Acoustic response properties of lagenar nerve fibers in the sleeper goby,<Emphasis Type="Italic"> Dormitator latifrons</Emphasis>

Auditory and vestibular functions of otolithic organs vary among vertebrate taxa. The saccule has been considered a major hearing organ in many fishes. However, little is known about the auditory role of the lagena in fishes. In this study we analyzed directional and frequency responses from single lagenar fibers of Dormitator latifrons to linear accelerations that simulate underwater acoustic particle motion. Characteristic frequencies of the lagenar fibers fell into two groups: 50 Hz and 80–125 Hz. We observed various temporal response patterns: strong phase-locking, double phase-locking, phase-locked bursting, and non-phase-locked bursting. Some bursting responses have not been previously observed in vertebrate otolithic nerve fibers. Lagenar fibers could respond to accelerations as small as 1.1 mm s^–2. Like saccular fibers, lagenar fibers were directionally responsive and decreased directional selectivity with stimulus level. Best response axes of the lagenar fibers clustered around the lagenar longitudinal axis in the horizontal plane, but distributed in a diversity of axes in the mid-sagittal plane, which generally reflect morphological polarizations of hair cells in the lagena. We conclude that the lagena of D. latifrons plays a role in sound localization in elevation, particularly at high stimulus intensities where responses of most saccular fibers are saturated.Abbreviations BRA best response axis/axes - BS best sensitivity - CF characteristic frequency - CV coefficient of variation - DI directionality index - ISIH inter-spike interval histogram - PSTH peri-stimulus time histogram - SR spontaneous rate     

Effects of saccular otolith removal on hearing sensitivity of the sleeper goby (<Emphasis Type="Italic">Dormitator latifrons</Emphasis>)

It is not known to what extent the entire saccule contributes to overall hearing sensitivity in any fish species. Here we report directional and frequency sensitivity in a teleost fish (Dormitator latifrons) and effects of unilateral and bilateral removal of saccular otoliths on its hearing sensitivity. The fish had different hearing thresholds in the horizontal (-54.4 to -50.3 dB re: 1 micro m) and mid-sagittal (-58.6 to -53.1 dB) planes. At 100 Hz, unilateral otolith removal did not significantly change hearing sensitivity in the mid-sagittal plane, but caused selective reductions of auditory sensitivity by 3-7 dB in the azimuthal axes that are consistent with the longitudinal axis of the damaged saccule. Along the fish's longitudinal axis, unilateral otolith removal significantly decreased auditory sensitivity at 50 Hz and 400 Hz, but not at 100 Hz, 200 Hz, and 345 Hz. At 100 Hz, bilateral otolith removal resulted in robust hearing loss of 27-35 dB at different axes in both horizontal and mid-sagittal planes. Along the fish's longitudinal axis, the bilateral removal reduced auditory sensitivity by 13-27 dB at the different frequencies. Therefore, these results demonstrate that the saccule plays important roles in directional hearing and frequency responses.     

Response characteristics of vibration-sensitive neurons in the midbrain of the grassfrog,Rana temporaria

Summary European grassfrogs (Rana temporaria) were stimulated with pulsed sinusoidal, vertical vibrations (10–300 Hz) and the responses of 46 single midbrain neurons were recorded in awake, immobilized animals.Most units (40) had simple V-shaped excitatory vibrational tuning curves. The distribution of best frequencies (BF's) was bimodal with peaks at 10 and 100 Hz and the thresholds ranged from 0.02 to 1.28cm/s² at the BF.Twenty-three neurons showed phasic-tonic and 11 neurons phasic responses. The dynamic range of seismic intensity for most neurons was 20–30 dB.In contrast to the sharp phase-locking in peripheral vibration-sensitive fibers, no phase-locking to the sinusoidal wave-form was seen in the midbrain neurons. The midbrain cells did not respond at low stimulus intensities (below 0.01–0.02 cm/s²) where a clear synchronization response occurs in saccular fibers.Six midbrain neurons had more complex response characteristics expressed by inhibition of their spontaneous activity by vibration or by bi-and trimodal sensory sensitivities.In conclusion, the vibration sensitive cells in the midbrain of the grassfrog can encode the frequency, intensity, onset and cessation of vibration stimuli. Seismic stimuli probably play a role in communication and detection of predators and the vibration-sensitive midbrain neurons may be involved in the central processing of such behaviorally significant stimuli.Abbreviation BF best frequency     

Auditory sensitivity of the cichlid fish Astronotus ocellatus (Cuvier)

Summary Auditory sensitivity was determined for the oscar, Astronotus ocellatus, a cichlid fish that has no known structural specializations to enhance hearing. Trained A. ocellatus behaviorally responded to sound stimuli from 200 Hz to 800 Hz with best sensitivity of 18 dB (re: 1 bar) to 21 dB for frequencies between 200 and 400 Hz. This is significantly poorer than hearing sensitivity for fish classified as hearing specialists, but well within the range of hearing capabilities reported for non-specialist teleost species.     

Development of the Acoustically Evoked Behavioral Response in Larval Plainfin Midshipman Fish,Porichthys notatus

The ontogeny of hearing in fishes has become a major interest among bioacoustics researchers studying fish behavior and sensory ecology. Most fish begin to detect acoustic stimuli during the larval stage which can be important for navigation, predator avoidance and settlement, however relatively little is known about the hearing capabilities of larval fishes. We characterized the acoustically evoked behavioral response (AEBR) in the plainfin midshipman fish, Porichthys notatus, and used this innate startle-like response to characterize this species'' auditory capability during larval development. Age and size of larval midshipman were highly correlated (r² = 0.92). The AEBR was first observed in larvae at 1.4 cm TL. At a size ≥1.8 cm TL, all larvae responded to a broadband stimulus of 154 dB re1 µPa or −15.2 dB re 1 g (z-axis). Lowest AEBR thresholds were 140–150 dB re 1 µPa or −33 to −23 dB re 1 g for frequencies below 225 Hz. Larval fish with size ranges of 1.9–2.4 cm TL had significantly lower best evoked frequencies than the other tested size groups. We also investigated the development of the lateral line organ and its function in mediating the AEBR. The lateral line organ is likely involved in mediating the AEBR but not necessary to evoke the startle-like response. The midshipman auditory and lateral line systems are functional during early development when the larvae are in the nest and the auditory system appears to have similar tuning characteristics throughout all life history stages.     

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     

The response characteristics of vibration-sensitive saccular fibers in the grassfrog,Rana temporaria

Summary The response characteristics of saccular nerve fibers in European grassfrogs (Rana temporaria) subjected to dorso-ventral, 10–200 Hz sinusoidal vibrations were studied.Only 4 fibers out of a total of 129 did not respond to the vibrations.70 fibers had an irregular spontaneous activity of 2–48 spikes/s. These fibers were very vibration-sensitive. The synchronization thresholds at 10–20 Hz varied from below 0.005 to 0.02 cm/s².In contrast to earlier results, all these fibers had low-pass characteristics (with respect to acceleration) and responded maximally at 10 and 20 Hz.55 fibers had spontaneous activities from 0–2 spikes/s. These fibers were less sensitive than the fibers with higher spontaneous activity. The spike-rate thresholds varied from about 0.04 to above 1.28 cm/s², giving a considerable range fractionation. Most of these fibers also had low-pass characteristics with respect to acceleration, but 8 fibers showed band-pass characteristics with maximal synchronizations and spike-rates occurring at 40–80 Hz.At high acceleration levels, most spikes fell within 5–10 degrees of the stimulus cycle. The phase-locking of the saccular fibers is therefore very acute at low frequencies.The phase angles preferred by the fibers at 10 Hz were bimodally distributed with the two peaks about 180° apart. This finding probably reflects the morphological observation that the saccular macula contains two oppositely oriented hair-cell populations. The results also indicate that the actual motion of the otolith relative to the macula is complex.No behavioral role of a vibration receptor has been demonstrated in the grassfrog. A use in predator avoidance is likely, and it is possible that the sacculus is used for detection of water surface-waves. The vibration sense could therefore be of importance in the detection and localization of conspecifics in the breeding ponds.     

The release of attack and escape behavior by vibratory stimuli in a wandering spider (Cupiennim salei keys.)

Summary The wandering spiderCupiennius salei responds to vibration of the substrate either with predatory behavior (approach) or with a startle reaction or escape behavior (withdrawal) (Fig. 3). The effects of different parameters of the signal in releasing this behavior were studied by applying various artificial stimuli to a spider standing on a vibrating platform with one or more legs. Receptors sensitive to substrate vibration and the trichobothria, which respond to airborne vibration, together determine the response. Spiders without trichobothria: The type of response to vertical vibrations isfrequency-dependent (Fig. 4a), with predatory reactions predominant at low frequencies (3–4 Hz), and withdrawal reactions at high frequencies (350–460 Hz). Whereas approach is most likely to occur at an intermediate, frequency-dependentamplitude, the probability of withdrawal increases continuously with increasing amplitude (Fig. 6). With sine wave stimuli the lowest threshold amplitude for approach is 9 m peak-to-peak (550 Hz, range tested 1–550 Hz) whereas that for withdrawal is 17 m (800 Hz, range tested 1–800 Hz). The threshold for approach is lower by 6–8 dB whenband-limited noise is used, and the probability of an approach response increases as the bandwidth is expanded. The threshold curve for withdrawal, however, is the same in all cases (Fig. 4b and 5). The spider is capable of both frequency and amplitude discrimination.The metatarsal and pretarsal slit sense organs contribute to these responses as is shown by increased thresholds following their destruction (Fig- 7). Intact animals, with functional trichobothria as well as slit sense organs: They have lower thresholds for withdrawal (by ca. 10 dB; Fig. 9) and shorter reaction times than do spiders without trichobothria. Unlike animals without trichobothria the amplitude thresholds of intact animals to bandlimited noise are ca. 7.5 dB lower than those to sine wave stimuli. The approach threshold is the same as that of spiders without trichobothria. According to direct observation the trichobothria are deflected by airborne sound generated by the substrate motion; the deflection angle increases with both amplitude and frequency of substrate vibration (Fig. 10).There is acentral nervous interaction between the signals from the trichobothria and the slit sense organs with the following basic properties: when both of the two receptor systems receive either a prey-like stimulus or a stimulus eliciting withdrawal their effects add, but when the trichobothria receive stimuli unlike prey they inhibit the approach reaction that would otherwise be triggered by substrate vibration.     

Vibrations in the orb web of the spider Nephila clavipes: cues for discrimination and orientation

Transmission of natural and artificial vibrations in webs of Nephila clavipes was examined using laser Doppler vibrometry to determine how this spider discriminates and localizes stimuli. 1. Vibration signals of four entrapped insect species peaked at different frequencies from 5–30 Hz, but their spectra overlapped considerably. Peak amplitudes spanned 50 dB. 2. Transmission of longitudinal vibrations along individual radii was attenuated over ca. 12 cm by 4.0 ± 2.7 dB; attenuation values for transverse and lateral vibrations were 22.2 ± 4.6 dB and 26.2 ± 4.3 dB, respectively. Some transmission spectra characteristics may be explained by resonances of the spider and threads. 3. Radial thread transmission increased by 2.2–5.8 dB after cutting the connecting auxiliary spirals, demonstrating that vibrations leak from stimulated radii via these threads. Auxiliary spirals provide structural support to Nephila webs at the expense of degraded directional transmission. 4. Upon single-point stimulation, vibrations measured around the web hub and at the spider's tarsi revealed 2-D vibration amplitude gradients of 20–30 dB indicating the stimulus direction. In contrast, measured vibration propagation velocities of 70–1500 m/s resulted in time-of-arrival differences at the spiders tarsi of < 1.5 ms, which may be too brief for stimulus direction determination.Abbreviations A area - C propagation velocity - E Young's modulus - LDV laser Doppler vibrometer/-metry - r.h. relative humidity - T tension - space constant - radius of gyration - density - 2-D two-dimensional - 3-D three-dimensional     

Directionality of auditory nerve fiber responses to pure tone stimuli in the grassfrog, Rana temporaria. II. Spike timing

We studied the directionality of spike timing in the responses of single auditory nerve fibers of the grass frog, Rana temporaria, to tone burst stimulation. Both the latency of the first spike after stimulus onset and the preferred firing phase during the stimulus were studied. In addition, the directionality of the phase of eardrum vibrations was measured. The response latency showed systematic and statistically significant changes with sound direction at both low and high frequencies. The latency changes were correlated with response strength (spike rate) changes and were probably the result of directional changes in effective stimulus intensity. Systematic changes in the preferred firing phase were seen in all fibers that showed phaselocking (i.e., at frequencies below 500–700 Hz). The mean phase lead for stimulation from the contralateral side was approximately 140° at 200 Hz and decreased to approximately 100° at 700 Hz. These phaseshifts correspond to differences in spike timing of approximately 2 ms and 0.4 ms respectively. The phaseshifts were nearly independent of stimulus intensity. The phase directionality of eardrum vibrations was smaller than that of the nerve fibers. Hence, the strong directional phaseshifts shown by the nerve fibers probably reflect the directional characteristics of extratympanic pathways. Accepted: 23 November 1996     

Electromotility of outer hair cells from the cochlea of the echolocating bat,Carollia perspicillata

Isolated outer hair cells (OHCs) and explants ot the organ of Corti were obtained from the cochlea of the echolocating bat, Carollia perspicillata, whose hearing range extends up to about 100 kHz. The OHCs were about 10–30 m long and produced resting potentials between-30 to -69 mV. During stimulation with a sinusoidal extracellular voltage field (voltage gradient of 2 mV/m) cyclic length changes were observed in isolated OHCs. The displacements were most prominent at the level of the cell nucleus and the cuticular plate. In the organ of Corti explants, the extracellular electric field induced a radial movement of the cuticular plate which was observed using video subtraction and photodiode techniques. Maximum displacements of about 0.3–0.8 m were elicited by stimulus frequencies below 100 Hz. The displacement amplitude decreased towards the noise level of about 10–30 nm for stimulus frequencies between 100–500 Hz, both in apical and basal explants. This compares well with data from the guinea pig, where OHC motility induced by extracellular electrical stimulation exhibits a low pass characteristic with a corner frequency below 1 kHz. The data indicate that fast OHC movements presumably are quite small at ultrasonic frequencies and it remains to be solved how they participate in amplifying and sharpening cochlear responses in vivo.Abbreviations BM basilar membrane - FFT fast Fourier Transfer - IHC inner hair cell - OHC outer hair cell     

Central auditory neurophysiology of a sound-producing fish: the mesencephalon of Pollimyrus isidori (Mormyridae)

This paper describes the auditory neurophysiology of the mesencephalon of P. isidori, a soundproducing mormyrid fish. Mormyrids have a specialized pressure-sensitive auditory periphery, and anatomical studies indicate that acoustic information is relayed to the mesencephalic nucleus MD. Fish were stimulated with tone bursts and clicks, and responses of MD neurons were recorded extracellularly. Auditory neurons had best frequencies (BF) and best sensitivities (BS) that fell within the range of frequencies and levels of the natural communication sounds of these fish. BSs were in the range of 0 to — 35 dB (re. 1.0 dyne/cm²). Many of the neurons were tuned (Q_{10 dB}: 2–6), and had BFs in the range of 100–300 Hz where the animal's sounds have their peak energy. A range of different physiological cell types were encountered, including phasic, sustained, and complex neurons. Some of the sustained neurons showed strong phase-locking to tones. Many neurons exhibited non-monotonic rate-level functions. Frequencies flanking the BF often caused a reduction in spontaneous activity suggesting inhibition. Many neurons showed excellent representation of click-trains, and some showed a temporal representation of inter-click-intervals with errors less than 1 ms.Abbreviations BF best frequency - BS best sensitivity - ELa anterior exterolateral toral nucleus - ELp posterior exterolateral toral nucleus - EOCD electric organ command discharge - FFT fast Fourier transform - HRP horseradish peroxidase - ICI inter-clickinterval - MD mediodorsal toral nucleus (=auditory nucleus) - OR onset response rate - PSTH peri-stimulus-time-histogram - R synchronization coefficient - RA response area - SS steady state response rate     

Frequency response characteristics of electroreceptors in weakly electric fish (Gymnotoidei) with a pulse discharge

Summary Three species of Gymnotid fish, two species ofHypopomus andRhamphichthys rostratus, each having pulse type electric organ discharges (EOD) of different durations were studied to learn if any correlation exists between the spectral composition of the species specific EOD pulse and the frequency response characteristics of that species' electroreceptors. The receptor population consisted of two major categories (examples in Fig. 3). One category, termed pulse marker receptors, responded to suprathreshold stimulus pulses with a single spike at a short (<2 ms) latency. These receptors were tuned to the higher frequency components of a species' EOD (Fig. 4A) and were always 5 to 10 dB less sensitive than any other electroreceptors within a given species. The second major receptor category, burst duration coders, responded to an electrical stimulus with a burst of spikes at a longer latency, burst length was a function of stimulus amplitude. This second category could be further divided into three sub-categories according to the receptors' frequency response characteristics. The most commonly seen subcategory, wide band receptors (Fig. 4B), responded best to stimuli having frequencies equal to the dominant frequency component of the species' EOD in the two species ofHypopomus studied. A second subcategory, narrow band receptors (Fig. 4 A), had frequency response characteristics similar to those of the pulse marker receptors; however, these had thresholds 10 dB lower than those of the pulse marker. The third subcategory of burst duration coders, low frequency receptors (Fig. 4 C, D), responded best to stimulus frequencies ranging from about 50 to 150 Hz. Mechanisms of coding stimulus amplitude and responses to prolonged sinusoidal electrical stimuli were also studied in the various receptor types.It is suggested that the differences in the major receptor types and the different frequency response characteristics of the electroreceptors within a given species allows the animals to identify and evaluate signals resulting from their own EOD, the EODs of conspecifics and electrical stimuli generated by other species of electric fish.Supported by NIH Grant #1 RO1 NS 12337-01     

Modulation rate transfer functions in bottlenose dolphins (<Emphasis Type="Italic">Tursiops truncatus</Emphasis>) with normal hearing and high-frequency hearing loss

Envelope following responses were measured in two bottlenose dolphins in response to sinusoidal amplitude modulated tones with carrier frequencies from 20 to 60 kHz and modulation rates from 100 to 5,000 Hz. One subject had elevated hearing thresholds at higher frequencies, with threshold differences between subjects varying from ±4 dB at 20 and 30 kHz to +40 dB at 50 and 60 kHz. At each carrier frequency, evoked response amplitudes and phase angles were plotted with respect to modulation frequency to construct modulation rate transfer functions. Results showed that both subjects could follow the stimulus envelope components up to at least 2,000 Hz, regardless of carrier frequency. There were no substantial differences in modulation rate transfer functions for the two subjects suggesting that reductions in hearing sensitivity did not result in reduced temporal processing ability. In contrast to earlier studies, phase data showed group delays of approximately 3.5 ms across the tested frequency range, implying generation site(s) within the brainstem rather than the periphery at modulation rates from 100 to 1,600 Hz. This discrepancy is believed to be the result of undersampling of the modulation rate during previous phase measurements.     

Tympanic and extratympanic sound transmission in the leopard frog

Summary The inner ear of the leopard frog,Rana pipiens, receives sound via two separate pathways: the tympanic-columellar pathway and an extratympanic route. The relative efficiency of the two pathways was investigated. Laser interferometry measurements of tympanic vibration induced by free-field acoustic stimulation reveal a broadly tuned response with maximal vibration at 800 and 1500 Hz. Vibrational amplitude falls off rapidly above and below these frequencies so that above 2 kHz and below 300 Hz tympanic vibration is severely reduced. Electrophysiological measurements of the thresholds of single eighth cranial nerve fibers from both the amphibian and basilar papillae in response to pure tones were made in such a way that the relative efficiency of tympanic and extratympanic transmission could be assessed for each fiber. Thresholds for the two routes are very similar up to 1.0 kHz, above which tympanic transmission eventually becomes more efficient by 15–20 dB. By varying the relative phase of the two modes of stimulation, a reduction of the eighth nerve response can be achieved. When considered together, the measurements of tympanic vibration and the measurements of tympanic and extratympanic transmission thresholds suggest that under normal conditions in this species (1) below 300 Hz extratympanic sound transmission is the main source of inner ear stimulation; (2) for most of the basilar papilla frequency range (i.e., above 1.2 kHz) tympanic transmission is more important; and (3) both routes contribute to the stimulation of amphibian papilla fibers tuned between those points. Thus acoustic excitation of the an uran's inner ear depends on a complex interac tion between tympanic and extratympanic sound transmission.Abbreviations dB SPL decibels sound pressure level re: 20 N/ m² - AP amphibian papilla - BP basilar papilla - BEF best excitatory frequency     

Der „Kniesehnenreflex” bei Carausius morosus: Übergangsfunktion und Frequenzgang

Stretching and releasing the femoral chordotonal organ caused by a movement of the tendon of the organ gives rise to a movement of the tibia. This reaction is called Kniesehnenreflex (knee-tendon-reflex). Its step response can be described in the following manner: After a certain reaction-time (at flexion 0.02–0.06 sec, at extension 0.06–0.2 sec) the tibia moves with a maximum speed between 150°/sec and 1000°/sec at extension and between 20°/sec and 450°/sec at flexion. The amplitude of the movement and the maximum speed of tibia movement are correlated. After reaching the extreme position the tibia returnes nearly to its starting-point with half lifes of 3–58 sec after a flexion and 7–232 sec after an extension. — The frequency response shows a strong decrease of the amplitude of the tibia at about 1 Hz. Above 2 Hz the amplitude is only a few degrees. The phase shift between stimulus and reaction increases with increasing frequency. Big individual differences are observed. A step stimulus, which is given in addition to a sinoidal stimulus causes a response at all frequencies. — Slow stretching and releasing the chordotonal organ with constant speeds causes movements of the tibia even at stimulus speeds of 0.002 mm/min. — It is discussed: the significance of the results for the theory of the control mechanism at walk, the stability of the control system in connection with the rocking-movements of the animal and the control of Flexibilitas cerea.     

Directionality of auditory nerve fiber responses to pure tone stimuli in the grassfrog, Rana temporaria. I. Spike rate responses

We studied the directionality of spike rate responses of auditory nerve fibers of the grassfrog, Rana temporaria, to pure tone stimuli. All auditory fibers showed spike rate directionality. The strongest directionality was seen at low frequencies (200 – 400 Hz), where the spike rate could change by up to nearly 200␣spikes s⁻¹. with sound direction. At higher frequencies the directional spike rate changes were mostly below 100 spikes s⁻¹. In equivalent dB SPL terms (calculated using the fibers' rate-intensity curves) the maximum directionalities were up to 15 dB at low frequencies and below 10 dB at higher frequencies. Two types of directional patterns were observed. At frequencies below 500 Hz relatively strong responses were evoked by stimuli from the ipsilateral (+90^o) and contralateral (−90^o) directions while the weakest responses were evoked by stimuli from frontal (0^o or +30^o) or posterior (−135^o) directions. At frequencies above 800 Hz the strongest responses were evoked by stimuli from the ipsilateral direction while gradually weaker responses were seen as the sound direction shifted towards the contralateral side. At frequencies between 500 and 800 Hz both directional patterns were seen. The directionality was highly intensity dependent. No special adaptations for localization of conspecific calls were found. Accepted: 23 November 1996     

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