120–130 kHz. An additional sensitivity peak was observed between 20 and 30 kHz. The minimal EP threshold to noise burst was 3·10^–4–10^/s-3 Pa. The threshold for response to modulations in sound intensity was below 0.5 dB and about 0.1% for frequency modulations. Special attention was paid to the dependence of the auditory cortex EP on the temporal parameters of the acoustic stimuli: sound burst duration, rise time, and repetition rate. The data indicate that the porpoise auditory cortex is adapted to detect ultrasonic, brief, fast rising, and closely spaced sounds like echolocating clicks.Abbreviation EP evoked potential     

Ossicular differentiation of airborne and seismic stimuli in the Cape golden mole (Chrysochloris asiatica)

Comparison between the middle ear anatomy of the Cape golden mole (Chrysochloris asiatica), which exhibits a club-shaped malleus head, and the Desert golden mole (Eremitalpa granti), with a ball-shaped malleus head, suggests differences in sensitivity to airborne sound. Scanning laser Doppler vibrometric measurements of the ossicular behavior in response to both vibration and airborne sound were made in C. asiatica. Two distinct vibrational modes were observed. In response to low-frequency vibration (70–200 Hz), the malleus oscillates about the ligament of the short process of the incus, whereas in response to high-frequency airborne sound (1–6 kHz) the ossicular chain rotates about the long axis of malleus. It is proposed that the club-shaped malleus head in C. asiatica constitutes an adaptation towards bimodal hearing—sensitivity to substrate vibrations and airborne sound. Possible functional differences between these two middle ear types are discussed.     

Physics of directional hearing in the cricket Gryllus bimaculatus

In the cricket ear, sound acts on the external surface of the tympanum and also reaches the inner surface after travelling in at least three pathways in the tracheal system. We have determined the transmission gain of the three internal sound pathways; that is, the change of amplitude and phase angle from the entrances of the tracheal system to the inner surface of the tympanum. In addition, we have measured the diffraction and time of arrival of sound at the ear and at the three entrances at various directions of sound incidence. By combining these data we have calculated how the total driving force at the tympanum depends on the direction of sound. The results are in reasonable agreement with the directionality of the tympanal vibrations as determined with laser vibrometry.At the frequency of the calling song (4.7 kHz), the direction of the sound has little effect on the amplitudes of the sounds acting on the tympanum, but large effects on their phase angles, especially of the sound waves entering the tracheal system at the contralateral side of the body. The master parameter for causing the directionality of the ear in the forward direction is the sound wave entering the contralateral thoracic spiracle. The phase of this sound component may change by 130–140° with sound direction. The transmission of sound from the contralateral inputs is dominated by a very selective high-pass filter, and large changes in amplitude and phase are seen in the transmitted sounds when the sound frequency changes from 4 to 5 kHz. The directionality is therefore very dependent on sound frequency.The transmission gains vary considerably in different individuals, and much variation was also found in the directional patterns of the ears, especially in the effects of sounds from contralateral directions. However, the directional pattern in the frontal direction is quite robust (at least 5 dB difference between the 330° and 30° directions), so these variations have only little effect on how well the individual animals can approach singing conspecifics.Abbreviations CS contralateral spiracle - CT contralateral tympanum - IS ipsilateral spiracle - IT ipsilateral tympanum - P the vectorial sum of the sounds acting on the tympanum     

Specializations for aerial hawking in the echolocation system of<Emphasis Type="Italic"> Molossus molossus</Emphasis> (Molossidae,Chiroptera)

While searching for prey, Molossus molossus broadcasts narrow-band calls of 11.42 ms organized in pairs of pulses that alternate in frequency. The first signal of the pair is at 34.5 kHz, the second at 39.6 kHz. Pairs of calls with changing frequencies were only emitted when the interpulse intervals were below 200 ms. Maximum duty cycles during search phase are close to 20%. Frequency alternation of search calls is interpreted as a mechanism for increasing duty cycle and thus the temporal continuity of scanning, as well as increasing the detection range. A neurophysiological correlate for the processing of search calls was found in the inferior colliculus. 64% of neurons respond to frequencies in the 30- to 40-kHz range and only in this frequency range were closed tuning curves found for levels below 40 dB SPL. In addition, 15% of the neurons have double-tuned frequency-threshold curves with best thresholds at 34 and 39 kHz. Differing from observations in other bats, approach calls of M. molossus are longer and of higher frequencies than search calls. Close to the roost, the call frequency is increased to 45.0–49.8 kHz and, in addition, extremely broadband signals are emitted. This demonstrates high plasticity of call design.Abbreviations BF best frequency - CF constant frequency - IC inferior colliculus - F_max maximal frequency - F_min minimal frequency - PF peak frequency - PSTH post-stimulus time histogram - QCF quasi-constant frequency - SPL sound pressure level     

Directional hearing of a cicada: biophysical aspects

The directional hearing of male and female cicadas of the species Tympanistalna gastrica was investigated by means of laser vibrometry. The results show that the tympanic organs act as pressure difference receivers. This mechanism can produce left-right differences of more than 10 dB. The main acoustic inputs to the inner surfaces of the ears are the tympana, in males supplemented by the timbals, and by the third spiracles in females. In addition the hollow abdomen of males seems to play a minor role. Tympanic membrane input is the source of left-right differences in the tympanic vibration velocity at frequencies below 9 kHz in males and below 15–18 kHz in females. The input via the (contralateral) timbal in males is responsible for a null in vibration velocity appearing between 12 and 14 kHz when the sound is coming from the contralateral direction. The highest energy components of the calling song are found in this frequency range. The mechanical sensitivity of the ears depends upon the sex. At low frequencies males are about 10 dB more sensitive than females.     

Echoortung bei der Fledermaus Chilonycteris rubiginosa

Zusammenfassung Chilonycteris rubiginosa erzeugt in allen Orientierungsituationen dreiteilige Ortungslaute. Im Anfangsteil steigt die Frequenz um etwa 1–2 kHz an. Der folgende Mittelteil hat eine konstante Frequenz von etwa 57 bis 57,6 kHz. Im Endteil fällt die Frequenz um etwa 8 kHz ab. Die Laute werden in Folgen von Lautgruppen ausgesendet.CR erzeugt pro Flügelschlag eine Lautgruppe. Im freien Flug zeigt CR Gruppen mit 2 Lauten von etwa 17–23 msec Dauer. Landende Fledermäuse senden in der Annäherungsphase Gruppen mit einer zunehmenden Zahl immer kürzerer Laute und in der Schlußphase eine längere Gruppe mit vielen kurzen Lauten.Fliegende Tiere senken die Frequenz des konstantfrequenten Mittelteils immer um etwa den Betrag der durch die Fluggeschwindigkeit bedingten Dopplereffekte ab, so daß die von den Tieren gehörte Echofrequenz nahezu konstant in Höhe der vor dem Flug ausgesendeten Frequenz gehalten wird.CR zeigt Kopf- und Ohrbewegungen. Die Ohrbewegungen stehen in Beziehung zur Lautaussendung.

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Echolocation by the bat Chilonycteris rubiginosa

Summary Chilonycteris rubiginosa (CR) produces tripartite ultrasonic sounds in all orientation situations. During the first part the frequency rises by 1–2 kHz. The following middle part has a constant frequency of about 57–57,6 kHz. In the terminal part the frequency decreases by about 8 kHz. The sounds are emitted as a sequence of groups of sounds.In flight they produce per wingbeat one group of sounds at a repetition rate of 10–11 Hz. In free flight CR emits groups of 2 sounds of about 17–23 msec duration. During the approach landing bats emit groups consisting of an increasing number of sounds of decreasing duration. During the terminal phase the group is longer in duration and consists of many short sounds.Flying CR lower the frequency of the middle part by an amount which compensates for Doppler shifts caused by the flight velocity. The frequency heard by the bats is, thus, always kept constant and equal to a frequency which is about 100–150 Hz above the medium frequency emitted before the flight. CR shows head and ear movements. The ear movements are correlated to the sound emission.

     

Acoustic irradiation produced by flying moths (Lepidoptera,Noctuidae)

Characteristics of acoustic waves accompanying the flight of noctuid moths (Noctuidae) were measured. The low-frequency part of the spectrum is formed of a series of up to 17 harmonics of the wingbeat frequency (30–50 Hz) with a general tendency toward the decrease in the spectral density and the increase in the sound frequency. The root-mean-square level of the sound pressure from flapping wings was found to be 70–78 dB SPL. Besides low-frequency components, the flight of moths was accompanied by short ultrasonic pulses, which appeared with every wingbeat. Most of the spectral energy was concentrated within a range of 7–150 kHz with the main peaks at 60–110 kHz. The short-term pulses were divided into two or more subpulses with different spectra. The high-frequency pulses were produced at two phases of the wingbeat cycle: during the pronation of the wings at the highest point and at the beginning of their upward movement from the lowest point. In most of the specimens tested, the peak amplitude of sounds varied from 55 to 65 dB SPL at a distance of 6 cm from the insect body. However, in nine noctuid species, no high-frequency acoustic components were recorded. In these experiments, the acoustic flow from the flying moth within a frequency range of 2 to 20 kHz did not exceed the self-noise level of the microphone amplifier (RMS 18 dB SPL). Probable mechanisms of the high frequency acoustic emission during flight, the effect of these sounds on the auditory sensitivity of moths, and the possibility of their self-revealing to insectivorous bats are discussed. In addition, spectral characteristics of the moth echolocation clicks were more precisely determined within the higher frequency range (>100 kHz).