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PubMed1982

Origin of ascending projections to inferior colliculus in the mustache bat, Pteronotus parnellii.

The origins of pathways to the inferior colliculus of the mustache bat were identified by retrograde transport of horseradish peroxidase (HRP). A specific goal of this study was to obtain evidence that would help determine whether the nuclei, shown in the previous paper to have unusual cytoarchitectural features, are unique to bats, or whether they are homologous to areas that are not well differentiated in other mammals. The auditory pathways in the lower brain stem of Pteronotus appear to conform to the same basic organization as in other mammals: After injection of HRP into one inferior colliculus, labeled cells are located contralaterally in the cochlear nucleus, ipsilaterally in the medical superior olive, bilaterally in the lateral superior olive, ipsilaterally in the ventral and intermediate nuclei of the lateral lemniscus, and bilaterally in the dorsal nucleus of the lateral lemniscus. These patterns of labeling provide a basis for understanding how the specialized auditory areas of the bat may be organized within a basic plan of mammalian auditory systems. In the anteroventral cochlear nucleus the unusually small spherical cells seem to be homologous to stellate cells in the anteroventral cochlear nucleus of the cat. In the superior olive, differences in patterns of labeled cells distinguish the medial from the lateral superior olive. In the lateral lemniscus the pattern of labeled cells shows clear differences between the two special parts, intermediate and ventral nuclei, as well as between these and the dorsal nucleus of the lateral lemniscus. The results are consistent with the hypothesis that the unusual auditory nuclei of the bat have homologues in mammals whose auditory systems are not specialized for echolocation.

Animals↗
PubMed1989

Functional organization of mustached bat inferior colliculus: II. Connections of the FM2 region.

Echolocating bats estimate target distance by analyzing the time delay between frequency-modulated portions of their emitted ultrasonic vocalizations and the resultant echoes. In the companion paper we investigated, in the central nucleus of the inferior colliculus, the representation of the predominant second-harmonic frequency-modulated component (FM2) of the mustached bat biosonar signal (O'Neill et al.: J. Comp. Neurol. 283:000-000,'89). In the present paper we report the connections of this part of the colliculus, as determined by focal, iontophoretic injections of HRP following single-unit mapping of the FM2 representation. It was found that the major inputs to the FM2 region of the inferior colliculus come from the contralateral cochlear nucleus; ipsilaterally from the medial superior olive, periolivary nuclei, and ventral and intermediate nuclei of the lateral lemniscus; and bilaterally from the lateral superior olive and dorsal nucleus of the lateral lemniscus. This study identifies for the first time those specific regions of brainstem nuclei providing input to the central nucleus of the inferior colliculus that process FM2 information in the mustached bat. The primary outputs of the FM2 region project to the medial and dorsal divisions of the medial geniculate body. In sharp contrast to other mammals, we found little evidence of connections to the ventral division of the medial geniculate. Other regions receiving significant inputs from the FM2 area include the deep superior colliculus ipsilaterally and the ipsilateral lateral pontine nuclei. Some fibers also terminated near the midline in the dorsal midbrain periaqueductal gray. Sparse intrinsic connections were also seen to the ipsilateral dorsoposterior division of the central nucleus and to the contralateral inferior colliculus at a location homologous to the injection site in the anterolateral division. The finding that FM2 projections to the medial geniculate heavily favor the medial and dorsal divisions is consistent with the location of "FM-FM" delay-dependent facilitation neurons found by Olsen (Processing of Biosonar Information by the Medical Geniculate Body of the Mustached Bat, Pteronotus parnellii. Dissertation, Washington Univ., St. Louis, '86) in these divisions, and with thalamocortical projection patterns in this species. These findings demonstrate that for the FM portions of the biosonar signal, a transformation from a tonotopic form of processing to a more specialized, convergent pattern of organization occurs at the level of the inferior colliculus outputs.

Animals↗
PubMed1996

Neurons with different temporal firing patterns in the inferior colliculus of the little brown bat differentially process sinusoidal amplitude-modulated signals.

We examined how well single neurons in the inferior colliculus (IC) of an FM bat (Myotis lucifugus) processed simple tone bursts of different duration and sinusoidal amplitude-modulated (SAM) signals that approximated passively heard natural sounds. Units' responses to SAM tones, measured in terms of average spike count and firing synchrony to the modulation envelope, were plotted as a function of the modulation frequency to construct their modulation transfer functions. These functions were classified according to their shape (e.g., band-, low-, high-, and all-pass). IC neurons having different temporal firing patterns to simple tone bursts (tonic, chopper, onset-late, and onset-immediate) exhibited different selectivities for SAM signals. All tonic and 83% of chopper neurons responded robustly to SAM signals and displayed a variety of spike count-based response functions. These neurons showed a decreased level of time-locking as the modulation frequency was increased, and thereby gave low-pass synchronization-based response functions. In contrast, 64% of onset-immediate, 37% of onset-late and 17% of chopper units failed to respond to SAM signals at any modulation frequency tested (5-800 Hz). Those onset neurons that did respond to SAM showed poor time-locking (i.e., non-significant levels of synchronization). We obtained evidence that the poor SAM response of some onset and chopper neurons was due to a preference for short-duration signals. These data suggest that tonic and most chopper neurons are better-suited for the processing of long-duration SAM signals related to passive hearing, whereas onset neurons are better-suited for the processing of short, pulsatile signals such as those used in echolocation.

Animals↗
PubMed1990

Ontogenesis of auditory fovea representation in the inferior colliculus of the Sri Lankan rufous horseshoe bat, Rhinolophus rouxi.

This report describes the ontogenesis of tonotopy in the inferior colliculus (IC) of the rufous horseshoe bat (Rhinolophus rouxi). Horseshoe bats are deaf at birth, but consistent tonotopy with a low-to-high frequency gradient from dorsolateral to ventromedial develops from the 2nd up to the 5th week. The representation of the auditory fovea is established in ventro-medio-caudal parts of the IC during the 3rd postnatal week (Fig. 3). Then, a narrow frequency band 5 kHz in width, comprising 16% of the bat's auditory range, captures 50-60 vol% of the IC (Fig. 3c). However, foveal tuning is 10-12 kHz (1/3 octave) lower than in adults; foveal tuning in females (65-68 kHz) is 2-3 kHz higher than in males (62-65 Khz). Thereafter, foveal tuning increases by 1-1.5 kHz per day up to the 5th postnatal week, when the adult hearing range is established (Figs. 4, 5). The increase of sensitivity and of tuning sharpness of single units also follows a low-to-high frequency gradient (Fig. 6). Throughout this development the foveal tuning matches the second harmonic of the echolocation pulses vocalised by these young bats. The results confirm the hypothesis of developmental shifts in the frequency-place code for the foveal high frequency representation in the IC.

Acoustic Stimulation↗
PubMed1991

Encoding repetition rate and duration in the inferior colliculus of the big brown bat, Eptesicus fuscus.

1. Encoding of temporal stimulus parameters by inferior collicular (IC) neurons of Eptesicus fuscus was studied by recording their responses to a wide range of repetition rates (RRs) and durations at several stimulus intensities under free field stimulus conditions. 2. The response properties of 424 IC neurons recorded were similar to those reported in previous studies of this species. 3. IC neurons were classified as low-pass, band-pass, and high-pass according to their preference for RRs and/or durations characteristic of, respectively, search, approach, or terminal phases of echolocation. These neurons selectively process stimuli characteristic of the various phases of hunting. 4. Best RRs and best durations were not correlated with either the BFs or recording depths This suggests that each isofrequency lamina is capable of processing RRs and durations of all hunting phases. 5. Responses of one half of IC neurons studied were correlated with the stimulus duty cycle. These neurons may preferentially process terminal phase information when the bat's pulse emission duty cycle increases. 6. While the stimulus RR affected the dynamic range and overall profile of the intensity rate function, only little effect was observed with different stimulus durations.

Acoustic Stimulation↗
PubMed1993

Labile cochlear tuning in the mustached bat. II. Concomitant shifts in neural tuning.

Acoustic stimuli near 60 kHz elicit pronounced resonance in the cochlea of the mustached bat (Pteronotus parnellii parnellii). The cochlear resonance frequency (CRF) is near the second harmonic, constant frequency (CF2) component of the bat's biosonar signals. Within narrow bands where CF2 and third harmonic (CF3) echoes are maintained, the cochlea has sharp tuning characteristics that are conserved throughout the central auditory system. The purpose of this study was to examine the effects of temperature-related shifts in the CRF on the tuning properties of neurons in the cochlear nucleus and inferior colliculus. Eighty-two single and multi-unit recordings were characterized in 6 awake bats with chronically implanted cochlear microphonic electrodes. As the CRF changed with body temperature, the tuning curves of neurons sharply tuned to frequencies near the CF2 and CF3 shifted with the CRF in every case, yielding a change in the unit's best frequency. The results show that cochlear tuning is labile in the mustached bat, and that this lability produces tonotopic shifts in the frequency response of central auditory neurons. Furthermore, results provide evidence of shifts in the frequency-to-place code within the sharply tuned CF2 and CF3 regions of the cochlea. In conjunction with the finding that biosonar emission frequency and the CRF shift concomitantly with temperature and flight, it is concluded that the adjustment of biosonar signals accommodates the shifts in cochlear and neural tuning that occur with active echolocation.

Acoustic Stimulation↗
PubMed1993

Discrimination of two-wavefront echoes by the big brown bat, Eptesicus fuscus: behavioral experiments and receiver simulations.

1. Echolocating bats (Eptesicus fuscus) were trained to discriminate between simulated targets consisting of one or two echo-wavefronts with internal time delays of up to 100 microseconds. Spectral and temporal properties and total signal energy of the targets were evaluated and predictions for performances of bats derived from receiver models were compared with measured performances. 2. Eptesicus fuscus was able to discriminate a one-wavefront target from two-wavefront targets with distinct internal time delays (12 microseconds, 32-40 microseconds and 52-100 microseconds). Performance was not affected by changes in total signal energy. Bats also successfully discriminated between two-wavefront targets with different internal time delays. 3. Performance predicted from differences in total energy between targets did not match the measured performance, indicating that bats did not rely on total echo energy. This finding is also supported by the behavioral data. Performance predicted from spectral and temporal receiver models both matched the measured performance and, therefore, neither one of these models can be favored over the other. 4. The behavioral data suggest that Eptesicus fuscus did not transform echo information into estimates of target range separation and, therefore, did not perceive the two wavefronts of each simulated two-wavefront echo as two separate targets.

Acoustic Stimulation↗
PubMed1996

The orientation behaviour of the lesser spearnosed bat, Phyllostomus discolor (Chiroptera) in a model roost. Concurrence of visual, echoacoustical and endogenous spatial information.

The orientation behaviour of bats (Phyllostomus discolor, Phyllostomidae), flying inside an octagonal "roost-like" chamber (phi: 100 cm; h: 150 cm) was examined. It has been shown that the bats begin turning manoeuvres during flight by turning their head towards the direction they intend to proceed to. During early phases of the flights, cumulative navigation errors were evident, indicating that endogenous spatial information plays a major role in the orientation of the bats. During later phases of the flight this error is diminished again. So it can be concluded that the bats start to use exogenous spatial information for orientation while approaching the target. In order to investigate the relative importance of vision, echolocation and endogenous spatial information for approaching the roost, the landing lattices inside the test arena were changed for non-grid dummies. We found that: 1. combined visual and endogenous information are more important than echoacoustical cues, 2. the bats learned quickly to switch their orientation behaviour in order to get a better performance in avoiding the dummies, 3. the learning performance was influenced by the visual similarity of dummies and the real landing lattice.

Animals↗
PubMed1996

Hearing in the FM-bat Phyllostomus discolor: a behavioral audiogram.

Absolute auditory thresholds of six adult lesser spear-nosed bats Phyllostomus discolor (Chiroptera, Phyllostomidae) were determined in a two-alternative forced-choice procedure. Behavioral response to pure tone stimuli could be elicited throughout the tested frequency range of 5-142 kHz. The shape of the average audiogram is characterized by two sensitivity peaks and a pronounced increase of thresholds around 55 kHz, and towards the limits of the tested frequency range. The spectral extent of both sensitivity peaks shows a close relation to the bandwidth of two types of species-specific vocalizations. The first low threshold area (> 10 and < 55 kHz) of the audiogram seems perfectly adapted to the directive call used for intraspecifc communication, whereas the second sensitivity peak, centered around 85 kHz, covers most of the bandwidth of the species' echolocation calls.

Acoustic Stimulation↗
PubMed1986

The detection of phantom targets in noise by serotine bats; negative evidence for the coherent receiver.

Using a target simulator three serotine bats, Eptesicus serotinus, were trained to judge whether a phantom target was present or absent. The echolocation sounds emitted by the bats during the detection were intercepted by a microphone, amplified and returned by a loudspeaker as an artificial echo, with a delay of 3.2 ms and a sound level determined by the overall gain and cry amplitude. The cry level of each pulse was measured and the echo level received by the bat was calculated. The target was presented in 50% of the trials and the gain adjusted using conventional up/down procedures. Under these conditions between 40 and 48 dB peSPL were required for 50% detection (Figs. 2, 3). In a subsequent experiment the phantom target was masked with white noise (No) with a spectrum level of -113 dB re. 1 Pa X Hz-1/2. The thresholds were increased by 7-14 dB. Energy density (S) of a single pulse was measured and used to estimate S/No, which ranged from 36-49 dB at threshold. Theoretically the coherent receiver model predicts the ratio between hits and false alarms observed for the bats at a S/No of ca. 1-2 dB. Since the bats require 40-50 dB higher S/No (Fig. 3), this is taken as negative evidence for coherent reception (cross correlation). Furthermore, a strong sensitivity to clutter was found since there seemed to exist a fixed relationship between thresholds and clutter level.

Animals↗
PubMed1985

Responses to pure tones and linear FM components of the CF-FM biosonar signal by single units in the inferior colliculus of the mustached bat.

The responses of 682 single-units in the inferior colliculus (IC) of 13 mustached bats (Pteronotus parnellii parnellii) were measured using pure tones (CF), frequency modulations (FM) and pairs of CF-FM signals mimicking the species' biosonar signal, which are stimuli known to be essential to the responses of CF/CF and FM-FM facilitation neurons in auditory cortex. Units were arbitrarily classified into 'reference frequency' (RF), 'FM2' and 'Non-echolocation' (NE) categories according to the relationship of their best frequencies (BF) to the biosonar signal frequencies. RF units have high Q10dB values and are tuned to the reference frequency of each bat, which ranged between 60.73 and 62.73 kHz. FM2 units had BF's between 50 and 60 kHz, while NE units had BF's outside the ranges of the RF and FM2 classes. PST histograms of the responses revealed discharge patterns such as 'onset', 'onset-bursting' (most common), 'on-off', 'tonic-on','pauser', and 'chopper'. Changes in discharge patterns usually resulted from changes in the frequency and/or intensity of the stimuli, most often involving a change from onset-bursting to on-off. Different patterns were also elicited by CF and FM stimuli. Frequency characteristics and thresholds to CF and FM stimuli were measured. RF neurons were very sharply tuned with Q10dB's ranging from 50-360. Most (92%) also responded to FM2 stimuli, but 78% were significantly more sensitive (greater than 5 dB) to CF stimuli, and only 3% had significantly lower thresholds to FM2. The best initial frequency for FM2 sweeps in RF units was 65.35 +/- 2.138 kHz (n = 118), well above the natural frequency of the 2nd harmonic. FM2 and NE units were indistinguishable from each other, but were quite different from RF units: 41% of these two classes had lower thresholds to CF, 49% were about equally sensitive, and 10% had lower thresholds to FM. For FM2 units, mean best initial frequency for FM was 60.94 kHz +/- 3.162 kHz (n = 114), which is closely matched to the 2nd harmonic in the biosonar signal. Very few units (5) responded only to FM signals, i.e., were FM-specialized. The characteristics of spike-count functions were determined in 587 units. The vast majority (79%) of RF units (n = 228) were nonmonotonic, and about 22% had upper-thresholds.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1976

Variations in lipid composition and sound velocity in melon from the North Atlantic Pilot whale, globicephala melaena melaena.

Nonpolar lipids and sound velocity at different distances from the skin surface within a sample of pilot whale melon were studied. Thin layer chromatography, sensitive radioisotopic methods, and an ultrasonic echo scanner were used. Wax esters had a maximum concentration at 11-12 cm from the skin surface. Non- and monoisovalero triglycerides both exhibited a minimum at 8-10 cm, whereas the diisovalero species increased steadily from the skin surface towards the center of the pilot whale head. A minimum of sound velocity, ca. 1340 m/s, was found at 9-11 cm, thus coinciding with a region rich in isovaleric acid esterified in wax and diisovalero triglycerides. These findings are compatible with predictions based on the known correlations between lipid structure and sound velocity. Consequently, additional evidence for refraction of sound in pilot whale melon is presented. This is important in understanding the system of echolocation of toothed whales.

Animals↗
PubMed2001

Cryptic mammalian species: a new species of whiskered bat (Myotis alcathoe n. sp.) in Europe.

The analysis of morphological, behavioural and genetic characters of whiskered bats revealed a new European bat species within the family Vespertilionidae. We describe the morphology, karyology, genetic similarity, ecology and distribution of Myotis alcathoe n. sp. It closely resembles Myotis mystacinus, Myotis brandtii and Myotis ikonnikovi in morphology, but all four species show clear genetic differences in two mitochondrial genes (ND1 and 12S rRNA). Myotis alcathoe n. sp. is the smallest species among the European whiskered bats and uses the highest-frequency echolocation calls of all the European Myotis species. It prefers to hunt in small valleys with deciduous trees and flowing water, which is an endangered habitat. Records from Greece and Hungary indicate a distribution range in south-eastern Europe.

Animals↗
PubMed2001

Interaction of vibrational and visual cues in parasitoid host location.

Female parasitoids are guided by multisensory information during host finding. Individual cues are used in an interactive or a hierarchical manner according to the relative importance on the spatial scale of their effect. Unlike most studies that concentrate on single cues, the present paper investigates the interaction of two physical cues. The interaction of mechanosensory and visual cues was studied in the pupal parasitoid Pimpla turionellae (Hymenoptera: Ichneumonidae). This species uses, amongst other senses, vibrational sounding (echolocation in a solid substrate) to find its mainly endophytic hosts. Location and frequency of ovipositor insertions were scored on cylindrical plant stem models with single or combined cues. Single-cue experiments show that parasitoids use both visual and mechanosensory cues and achieve a similar precision of host location with either cue. The combination of vision and vibrational sounding increased the precision of host location by a factor of approximately two to three. We conclude that the two senses interact, resulting in an additive accuracy. Neither the visual nor the mechanosensory cue was favored when offered adjacent to each other on the same stem model. On the investigated spatial scale, both physical cues are used and seem to be equally important for host location in this species.

Animals↗
PubMed2003

Fine control of call frequency by horseshoe bats.

The auditory system of horseshoe bats is narrowly tuned to the sound of their own echoes. During flight these bats continuously adjust the frequency of their echolocation calls to compensate for Doppler-effects in the returning echo. Horseshoe bats can accurately compensate for changes in echo frequency up to 5 kHz, but they do so through a sequence of small, temporally-independent, step changes in call frequency. The relationship between an echo's frequency and its subsequent impact on the frequency of the very next call is fundamental to how Doppler-shift compensation behavior works. We analyzed how horseshoe bats control call frequency by measuring the changes occurring between many successive pairs of calls during Doppler-shift compensation and relating the magnitude of these changes to the frequency of each intervening echo. The results indicate that Doppler-shift compensation is mediated by a pair of (echo)frequency-specific sigmoidal functions characterized by a threshold, a slope, and an upper limit to the maximum change in frequency that may occur between successive calls. The exact values of these parameters necessarily reflect properties of the underlying neural circuitry of Doppler-shift compensation and the motor control of vocalization, and provide insight into how neural feedback can accommodate the need for speed without sacrificing stability.

Animals↗
PubMed2003

Efficiency of vibrational sounding in parasitoid host location depends on substrate density.

Parasitoids of concealed hosts have to drill through a substrate with their ovipositor for successful parasitization. Hymenopteran species in this drill-and-sting guild locate immobile pupal hosts by vibrational sounding, i.e., echolocation on solid substrate. Although this host location strategy is assumed to be common among the Orussidae and Ichneumonidae there is no information yet whether it is adapted to characteristics of the host microhabitat. This study examined the effect of substrate density on responsiveness and host location efficiency in two pupal parasitoids, Pimpla turionellae and Xanthopimpla stemmator (Hymenoptera: Ichneumonidae), with different host-niche specialization and corresponding ovipositor morphology. Location and frequency of ovipositor insertions were scored on cylindrical plant stem models of various densities. Substrate density had a significant negative effect on responsiveness, number of ovipositor insertions, and host location precision in both species. The more niche-specific species X. stemmator showed a higher host location precision and insertion activity. We could show that vibrational sounding is obviously adapted to the host microhabitat of the parasitoid species using this host location strategy. We suggest the attenuation of pulses during vibrational sounding as the energetically costly limiting factor for this adaptation.

Animals↗
PubMed2005

Temporal resolution of the Risso's dolphin, Grampus griseus, auditory system.

Toothed whales and dolphins (Odontocetes) are known to echolocate, producing short, broadband clicks and receiving the corresponding echoes, at extremely rapid rates. Auditory evoked potentials (AEP) and broadband click stimuli were used to determine the modulation rate transfer function (MRTF) of a neonate Risso's dolphin, Grampus griseus, thus estimating the dolphin's temporal resolution, and quantifying its physiological delay to sound stimuli. The Risso's dolphin followed sound stimuli up to 1,000 Hz with a second peak response at 500 Hz. A weighted MRTF reflected that the animal followed a broad range of rates from 100 to 1,000 Hz, but beyond 1,250 Hz the animal's hearing response was simply an onset/offset response. Similar to other mammals, the dolphin's AEP response to a single stimulus was a series of waves. The delay of the first wave, PI, was 2.76 ms and the duration of the multi-peaked response was 4.13 ms. The MRTF was similar in shape to other marine mammals except that the response delay was among the fastest measured. Results predicted that the Risso's dolphin should have the ability to follow clicks and echoes while foraging at close range.

Animal Communication↗
PubMed2005

Serotonin modulates responses to species-specific vocalizations in the inferior colliculus.

Neuromodulators such as serotonin are capable of altering the neural processing of stimuli across many sensory modalities. In the inferior colliculus, a major midbrain auditory gateway, serotonin alters the way that individual neurons respond to simple tone bursts and linear frequency modulated sweeps. The effects of serotonin are complex, and vary among neurons. How serotonin transforms the responses to spectrotemporally complex sounds of the type normally heard in natural settings has been poorly examined. To explore this issue further, the effects of iontophoretically applied serotonin on the responses of individual inferior colliculus neurons to a variety of recorded species-specific vocalizations were examined. These experiments were performed in the Mexican free-tailed bat, a species that uses a rich repertoire of vocalizations for the purposes of communication as well as echolocation. Serotonin frequently changed the number of recorded calls that were capable of evoking a response from individual neurons, sometimes increasing (15% of serotonin-responsive neurons), but usually decreasing (62% of serotonin-responsive neurons), this number. A functional consequence of these serotonin-evoked changes would be to change the population response to species-specific vocalizations.

Acoustic Stimulation↗