Passive sound localization of prey by the pallid bat (Antrozous p. pallidus)

Summary The pallid bat (Antrozous p. pallidus) uses passive sound localization to capture terrestrial prey. This study of captive pallid bats examined the roles of echolocation and passive sound localization in prey capture, and focused on their spectral requirements for accurate passive sound localization.Crickets were used as prey throughout these studies. All tests were conducted in dim, red light in an effort to preclude the use of vision. Hunting performance did not differ significantly in red light and total darkness, nor did it differ when visual contrast between the terrestrial prey and the substrate was varied, demonstrating that the bats did not use vision to locate prey.Our bats apparently used echolocation for general orientation, but not to locate prey. They did not increase their pulse emission rate prior to prey capture, suggesting that they were not actively scanning prey. Instead, they required prey-generated sounds for localization. The bats attended to the sound of walking crickets for localization, and also attacked small, inanimate objects dragged across the floor. Stationary and/or anesthetized crickets were ignored, as were crickets walking on substrates that greatly attenuated walking sounds. Cricket communication sounds were not used in prey localization; the bats never captured stationary, calling crickets.The accuracy of their passive sound localization was tested with an open-loop passive sound localization task that required them to land upon an anesthetized cricket tossed on the floor. The impact of a cricket produced a single 10–20 ms duration sound, yet with this information, the bats were able to land within 7.6 cm of the cricket from a maximum distance of 4.9 m. This performance suggests a sound localization accuracy of approximately ±1° in the horizontal and vertical dimensions of auditory space. The lower frequency limit for accurate sound localization was between 3–8 kHz. A physiological survey of frequency representation in the pallid bat inferior colliculus suggests that this lower frequency limit is around 5 kHz.     

Azimuthal sound localization in the European starling (Sturnus vulgaris)

Summary The physical measurements reported here test whether the European starling (Sturnus vulgaris) evaluates the azimuth direction of a sound source with a peripheral auditory system composed of two acoustically coupled pressure-difference receivers (1) or of two decoupled pressure receivers (2).A directional pattern of sound intensity in the freefield was measured at the entrance of the auditory meatus using a probe microphone, and at the tympanum using laser vibrometry. The maximum differences in the soundpressure level measured with the microphone between various speaker positions and the frontal speaker position were 2.4 dB at 1 and 2 kHz, 7.3 dB at 4 kHz, 9.2 dB at 6 kHz, and 10.9 dB at 8 kHz. The directional amplitude pattern measured by laser vibrometry did not differ from that measured with the microphone. Neither did the directional pattern of travel times to the ear. Measurements of the amplitude and phase transfer function of the starling's interaural pathway using a closed sound system were in accord with the results of the free-field measurements.In conclusion, although some sound transmission via the interaural canal occurred, the present experiments support the hypothesis 2 above that the starling's peripheral auditory system is best described as consisting of two functionally decoupled pressure receivers.Abbreviations CM cochlear microphonics - ITD interaural time difference - IID interaural intensity difference - MRA minimum resolvable angle - dB SPL sound-pressure level (re 0.00002 Pa)     

A temporal window for lateralization of interaural time difference by barn owls

Summary Lateralization of interaural time difference by barn owls (Tyto alba) was studied in a dichotic masking experiment. Sound bursts consisted of two parts: binaurally time-shifted noise, termed the probe, was inserted between masking noise. The owls indicated that they detected and lateralized the time-shift in the probe by a head turn in the direction predicted from sign of the time-shift.The general characteristics of head turns in response to this stimulus was similar to the head turns elicited by free-field stimulation or to head turns in response to presentation of the probe alone.The owls could easily lateralize stimuli containing long probes. The number of correct turns decreased as probe duration decreased, demonstrating that the masking noise interfered with the owls' ability to lateralize the probe. The minimal probe duration that the animals could lateralize (minimal duration) became shorter as burst duration decreased. Minimal durations ranged from 1 ms to 15 ms for the two subjects and burst durations from 10 to 100 ms.These findings suggested that owls possess a temporal window. A fitting procedure proposed by Moore et al. (1988) was used to determine the shape of the temporal window. The fitting procedure showed that the shape of the owls' binaural temporal window could be described by the same algorithms as the human monaural temporal window. Thus, the temporal window is composed of a short time constant that determines the central part of the window, and a longer time constant that determines the shape at the skirts of the window.Abbreviations ERD equivalent rectangular duration - ILD interaural level difference - ITD interaural time difference - RSE relative signal energy - SNR signal-to-noise ratio     

Precognitive and cognitive elements in sound localization

Sound localization behavior is of great importance for an animal's survival. To localize a sound, animals have to detect a sound source and assign a location to it. In this review we discuss recent results on the underlying mechanisms and on modulatory influences in the barn owl, an auditory specialist with very well developed capabilities to localize sound. Information processing in the barn owl auditory pathway underlying the computations of detection and localization is well understood. This analysis of the sensory information primarily determines the following orienting behavior towards the sound source. However, orienting behavior may be modulated by cognitive (top-down) influences such as attention. We show how advanced stimulation techniques can be used to determine the importance of different cues for sound localization in quasi-realistic stimulation situations, how attentional influences can improve the response to behaviorally relevant stimuli, and how attention can modulate related neural responses. Taken together, these data indicate how sound localization might function in the usually complex natural environment.     

Spatial orientation in the bushcricket <Emphasis Type="Italic">Leptophyes punctatissima</Emphasis> (Phaneropterinae; Orthoptera): I. Phonotaxis to elevated and depressed sound sources

Many species of acoustically interacting insects live in a complex, arboreal or semi-arboreal habitat. Thus mate finding by phonotaxis requires sound localization in the horizontal and vertical plane. Here we investigated the ability of the duetting bushcricket Leptophyes punctatissima to orient to one of three speakers, positioned at different levels in an artificial grid system, where each point in space could be reached by the male with almost equal probability. The system was designed analogous to a spherical calotte model of bismuth, where, once the male arrived at any nodal point had to decide between only three directions: either up or down and/or left and right. This design does not favour any phonotactic path of the males. All 12 males tested reached the three speaker positions (one in the horzontal plane, one elevated by 45°, one depressed by 45° relative to the starting position) with only little deviation from the shortest possible path. There was no significant difference with respect to the whole phonotactic time needed, the number of segments passed, or the number of stimuli received for the different speaker positions. This remarkable spatial orientation is achieved although the insects have no specialized external ear structures such as mammals, or some owls.     

Dipole source localization by mottled sculpin. III. Orientation after site-specific, unilateral denervation of the lateral line system

To test the hypothesis that spatial excitation patterns along the lateral-line system underlie source localization, we videotaped the orientation behavior of blinded mottled sculpin in response to a small dipole source (50-Hz vibrating sphere) before and after unilateral denervation of the lateral line system on different body regions (head, trunk and head + trunk). Approach pathways were qualitatively similar to those followed by normal intact animals. Abnormal behavior (turning in circles) was not observed. However, the frequency with which fish placed their intact side facing the source increased by 12–89%, depending on the denervation site. The angular accuracy of orientation decreased by 20° to 60° (100% to 370% change) depending on source location and region of lateral line denervated. Deficits tended to be site-specific. For example, unilaterally denervating lateral-line organs on the head resulted in less accurate orienting responses when the source was located on the denervated side of the head, but not on the opposite side of the head or on either side of the trunk. Site-specific deficits and the absence of abnormal approach pathways argue that animals are relying on a point-by-point spatial representation of source location along the sensory surface rather than computations based on bilateral comparisons. Accepted: 28 May 1998     

Spatial orientation in the bushcricket <Emphasis Type="Italic">Leptophyes punctatissima</Emphasis> (Phaneropterinae; Orthoptera): II. Phonotaxis to elevated sound sources on a walking compensator

The ability of the bushcricket Leptophyes punctatissima to orient to elevated sound sources was investigated. Males were placed on a walking compensator and oriented in response to a synthetic female reply, which was broadcast via one of five loudspeakers placed at elevations of 0°, 30°, 60°, 75° and 90°. Forward and backward movements were compensated, so that males remained at the same distance and elevation to the sound source. With increasing loudspeaker elevation, the males meandered more, and the ratio of the ideal path length to the actual path length decreased. The same was true for the correlation between stimulus angle and turn angle, and there were more turns to the wrong side with increasing loudspeaker elevation. Most males performed phonotaxis with a high acuity up to an elevation of 60°. Individuals varied strongly in their performance especially at a source elevation of 75°, where some were still very accurate in their approach, whereas the acuity of others decreased rapidly. We also describe a behaviour where males tilt their body axis to more anterior and sideward positions, both during walking and while calling on the spot. This behaviour is interpreted as a kind of directional scanning in order to actively induce changes in binaural cues.     

Effects of sound direction on the processing of amplitude-modulated signals in the frog inferior colliculus

Single-unit recordings were made from 143 neurons in the frog (Rana p. pipiens) inferior colliculus (IC) to investigate how free-field sound direction influenced neural responses to sinusoidal-amplitude-modulated (SAM) tone and/or noise. Modulation transfer functions (MTFs) were derived from 3 to 5 sound directions within 180° of frontal field. Five classes of MTF were observed: low-pass, high-pass, band-pass, multi-pass, and all-pass. For 64% of IC neurons, the MTF class remained unchanged when sound direction was shifted from contralateral 90° to ipsilateral 90°. However, the MTFs of more than half of these neurons exhibited narrower bandwidths when the loudspeaker was shifted to ipsilateral azimuths. There was a decrease in the cut-off frequency for neurons possessing low-pass MTFs, an increase in cut-off frequency for neurons showing high-pass MTFs, or a reduction in the pass-band for neurons displaying bandpass MTFs. These results suggest that sound direction can influence amplitude modulation (AM) frequency tuning of single IC neurons.Since changes in periodicity of SAM tones alter both the temporal parameters of sounds as well as the sound spectrum, we examined whether directional effects on spectral selectivity play a role in shaping the observed direction-dependent AM selectivity. The directional influence on AM selectivity to both SAM tone and SAM noise was measured in 62 neurons in an attempt to gain some insight into the mechanisms that underlie directionally-induced changes in AM selectivity. Direction-dependent changes in the shapes of the tone and noise derived MTFs were different for the majority of IC neurons (55/62) tested. These data indicate that a spectrally-based and a temporally-based mechanism may be responsible for the observed results.Abbreviations AM amplitude modulation - CF characteristic frequency - DI direction index - FR isointensity frequency response - GABA gamma-aminobutyric acid - IC inferior colliculus - ICc central nucleus of the inferior colliculus - ITD interaural time difference - MTF modulation transfer function - PSTH peri-stimulus time histogram - SAM sinusoidal-amplitude-modulated - SC synchronization coefficient - CN cochlear nucleus     

Spatial localization of electrotactile stimuli on the fingertip in humans

This study was designed to determine the extent to which sensations elicited by discrete electrotactile stimulation can be spatially localized, with a qualitative comparison to mechanical stimulation, in a 2 × 2 electrode array on the fingertip. Electrotactile stimulation was delivered in two modes: (1) same current to all locations (constant) or (2) current adjusted to perceptual threshold of each location (varied). For each stimulus location, subjects were asked to identify the location of the stimulus. Mechanical stimulation of the same locations on the fingerpad was delivered through von Frey hairs (0.07, 0.2 and 0.4 g). The percentage of accurate responses was computed for all stimulation modes. We found that the accuracy of discrimination of stimulus location in both the constant (46%%) and varied (40%%) electrotactile stimulation modes was significantly higher than chance level (25%%; p < 0.01). Furthermore, subjects were significantly more accurate in discriminating electrotactile stimuli in the constant than in the varied mode (p < 0.05). We also found that the accuracy of spatial discrimination was dependent on stimulation site for mechanical, but not electrotactile stimulation. Finally, we found a significant difference in accuracy over the duration of the experiment only for mechanical modes, which may indicate that electrotactile stimuli are less biased over time. These results suggest that, although low in accuracy, human subjects are able to extract spatial information from electrotactile stimuli. Further research is needed to optimize the amount of the information that can be delivered through electrotactile stimulation.     

Regulation of ROCKII by localization to membrane compartments and binding to DynaminI

ROCKII kinase activity is known to be regulated by Rho GTPase binding; however, the context-specific regulation of ROCKII is not clearly understood. We pursued the C-terminal PH domain as a candidate domain for regulating ROCKII function. A proteomics-based screen identified potential ROCKII signaling partners, a large number of which were associated with membrane dynamics. We used subcellular fractionation to demonstrate that ROCKII is localized to both the plasma membrane and internal endosomal membrane fractions, and then used microscopy to show that the C-terminal PH domain can localize to internal or peripheral membrane compartments, depending on the cellular context. Co-immunoprecipitation demonstrated that Dynamin1 is a novel ROCKII binding partner. Furthermore, blocking Dynamin function with a dominant negative mutant mimicked the effect of inhibiting ROCK activity on the actin cytoskeleton. Our data suggest that ROCKII is regulated by localization to specific membrane compartments and its novel binding partner, Dynamin1.     

Spatial tuning of neurons in the inferior colliculus of the big brown bat: effects of sound level, stimulus type and multiple sound sources

We examined factors that affect spatial receptive fields of single units in the central nucleus of the inferior colliculus of Eptesicus fuscus. Pure tones, frequency- or amplitude-modulated sounds, or noise bursts were presented in the free-field, and responses were recorded extracellularly. For 58 neurons that were tested over a 30 dB range of sound levels, 7 (12%) exhibited a change of less than 10° in the center point and medial border of their receptive field. For 28 neurons that were tested with more than one stimulus type, 5 (18%) exhibited a change of less than 10° in the center point and medial border of their receptive field.The azimuthal response ranges of 19 neurons were measured in the presence of a continuous broadband noise presented from a second loudspeaker set at different fixed azimuthal positions. For 3 neurons driven by a contralateral stimulus only, the effect of the noise was simple masking. For 11 neurons driven by sound at either side, 8 were unaffected by the noise and 1 showed a simple masking effect. For the remaining 2, as well as for 5 neurons that were excited by contralateral sound and inhibited by ipsilateral sound, the peak of the azimuthal response range shifted toward the direction of the noise.Abbreviations E/E excitation at either ear - I/E inhibition at the ipsilateral ear, excitation at the contralateral ear - O/E no effect from the ipsilateral ear, excitation at the contralateral ear - FM downward frequency modulation - FM upward frequency modulation - IC inferior colliculus - ICC central nucleus of the inferior colliculus - ILD interaural level difference - ITD interaural time difference - PT pure tone - SAM sinusoidally amplitude modulated sounds - SFM sinusoidally frequency modulated sounds     

Directional orientation of pomacentrid larvae to ambient reef sound

The mechanisms by which reef fish larvae locate settlement habitat at the end of their pelagic phase are unclear. We used an in situ binary choice chamber and an artificial source of reef sound to determine whether pomacentrid larvae can use ambient sound to locate reefs. Larvae were caught in light traps and then placed in a submerged binary choice chamber with an artificial source of reef sound ~80 m from one end of the chamber. At night, larvae moved towards the sound source; during the day, larvae showed no preference. These results suggest that pomacentrid larvae can detect reef sound and are capable of directional hearing. While other studies have shown that reef fish larvae respond to reef sound, and that the adults of some species can localize underwater sound sources, the localization of underwater sound by fish larvae has not been demonstrated previously.Communicated by Ecological Editor P.F. Sale     

Mechanisms of experience-dependent plasticity in the auditory localization pathway of the barn owl

Sound localization is a computational process that requires the central nervous system to measure various auditory cues and then associate particular cue values with appropriate locations in space. Behavioral experiments show that barn owls learn to associate values of cues with locations in space based on experience. The capacity for experience-driven changes in sound localization behavior is particularly great during a sensitive period that lasts until the approach of adulthood. Neurophysiological techniques have been used to determine underlying sites of plasticity in the auditory space-processing pathway. The external nucleus of the inferior colliculus (ICX), where a map of auditory space is synthesized, is a major site of plasticity. Experience during the sensitive period can cause large-scale, adaptive changes in the tuning of ICX neurons for sound localization cues. Large-scale physiological changes are accompanied by anatomical remodeling of afferent axons to the ICX. Changes in the tuning of ICX neurons for cue values involve two stages: (1) the instructed acquisition of neuronal responses to novel cue values and (2) the elimination of responses to inappropriate cue values. Newly acquired neuronal responses depend differentially on NMDA receptor currents for their expression. A model is presented that can account for this adaptive plasticity in terms of plausible cellular mechanisms. Accepted: 17 April 1999     

Use of local and global geometry from object arrays by adult humans

Disoriented men and women were trained to search for a goal hidden in front of one of four objects forming a rectangular-shaped array. The angular properties of these objects (either 50° or 75°) served as local geometric cues and the rectangular shape of the array served as global geometric cues. Upon successful completion of training, transformation tests were conducted during which either the local angle cues were removed and the global geometry was preserved (Global Cues test) or the local angles were preserved and the global geometry was removed (Local Cues test). A Cue Conflict test was also conducted which placed the local geometry in direct competition with the global geometry for control of search behavior. Results from testing showed that neither men nor women could successfully use only the global geometric cues provided by the shape of the array to reorient. Analyses of sex differences for the use of local cues revealed that men showed clear evidence that they had successfully encoded the local angular cues whereas women did not. Furthermore, the size of the training angle may have affected the encoding of local cues.     

Regulation of subcellular localization of the antiproliferative protein Tob by its nuclear export signal and bipartite nuclear localization signal sequences

Tob, a member of the Tob and BTG antiproliferative protein family, plays an important role in many cellular processes including cell proliferation. In this study, we have addressed molecular mechanisms regulating subcellular localization of Tob. Treatment with leptomycin B, an inhibitor of nuclear export signal (NES) receptor, resulted in a change in subcellular distribution of Tob from its pan-cellular distribution to nuclear accumulation, indicating the existence of NES in Tob. Our results have then identified an N-terminal region (residues 2-14) of Tob as a functional NES. They have also shown that Tob has a functional, bipartite nuclear localization signal (NLS) in residues 18-40. Thus, Tob is shuttling between the nucleus and the cytoplasm by its NES and NLS. To examine a possible relationship between subcellular distribution of Tob and its function, we exogenously added a strong NLS sequence or a strong NES sequence or both to Tob. The obtained results have demonstrated that the strong NLS-added Tob has a much weaker activity to inhibit cell cycle progression from G0/G1 to S phase. These results suggest that cytoplasmic localization or nucleocytoplasmic shuttling is important for the antiproliferative function of Tob.