A novel circuit in the bat's midbrain recruits neurons into sound localization processing.
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The role of GABAergic inhibition in shaping directional selectivity of bat inferior collicular neurons determined with temporally patterned pulse trains.
This study examined the role of GABAergic inhibition in shaping directional selectivity of neurons in the inferior colliculus of the big brown bat, Eptesicus fuscus. When determined with temporally patterned pulse trains at different pulse repetition rates, 93 inferior colliculus neurons displayed three types of directional selectivity curves. A directionally selective curve always showed a maximum to a certain azimuthal angle (the best angle). A hemifield curve showed a maximum to a range of contralateral azimuthal angles. A non-directional curve did not show a maximum to any particular azimuthal angles. Directional selectivity curves of 42% neurons changed from hemifield or non-directional to directionally selective and the best angles of 16-21% neurons shifted toward the midline with increasing pulse repetition rate of pulse trains. Directional selectivity curves of most (74%) neurons that discharged impulses to each pulse of a pulse train also became sharper with increasing pulse repetition rate of pulse trains. Bicuculline application produced more pronounced broadening of directional selective curves of inferior colliculus neurons at higher than at lower pulse repetition rates. As a result, pulse repetition rate-dependent directional selectivity of inferior colliculus neurons was abolished. Possible mechanisms and biological significance of these findings are discussed.
Action Potentials↗
The effect of temporal structure on rustling-sound detection in the gleaning bat, Megaderma lyra.
For a gleaning bat hunting prey from the ground, rustling sounds generated by prey movements are essential to invoke a hunting behaviour. The detection of prey-generated rustling sounds may depend heavily on the time structure of the prey-generated and the masking sounds due to their spectral similarity. Here, we systematically investigate the effect of the temporal structure on psychophysical rustling-sound detection in the gleaning bat, Megaderma lyra. A recorded rustling sound serves as the signal; the maskers are either Gaussian noise or broadband noise with various degrees of envelope fluctuations. Exploratory experiments indicate that the selective manipulation of the temporal structure of the rustling sound does not influence its detection in a Gaussian-noise masker. The results of the main experiment show, however, that the temporal structure of the masker has a strong and systematic effect on rustling-sound detection: When the width of irregularly spaced gaps in the masker exceeded about 0.3 ms, rustling-sound detection improved monotonically with increasing gap duration. Computer simulations of this experiment reveal that a combined detection strategy of spectral and temporal analysis underlies rustling-sound detection with fluctuating masking sounds.
Acoustic Stimulation↗
Echo delay versus spectral cues for temporal hyperacuity in the big brown bat, Eptesicus fuscus.
Big brown bats can discriminate between echoes that alternate in delay (jitter) by as little as 10-15 ns and echoes that are stationary in delay. This delay hyperacuity seems so extreme that it has been rejected in favor of an explanation in terms of artifacts in echoes, most likely spectral in nature, that presumably are correlated with delay. Using different combinations of digital, analog, and cable delays, we dissociated the overall delay of jittering echoes from the size of the analog component of delay, which alone is presumed to determine the strength of the apparatus artifact. The bats' performance remains invariant with respect to the overall delay of the jittering echoes, not with respect to the amount of analog delay. This result is not consistent with the possible use of delay-related artifacts produced by the analog delay devices. Moreover, both electronic and acoustic measurements disclose no spectral cues or impedance-mismatch reflections in delayed signals, just time-delays. The absence of artifacts from the apparatus and the failure of overlap and interference from reverberation to account for the 10-ns result means that closing the gap between the level of temporal accuracy plausibly explained from physiology and the level observed in behavior may require a better understanding of the physiology.
Acoustic Stimulation↗
Ambient temperature affects mechanosensory host location in a parasitic wasp.
Certain parasitic wasps (Ichneumonidae, Pimplinae) use self-produced vibrations transmitted on plant substrate to locate their immobile concealed hosts (i.e. lepidopteran pupae). This mechanosensory mechanism, called the vibrational sounding, depends both on physical cues of the environment and physical activity of the parasitoid and is postulated to depend on ambient temperature. We analysed the influences of temperature on vibrational sounding by choice experiments using plant-stem models with hidden host mimics in the temperate species Pimpla turionellae. The results show a significant effect of temperature on host-location activity and on the success of this process. Outside an optimum range, the performance of the wasps decreased both at low and high temperatures. Below 10 degrees C and beyond 24 degrees C, the wasps displayed (1) substantial reduction in responsiveness, i.e. proportion of females showing ovipositor insertions, (2) reduction of quantitative activity with ovipositor insertions in the individuals, and (3) reduced precision of mechanosensory host location. Nevertheless, female wasps were able to locate their host over a surprisingly broad range of ambient temperatures which indicates that the wasps are able to compensate for temperature effects on vibrational sounding.
Animals↗
Diversity in sound pressure levels and estimated active space of resident killer whale vocalizations.
Signal source intensity and detection range, which integrates source intensity with propagation loss, background noise and receiver hearing abilities, are important characteristics of communication signals. Apparent source levels were calculated for 819 pulsed calls and 24 whistles produced by free-ranging resident killer whales by triangulating the angles-of-arrival of sounds on two beamforming arrays towed in series. Levels in the 1-20 kHz band ranged from 131 to 168 dB re 1 microPa at 1 m, with differences in the means of different sound classes (whistles: 140.2+/-4.1 dB; variable calls: 146.6+/-6.6 dB; stereotyped calls: 152.6+/-5.9 dB), and among stereotyped call types. Repertoire diversity carried through to estimates of active space, with "long-range" stereotyped calls all containing overlapping, independently-modulated high-frequency components (mean estimated active space of 10-16 km in sea state zero) and "short-range" sounds (5-9 km) included all stereotyped calls without a high-frequency component, whistles, and variable calls. Short-range sounds are reported to be more common during social and resting behaviors, while long-range stereotyped calls predominate in dispersed travel and foraging behaviors. These results suggest that variability in sound pressure levels may reflect diverse social and ecological functions of the acoustic repertoire of killer whales.
Animals↗
Intra-individual variation in the vocalized frequency of the Taiwanese leaf-nosed bat, Hipposideros terasensis, influenced by conspecific colony members.
We examined the intra-individual variation in resting frequency of the constant-frequency component of the second harmonic of the pulse (F (rest)) over 4 years in a laboratory colony of the Taiwanese leaf-nosed bat (Hipposideros terasensis). Patterns of change in F (rest) were observed when individuals were added to or removed from the colony so that we investigated whether F (rest) was affected by neighboring colony members. F (rest) of each bat continually showed a long-term gradual change throughout the year, and all bats in the colony increased or decreased their F (rest) in the same direction as a group non-seasonally. The greatest short-term changes were observed when new bats with a relatively low F (rest) joined the colony and F (rest) of new bats converged with those of the original colony members around 8 -16 days after their introduction. Conversely, a single individual showed sudden short-term decrease in F (rest) after its isolation from other colony members. These findings strongly indicate that F (rest) is flexible according to the presence of neighboring conspecific bats. We suggest that the audio-vocal feedback for conspecific pulses appears to be involved in the short- or long-term intra-individual variation in F (rest) other than factors previously thought such as age or season.
Adaptation, Physiological↗
GABA-mediated echo duration selectivity of inferior collicular neurons of Eptesicus fuscus, determined with single pulses and pulse-echo pairs.
When insectivorous bats such as Eptesicus fuscus emit ultrasonic signals and analyze the returning echoes to hunt insects, duration selectivity of auditory neurons plays an important role in echo recognition. The success of prey capture indicates that they can effectively encode progressively shortened echo duration throughout the hunting process. The present study examines the echo duration selectivity of neurons in the central nucleus of the bat inferior colliculus (IC) under stimulation conditions of single pulses and pulse-echo (P-E) pairs. This study also examines the role of gamma-aminobutyric acid (GABA)ergic inhibition in shaping echo duration selectivity of IC neurons. The data obtained show that the echo duration selectivity of IC neurons is sharper when determined with P-E pairs than with single pulses. Echo duration selectivity also sharpens with shortening of pulse duration and P-E gap. Bicuculline application decreases and GABA application increases echo duration selectivity of IC neurons. The degree of change in echo duration selectivity progressively increases with shortening of pulse duration and P-E gap during bicuculline application while the opposite is observed during the GABA application. These data indicate that the GABAergic inhibition contributes to sharpening of echo duration selectivity of IC neurons and facilitates echo recognition by bats throughout different phases of hunting.
Acoustic Stimulation↗
Detection of jitter in intertarget spacing by the big brown bat Eptesicus fuscus.
We trained bats to detect intertarget jitter, i.e., relative motion between two virtual (electronically synthesized) targets. Both targets were themselves moving with respect to nearby objects (e.g., the microphone and speaker used to create the virtual targets) so that the only reliable cue available to the bats was variation in intertarget spacing. Given a target at 80 cm and another at 95, 110 or 125 cm, the threshold for intertarget jitter (ITJ) of the two bats tested was < 10 microseconds, corresponding to < 1.7 mm of range. When, for one bat, we increased the range instability of the targets by adding varying amounts of random range shift to the target complex (while preserving the correct intertarget spacing), ITJ threshold worsened. When we presented three targets, one of which was jittering, the bat's threshold improved to 0.9 microsecond (equivalent to 0.16 mm). If no second target was presented, i.e., if the task was to detect jitter added to a single moving target, then bats' jitter threshold was very high (> 200 microseconds). Eptesicus fuscus appears to be very good at detecting changes in intertarget spacing, which might prove valuable for detecting targets moving relative to the background or for constructing a spatial image of a complex environment.
Animals↗
Echo SPL, training experience, and experimental procedure influence the ranging performance in the big brown bat, Eptesicus fuscus.
Four Eptesicus fuscus were trained in a range discrimination experiment to choose the closer of two phantom targets. Echo attenuation was roving between trials returning echoes ranging from -10 dB to -50 dB SPL (sound pressure level) relative to emission SPL. Discrimination thresholds were determined. After sufficient training, ranging performance was stable and about the same in the range between -20 dB and -50 dB with range difference thresholds around 300 microseconds. At -10 dB, performance was poor even after long training. After additional training at a constant relative echo SPL of -30 dB and a delay difference of 300 microseconds the performance measured with roving echo SPL improved at all relative echo SPL between -20 dB and -50 dB but not at -10 dB. The new experimental procedure improved the performance by additional learning, and the bats generalized over a wide range of relative echo SPL. Threshold improved to 100 microseconds when measured at a constant relative echo SPL of -30 dB, again indicating the influence of the experimental procedure. In correspondence to neurophysiological data the ranging performance deteriorates if the echo SPL is close to the emission SPL. Signal duration and emission SPL were variable during range discrimination.
Animals↗
Temporally patterned pulse trains affect duration tuning characteristics of bat inferior collicular neurons.
This study examines the effect of temporally patterned pulse trains on duration tuning characteristics of inferior collicular neurons of the big brown bat. Eptesicus fuscus, under free-field stimulation conditions. Using a 50% difference between maximal and minimal responses as a criterion, the duration tuning characteristics of inferior collicular neurons determined with pulse trains of different pulse durations are described as band-pass, long-pass, short-pass, and all-pass. Each band-pass neuron discharged maximally to a specific pulse duration that was at least 50% larger than the neuron's responses to a long- and a short-duration pulse. In contrast, each long- or short-pass neuron discharged maximally to a range of long- or short-duration pulses that were at least 50% larger than the minimal responses. The number of impulses of an all-pass neuron never differed by more than 50%. When pulse trains were delivered at different pulse repetition rates, the number of short-pass and band-pass neurons progressively increased with increasing pulse repetition rates. The slope of the duration tuning curves also became sharper when determined with pulse trains at high pulse repetition rates. Possible mechanisms underlying these findings are discussed.
Acoustic Stimulation↗
Sound transmission in the nose of the sperm whale Physeter catodon. A post mortem study.
During a sperm whale stranding at Rømø, the Wadden Sea, Denmark, on 4 December 1997, we were notified in time to start acoustic transmission measurements in the spermaceti complex 1 h after the specimen was seen alive. Frequency-modulated sound pulses, sweeping from 30 kHz to 10 kHz in 25 ms, were injected at the frontal surface at two positions: at the distal sac, and at the center of the junk (a compartmentalized structure below the spermaceti organ). A hydrophone next to the projector served as receiver. The analyses of the recordings show a repetitive, decaying reflection pattern at both projection sites, reminiscent of the multi-pulse click peculiar to sperm whales, although with minor differences in the duration of the intra-click intervals. This experimental evidence supports the Norris and Harvey (1972) theory of click generation in the spermaceti organ. Accordingly, the click is composed of a primary event, followed by a train of reflected pulses, spaced by the time required for the event to travel back and forth between air sacs (reflectors) at each end of the organ. The results also show that the junk readily transmits sound and probably is in acoustic contact with the spermaceti organ.
Animals↗
Thalamic projections to the auditory cortex in the rufous horseshoe bat (Rhinolophus rouxi). II. Dorsal fields.
In this study, we analyzed the thalamic connections to the parietal or dorsal auditory cortical fields of the horseshoe bat, Rhinolophus rouxi. The data of the present study were collected as part of a combined investigation of physiologic properties, neuroarchitecture, and chemoarchitecture as well as connectivity of cortical fields in Rhinolophus, in order to establish a neuroanatomically and functionally coherent view of the auditory cortex. Horseradish peroxidase or wheat-germ-agglutinated horseradish peroxidase deposits were made into cortical fields after mapping response properties. The dorsal fields of the auditory cortex span nearly the entire parietal region and comprise more than half of the non-primary auditory cortex. In contrast to the temporal fields of the auditory cortex, which receive input mainly from the ventral medial geniculate body (or "main sensory nucleus"), the dorsal fields of the auditory cortex receive strong input from the "associated nuclei" of the medial geniculate body, especially from the anterior dorsal nucleus of the medial geniculate body. The anterior dorsal nucleus is as significant for the dorsal fields of the auditory cortex as the ventral nucleus of the medial geniculate body is for the temporal fields of the auditory cortex. Additionally, the multisensory nuclei of the medial geniculate body provide a large share of the total input to the nonprimary fields of the auditory cortex. Comparing the organization of thalamic auditory cortical afferents in Rhinolophus with other species demonstrates the strong organizational similarity of this bat's auditory cortex with that of other mammals, including primates, and provides further evidence that the bat is a relevant and valuable model for studying mammalian auditory function.
Animals↗
Mechanisms by which dolphins identify target movement characteristics and their technical use in hydrolocation and radiolocation.
The ability of the localizing system of the dolphin Tursiops truncatus to discriminate the dynamic characteristics of an object being located (a radially moving target) was studied. Measurements were made of the thresholds of the animals' sensitivity to the target speed (2.6 cm/sec) and acceleration (0.6 cm/sec(2)). For location of a target moving at constant speed, dolphins were found to use probe signals consisting of two impulses, while accelerating targets were located using probe signals consisting of three impulses. These characteristics were used to propose new and highly effective technical methods for hydrolocation and radiolocation.
Animals↗
Heterogeneity of lipid composition within the cephalic melon tissue of the pilot whale (Globicephala melaena).
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Adipose Tissue↗
Coding of fine frequency information by echoranging neurons in the inferior colliculus of the Mexican free-tailed bat.
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Acoustic Stimulation↗
Mapping of the auditory area in the cerebellar vermis and hemispheres of the little brown bats, Myotis lucifugus.
Mapping of auditory area in the cerebellar vermis and hemispheres of little brown bats, Myotis lucifugus, shows that a large area of the bat's cerebellum contains units responding to acoustic stimuli. These units had latencies between 4 and 34 msec and best frequencies between 33.0 and 92.5 kHZ. The Q10-dB values of their tuning curves ranged from 2.0 to 19.7. Most of the units studied fired only a few impulses during a stimulus with minimum thresholds between 22 and 90 dB SPL. Units in the cerebellar vermis tend to have higher best frequencies and shorter latencies than those in the cerebellar hemispheres. However, there is no evidence of clear tonotopic organization therein.
Animals↗
Localization of brain stem motoneurons innervating the laryngeal muscles in the rufous horseshoe bat, Rhinolophus rouxi.
The motoneurons innervating the laryngeal muscles were localized in the rufous horseshoe bat, Rhinolophus rouxi, using the HRP method. HRP was applied to the cricothyroid muscle and to the cut end of the recurrent laryngeal nerve. Labeled motoneurons were found in two completely separated regions of the nucleus ambiguus. The motoneurons innervating the cricothyroid muscle via the superior laryngeal nerve (SLN) are located within the ventrolateral portion of the nucleus reaching the caudal pole of the motor nucleus of the facial nerve. The motoneurons innervating the other intrinsic laryngeal muscles via the recurrent laryngeal nerve (RLN) are situated in the caudal half of the nucleus ambiguus. The innervation is strictly homolateral.
Animals↗