Behavioral detection of acoustic particle motion by a teleost fish (Astronotus ocellatus): sensitivity and directionality

The physiology and morphology of auditory interneurons of a fly, the parasitoid Therobia leonidei, are described for the first time. 1. The hearing threshold has been determined with summed recordings of the neck connective. Females are most sensitive in a frequency range from 16 to 40 kHz (thresholds: around 45 dB SPL). This broad hearing range matches with the peak frequencies of the song spectra of host bushcricket species. Male flies are 10–20 dB less sensitive than females. 2. The sensory cells of the prosternal tympanal organ of T. leonidei project into the thoracico-abdominal ganglion complex with arborizations in all three thoracic neuromeres. 3. Three types of ascending auditory interneurons were identified by their morphology and response properties. These have arborizations in all three thoracic neuromeres and terminate soma-contralaterally in the brain. At least three other neuron types were also identified according to response properties alone. The neurons show similar spectral tuning but different sensitivities.     

Correspondence between evoked vocal responses and auditory thresholds in Pleurodema thaul (Amphibia; Leptodactylidae)

Thresholds for evoked vocal responses and thresholds of multiunit midbrain auditory responses to pure tones and synthetic calls were investigated in males of Pleurodema thaul, as behavioral thresholds well above auditory sensitivity have been reported for other anurans. Thresholds for evoked vocal responses to synthetic advertisement calls played back at increasing intensity averaged 43 dB RMS SPL (range 31–52 dB RMS SPL), measured at the subjects’ position. Number of pulses increased with stimulus intensities, reaching a plateau at about 18–39 dB above threshold and decreased at higher intensities. Latency to call followed inverse trends relative to number of pulses. Neural audiograms yielded an average best threshold in the high frequency range of 46.6 dB RMS SPL (range 41–51 dB RMS SPL) and a center frequency of 1.9 kHz (range 1.7–2.6 kHz). Auditory thresholds for a synthetic call having a carrier frequency of 2.1 kHz averaged 44 dB RMS SPL (range 39–47 dB RMS SPL). The similarity between thresholds for advertisement calling and auditory thresholds for the advertisement call indicates that male P. thaul use the full extent of their auditory sensitivity in acoustic interactions, likely an evolutionary adaptation allowing chorusing activity in low-density aggregations.     

Otoacoustic emissions from insect ears having just one auditory neuron

Sensitive hearing organs often employ nonlinear mechanical sound processing which produces distortion-product otoacoustic emissions. Such emissions are also recorded from insect tympanal organs. Here we report high frequency distortion-product emissions, evoked by stimulus frequencies up to 95 kHz, from the tympanal organ of a notodontid moth, Ptilodon cucullina, which contains only a single auditory receptor neuron. The 2f1–f2 distortion-product emission reaches sound levels above 40 dB SPL. Most emission growth functions show a prominent notch of 20 dB depth (n = 20 trials), accompanied by an average phase shift of 119°, at stimulus levels between 60 and 70 dB SPL, which separates a low- and a high-level component. The emissions are vulnerable to topical application of ethyl ether which shifts growth functions by about 20 dB towards higher stimulus levels. For the mammalian cochlea, Lukashkin and colleagues have proposed that distinct level-dependent components of nonlinear amplification do not necessarily require interaction of several cellular sources but could be due to a single nonlinear source. In notodontids, such a physiologically vulnerable source could be the single receptor cell. Potential contributions from accessory cells to the nonlinear properties of the scolopidial hearing organ are still unclear.     

Psychophysical and neurophysiological hearing thresholds in the bat <Emphasis Type="Italic">Phyllostomus </Emphasis><Emphasis Type="Italic">discolor</Emphasis>

Absolute hearing thresholds in the spear-nosed bat Phyllostomus discolor have been determined both with psychophysical and neurophysiological methods. Neurophysiological data have been obtained from two different structures of the ascending auditory pathway, the inferior colliculus and the auditory cortex. Minimum auditory thresholds of neurons are very similar in both structures. Lowest absolute thresholds of 0 dB SPL are reached at frequencies from about 35 to 55 kHz in both cases. Overall behavioural sensitivity is roughly 20 dB better than neural sensitivity. The behavioural audiogram shows a first threshold dip around 23 kHz but threshold was lowest at 80 kHz (−10 dB SPL). This high sensitivity at 80 kHz is not reflected in the neural data. The data suggest that P. discolor has considerably better absolute auditory thresholds than estimated previously. The psychophysical and neurophysiological data are compared to other phyllostomid bats and differences are discussed. S. Hoffmann, L. Baier, F. Borina contributed equally to this work.     

Serially homologous ears perform frequency range fractionation in the praying mantis, Creobroter (Mantodea, Hymenopodidae)

Unlike most praying mantises that have a single region of auditory sensitivity, species in the genus Creobroter have equally sensitive hearing at 2–4 and at 25–50 kHz and and are relatively insensitivity at 10–15 kHz — they have a W-shaped audiogram. Ultrasonic sensitivity originates from an auditory organ in the ventral midline of the metathorax that closely resembles the ear of other mantises. Ablation experiments demonstrate that low frequency sensitivity derives from a serially homologous mesothoracic auditory organ. Extracellular recordings suggest that these two ears operate largely, if not entirely, independently of one another in the thorax. The low frequency response has a longer latency, more action potentials per stimulus, and different patterns of change with increasing SPL than the high frequency response. Separate interneurons mediate responses in the two frequency ranges, but our evidence suggests that they are two serially homologous sets of cells. Neither auditory organ shows any physiological evidence of directional sensitivity. Ultrasound triggers a set of behaviors in flying hymenopodid mantises much like those in other mantises, but the behavioral significance of low frequency hearing in these animals is still unknown.Abbreviations SPL sound pressure level - dB SPL sound pressure level re: 20 Pa - HF high frequency - LF low frequency     

Neuroethology of ultrasonic hearing in nocturnal butterflies (Hedyloidea)

Nocturnal Hedyloidea butterflies possess ultrasound-sensitive ears that mediate evasive flight maneuvers. Tympanal ear morphology, auditory physiology and behavioural responses to ultrasound are described for Macrosoma heliconiaria, and evidence for hearing is described for eight other hedylid species. The ear is formed by modifications of the cubital and subcostal veins at the forewing base, where the thin (1–3 μm), ovoid (520 × 220 μm) tympanal membrane occurs in a cavity. The ear is innervated by nerve IIN1c, with three chordotonal organs attaching to separate regions of the tympanal membrane. Extracellular recordings from IIN1c reveal sensory responses to ultrasonic (>20 kHz), but not low frequency (<10 kHz) sounds. Hearing is broadly tuned to frequencies between 40 and 80 kHz, with best thresholds around 60 dB SPL. Free flying butterflies exposed to ultrasound exhibit a variety of evasive maneuvers, characterized by sudden and unpredictable changes in direction, increased velocity, and durations of ∼500 ms. Hedylid hearing is compared to that of several other insects that have independently evolved ears for the same purpose-bat detection. Hedylid hearing may also represent an interesting example of evolutionary divergence, since we demonstrate that the ears are homologous to low frequency ears in some diurnal Nymphalidae butterflies.     

Responses and song pattern copying of Omega-type I-neurons in the cricket,Gryllus bimaculatus,at different prothoracic temperatures

Summary Omega-type I-neurons (ON/1) (Fig. 1A) were recorded intracellularly with the prothoracic ganglion kept at temperatures of either 8–9°, or 20–22° or 30–33 °C and the forelegs with the tympanal organs kept at ambient temperature (20–22 °C). The neurons were stimulated with synthetic calling songs (5 kHz carrier frequency) with syllable periods (SP in ms) varying between 20 and 100, presented at sound intensities between 40 and 80 dB SPL. The amplitude and duration of spikes as well as response latency decreased at higher temperatures (Figs. 1 B, 2, 6). At lower prothoracic temperatures (8–9 °C) the neuron's responses to songs with short SP (20 ms) failed to copy single syllables, or with moderate SP (40 ms) copied the syllable with low signal to noise ratio (Fig. 3). The auditory threshold of the ON/1 type neuron, when tested with the song model, was temperature-dependent. At 9° and 20 °C it was between 40 and 50 dB SPL and at 33 °C it was less than 40 dB SPL (Fig. 4). For each SP, the slope of the intensity-response function was positively correlated with temperature, however, at low prothoracic temperatures the slope was lower for songs with shorter SPs (Fig. 5). The poor copying of the syllabic structure of the songs with short SPs at low prothoracic temperatures finds a behavioral correlate because females when tested for phonotaxis on a walking compensator responded best to songs with longer SPs at a similar temperature.Abbreviations epsps excitatory postsynaptic potentials - ON/1 omega-type I-neuron - SP syllable period - SPL sound pressure level     

The tympanal hearing organ of the parasitoid fly Ormia ochracea (Diptera, Tachinidae, Ormiini)

Tympanate hearing has evolved in at least 6 different orders of insects, but had not been reported until recently in the Diptera. This study presents a newly discovered tympanal hearing organ, in the parasitoid tachinid fly, Ormia ochracea. The hearing organ is described in terms of external and internal morphology, cellular organization of the sensory organ and preliminary neuroanatomy of the primary auditory afferents. The ear is located on the frontal face of the prothorax, directly behind the head capsule. Conspicuously visible are a pair of thin cuticular membranes specialized for audition, the prosternal tympanal membranes. Directly attached to these membranes, within the enlarged prosternal chamber, are a pair of auditory sensory organs, the bulbae acusticae. These sensory organs are unique among all auditory organs known so far because both are contained within an unpartitioned acoustic chamber. The prosternal chamber is connected to the outside by a pair of tracheae. The cellular anatomy of the fly's scolopophorous organ was investigated by light and electron microscopy. The bulba acustica is a typical chordotonal organ and it contains approximately 70 receptor cells. It is similar to other insect sensory organs associated with tympanal ears. The similarity of the cellular organization and tympanal morphology of the ormiine ear to the ears of other tympanate insects suggests that there are potent constraints in the design features of tympanal hearing organs, which must function to detect high frequency auditory signals over long distances. Each sensory organ is innervated by a branch of the frontal nerve of the fused thoracic ganglia. The primary auditory afferents project to each of the pro-, meso-, and metathoracic neuropils. The fly's hearing organ is sexually dimorphic, whereby the tympanal membranes are larger in females and the spiracles larger in males. The dimorphism presumably reflects differences in the acoustic behavior in the two sexes.     

Involvement of the mechanosensory complex structures of the cricket <Emphasis Type="Italic">Phaeophilacris bredoides</Emphasis> in triggering of motor responses to sound

Using an ethological approach, we studied the possibility of sound perception as well as probable contribution of diverse mechanosensory systems composing the mechanosensory complex to triggering of motor responses to sound stimulation in imaginal crickets Phaeophilacris bredoides lacking the tympanal organs (“deaf”). It was shown that Ph. bredoides imagoes are able to perceive sounds and respond to sound cues by a locomotor reaction in a relatively broad frequency range which becomes narrower as sound intensity decreases [0.1–6.0 kHz (111 ± 3 dB SPL), 0.1–1.5 kHz (101 ± 3 dB SPL), 0.1–1.3 kHz (91 ± 3 dB SPL), 0.1–0.6 kHz (81 ± 3 dB SPL), and 0.1 kHz (71 ± 3 dB SPL)]. Sound perception and triggering ofmotor responses appear to involve the cercal organs (CO), subgenual organs (SO) and, probably, other distant mechanosensory organs (DMO). CO are essential for triggering of locomotor responses to sound within the ranges of 1.6–6.0 kHz (111 ± 3 dB SPL), 1–1.5 kHz (101 ± 3 dB SPL), 0.9–1.3 kHz (91 ± 3 dB SPL), and 0.5–0.6 kHz (81 ± 3 dB SPL). SO and, probably, other DMO provide locomotor responses to sound within the ranges of 0.1–6.0 kHz (111 ± 3 dB SPL), 0.1–0.8 kHz (101 ± 3 dB SPL), 0.1–0.4 kHz (91 ± 3 dB SPL), and 0.1–0.4 kHz (81 ± 3 dB SPL). From this, it follows that “deaf” (nonsinging) Ph. bredoides can perceive sounds using CO, SO and, probably, other DMO, which (as in singing crickets) are likely to compose an integrated mechanosensory complex providing adequate acoustic behavior of this cricket species. Performance efficiency and sensitivity of the mechanosensory complex (specifically, of CO) rely on the thoroughness of grooming. Following self-cleaning of CO, the level of cricket motor activity in response to cue presentation returned to the baseline and sometimes even increased. Whether or not crickets of this species communicate acoustically is yet to be found out, however, we suggest that the mechanosensory complex, which triggers motor responses to a sound, is normally involved in the defensive escape response aimed at rescuing from predators.     

Auditory response characteristics of the piebald odorous frog and their implications

The piebald odorous frog (Odorrana schmackeri), the large odorous frog (Odorrana livida) and the concave-eared torrent frog (Amolops tormotus) are sympatric species living near the same torrent streams in the vicinity of Mt. Huangshan, China. A recent study demonstrated that A. tormotus can use sound signals involving ultrasonic components for communication in a noisy environment, and another sympatric species, O. livida, can also perceive ultrasonic sound. Here we report data on the hearing range of O. schmackeri by studying auditory evoked potentials and single-unit data from the torus semicircularis. This frog exhibits its two most sensitive peaks at 2 kHz and 3.5–4.0 kHz with thresholds <42 dB SPL, with an upper frequency limit of hearing at 8.5 kHz with threshold of 87 dB SPL. The upper limit is much lower than those of O. livida and A. tormotus, at 22 and 34 kHz, respectively. It suggests that sympatric species may respond differently to similar environmental selection pressures sculpting auditory communication systems.     

Noise improves transfer of near-threshold, phase-locked activity of the cochlear nerve: evidence for stochastic resonance?

 Stochastic resonance can be described as improved detection of weak periodic stimuli by a dynamic nonlinear system, resulting from the simultaneous presentation of a restricted dynamic range of low-intensity noise. This property has been reported in simple physical and biological activities. The present study describes data consistent with the interpretation that stochastic resonance can be observed in the response of cochlear neurons. These experiments utilized low levels (−5 to 25 dB SPL) of stimuli and noise (5 to 30 dB SPL). Stimuli consisted of simultaneously presented 8 kHz (F ₁) and 8.8 kHz (F ₂) tone bursts, which generated an 800 Hz F ₂–F ₁ cochlear nerve envelope ensemble response in the gerbil. The mean response threshold was approximately −3 dB SPL. Simultaneous presentation of a low-intensity wideband noise increased the amplitude of this response. This was observed with tonal stimuli having intensities of 0–5 dB SPL; responses to stimulus levels >10 dB were attenuated by noise. Response amplitude was increased by noise levels of 10–15 dB; the amplitude was unaffected by lower levels of noise, and decreased in the presence of higher noise levels. These properties are compatible with those of stochastic resonance. Accepted: 11 March 1999     

Comparison of auditory sense organs in parasitoid Tachinidae (Diptera) hosted by Tettigoniidae (Orthoptera) and homologous structures in a non-hearing Phoridae (Diptera)

The dipteran parasitoids Therobia leonidei and Homotrixa alleni (Tachinidae) use acoustic cues to locate their calling tettigoniid (Ensifera, Orthoptera) hosts. The sexually dimorphic tympanal organs of both fly species are located at the prosternum. For comparison a homologous chordotonal organ in the non-hearing fly Phormia regina, Meigen (Phoridae) is also described. The scolopidial sense organs of the ears have approximately 180 sensory cells in Th. leonidei and 250 cells in H. alleni. Interspecific analysis indicates that the cell number and arrangement might be genus specific in Tachinidae. The mononematic scolopidia, each with one sensory cell, are of different sizes and insert at the tympanal membrane. Large scolopidial units (diameter of sensory cells up to 50 μm) extend longitudinally from the centre of the sensory organ towards the ligament, whereas small units (sensory cell diameter up to 10 μm) are arranged sequentially within the sensory organ. This arrangement is discussed to be a possible basis for frequency discrimination. The ultrastructure of the scolopidia is similar in the hearing and non-hearing flies. In both groups, the majority of scolopales has a diameter from 2 to 2.9 μm, although hearing species have additionally wider scolopales. The homologous chordotonal organ of Ph. regina consists of approximately 55 sensory cells of uniform direction. The data are discussed in comparison to the ears of other Diptera.     

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.     

Presynaptic contributions to response shape in an auditory neuron of the grasshopper

Neuron 714 is morphologically one of the most prominent neurons in the central auditory pathway of the grasshopper with arborizations extending from the abdominal neuromeres of the metathoracic ganglion to the brain. The aim of this study is to explore auditory information flow involving neuron 714 at the level of the ventral nerve cord. Paired intracellular recordings were made from neuron 714 in the mesothorax on the one hand, and from candidate presynaptic auditory neurons of the metathorax on the other. Electrical stimulation of the tympanal nerves provides an estimate of the synaptic distance between these interneurons and auditory afferents. Four, including neuron 714, are monosynaptically connected to afferents, the remainder disynaptically. Current-injection and spike-triggered averaging reveal that of nine neurons examined, seven make either monosynaptic, disynaptic or polysynaptic connections onto neuron 714. All connections are excitatory. Paired recordings show that response duration and response amplitude in synaptically linked cells vary according to the frequency of the stimulus. Measurements of the latency of the first excitatory post-synaptic potential evoked in neuron 714 by afferents and by metathoracic interneurons show how the synaptic drive from these sources shapes the auditory response of neuron 714. Accepted: 14 October 1998     

Involvement of Mechanosensory Complex Structures of the Cricket <Emphasis Type="Italic">Gryllus bimaculatus</Emphasis> Larvae (Orthoptera,Gryllidae) in Triggering of Motor Responses to Sound

Using an ethological approach, we studied the possibility of sound perception as well as probable contribution of diverse mechanosensory systems composing the mechanosensory complex to triggering of motor responses to sound stimulation in the cricket Gryllus bimaculatus larvae. It was shown that larvae can perceive sounds and respond to them by a locomotor reaction in a relatively broad frequency range, which becomes narrower as sound intensity decreases [0.1–6.6 kHz (111 ± 3 dB SPL), 0.1–1.4 kHz (101 ± 3 dB SPL), 0.1–0.8 kHz (91 ± 3 dB SPL]. Sound perception and triggering of motor responses appear to involve the cercal organs (CO), subgenual organs (SO) and, probably, other distant mechanosensory organs (DMO). Normal functioning of CO is essential for triggering locomotor responses to sound within the ranges of 1–1.4 kHz (101 ± 3 dB SPL) and 0.1–0.8 kHz (91 ± 3 dB SPL). CO are not necessary for triggering of motor responses to cues with an intensity of 111 ± 3 dB. SO and, probably, other DMO provide locomotor responses to sound within the ranges of 0.1–6.6 kHz (111 ± 3 dB SPL), 0.1–0.9 kHz (101 ± 3 dB SPL), and 0.1–0.3 kHz (91 ± 3 dB SPL). Thus, last instar larvae of G. bimaculatus lacking the tympanal organs can perceive sounds using CO, SO and, probably, other DMO, which (as in cricket imagoes) are likely to compose an integrated mechanosensory complex providing adequate acoustic behavior of this cricket species. Performance efficiency and sensitivity of the mechanosensory complex (specifically, CO) rely on the thoroughness of grooming. After self-cleaning of CO, the level of larval motor activity in response to cue presentation returned to the baseline and sometimes even increased. We assume that under normal conditions the mechanosensory complex, which triggers motor responses to a sound, is involved in the defensive escape response aimed at rescuing from predators.     

Directional characteristics of the auditory system of cicadas: is the sound producing tymbal an integral part of directional hearing?

Abstract. Directional hearing is investigated in males of two species of cicadas, Tympanistalna gastrica (Stål) and Tettigetta josei Boulard, that are similar in size but show different calling song spectra. The vibrational response of the ears is measured with laser vibrometry and compared with thresholds determined from auditory nerve recordings. The data are used to investigate to what extent the directional characteristic of the tympanal vibrations is encoded by the activity of auditory receptors. Laser measurements show complex vibrations of the tympanum, and reveal that directional differences are rather high (>15 dB) in characteristic but limited frequency ranges. At low frequencies, both species show a large directional difference at the same frequency (3–5 kHz) whereas, above 10 kHz, the directional differences correspond to the different resonant frequencies of the respective tymbals. Consequently, due to the mechanical resonance of the tymbal, the frequency range at which directional differences are high differs between the two species that otherwise show similar dimensions of the acoustic system. The directional differences observed in the tympanal vibrations are also observed in the auditory nerve activity. These recordings confirm that the biophysically determined directional differences are available within the nervous system for further processing. Despite considerable intra as well as interindividual variability, the ears of the cicadas investigated here exhibit profound directional characteristics, because the thresholds determined from recordings of the auditory nerve at 30° to the right and left of the longitudinal axis differ by more than 5 dB.     

Morphological and physiological regeneration in the auditory system of adult <Emphasis Type="Italic">Mecopoda elongata</Emphasis> (Orthoptera: Tettigoniidae)

Orthopterans are suitable model organisms for investigations of regeneration mechanisms in the auditory system. Regeneration has been described in the auditory systems of locusts (Caelifera) and of crickets (Ensifera). In this study, we comparatively investigate the neural regeneration in the auditory system in the bush cricket Mecopoda elongata. A crushing of the tympanal nerve in the foreleg of M. elongata results in a loss of auditory information transfer. Physiological recordings of the tympanal nerve suggest outgrowing fibers 5 days after crushing. An anatomical regeneration of the fibers within the central nervous system starts 10 days after crushing. The neuronal projection reaches the target area at day 20. Threshold values to low frequency airborne sound remain high after crushing, indicating a lower regeneration capability of this group of fibers. However, within the central target area the low frequency areas are also innervated. Recordings of auditory interneurons show that the regenerating fibers form new functional connections starting at day 20 after crushing.     

Effect of temperature upon auditory thresholds in two anuran species,Bombina v. variegata andAlytes o. obstetricans (Amphibia,Discoglossidae)

Summary Because it seemed likely that temperature affects not only the calling mechanism of anurans, but their auditory systems as well, we have measured the thresholds ofBombina variegata variegata andAlytes obstetricans obstetricans at 5°, 12°, 20° and 28°C by recording multiple-unit activity from the torus semicircularis. An increase in temperature from 5° to 28°C shortened the latencies considerably. InBombina v. variegata latencies fell from an average of 32 ms to 13 ms (600 Hz), and inAlytes o. obstetricans from an average of 22 ms to 11 ms (500 Hz). At frequencies below 500 Hz the decrease was still greater. Latency was also dependent on frequency, being shorter with high-frequency tones. At 5°C the auditory neurons ofBombina are rather insensitive and respond irregularly. At 12°C and at 20°C sensitivity is markedly increased. The minimum threshold in males was at 400–500 Hz (49 dB SPL), and that of females was at 450 Hz (47 dB SPL). There was no further increase in sensitivity at 28°C. InAlytes the auditory neurons were fully functional even at 5°C. At this temperature the audiogram had sensitivity maxima at 300, 1,100–1,300 and 1,800 Hz. In both males and females an increase in temperature to 20°C caused an extraordinary increase in sensitivity, primarily in the low-frequency range; the minimum threshold, at 400 Hz, was 44 dB SPL in males and 41 dB SPL in famales. In the intermediate frequency range there was also a marked increase in sensitivity, but not in the high-frequency range, where the best frequency was 1,800 Hz. At 28°C the threshold to low-frequency tones was increased.     

The generation of DPOAEs in the locust ear is contingent upon the sensory neurons

Tympanal organs of insects emit distortion-product otoacoustic emissions (DPOAEs) that are indicative of nonlinear ear mechanics. Our study sought (1) to define constraints of DPOAE generation in the ear of Locusta migratoria, and (2) to identify the sensory structures involved. We selectively destroyed the connection between the (peripheral) sensory ganglion and the tympanal attachment points of the “d-cell” dendrites; d-cells are most sensitive to sound frequencies above 12 kHz. This led to a decrease of DPOAEs that were evoked by f ₂ frequencies above 15 kHz (decrease of 15–40 dB; mean 28 dB; n = 12 organs). DPOAEs elicited by lower frequencies remained unchanged. Such frequency-specific changes following the exclusion of one scolopidial sub-population suggest that these auditory scolopidia are in fact the source of DPOAEs in insects. Electrical stimulation of the auditory nerve (with short current pulses of 4–10 μA or DC-currents of 0.5 μA) reversibly reduced DPOAEs by as much as 30 dB. We assume that retrograde electrical stimulation primarily affected the neuronal part of the scolopidia. Severing the auditory nerve from the central nervous system (CNS) did not alter the DPOAE amplitudes nor the effects of electrical stimulation.     

Characterization of auditory afferents in the tiger beetle,Cicindela marutha Dow

We have identified a nerve carrying auditory afferents and characterized their physiological responses in the tiger beetle,Cicindela marutha.