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ON-OFF units in the mustached bat inferior colliculus are selective for transients resembling "acoustic glint" from fluttering insect targets.

Of 311 single units studied in the central nucleus of the inferior colliculus (ICC) in 18 mustached bats (Pteronotus parnelli), a small but significant population (13%) of cells with on-off discharge patterns to tone bursts at best frequency (BF) was found in the dorsoposterior division. In contrast to units with the same BF's but other discharge patterns, the majority of ON-OFF units were unresponsive to sinusoidally amplitude-modulated tone bursts (SAM). To define the contribution of linear and nonlinear components to the responses of ICC neurons to amplitude modulation, we tested some of these neurons with a long, seamlessly repeating pseudorandom sequence of ternary amplitude-modulated tones at BF. Wiener-like kernels were subsequently derived from cross-correlation of spikes with acoustic events in the sequence. These kernels provided estimates of neural impulse responses that proved unusual in SAM-unresponsive ON-OFF units. First, their estimated impulse response had no linear component. Second, the predicted second-order impulse responses to both increments and decrements in stimulus intensity were long (about 20 ms) and nearly identical in shape: triphasic, with the positive phase bounded by leading and trailing negative periods. The similar shape of responses to increments and decrements in these neurons suggests a full-wave rectifier. The triphasic, initially negative second-order prediction of the impulse response accounted for an unusual result in experiments measuring the recovery cycle of ON-OFF units using a pair of identical stimulus pulses separated by various time delays. This recovery cycle can be related to their response to amplitude modulation. As the delay between two brief, near-threshold BF tone bursts decreased, the response to the first tone diminished, rather than to the second. The second-order prediction of this experiment derived from impulse responses obtained with pseudorandom noise suggests that, at short interpulse intervals, the initial negative phase of the response to the later stimulus cancels the positive phase of the response to the first. Such cancellation at short interpulse intervals may help explain why the majority of ON-OFF units are unresponsive to SAM. The unusual properties of these ON-OFF units make them ideally suited to respond selectively to infrequent acoustic transients superimposed on an ongoing background of modulation. Such patterns are commonly encountered by mustached bats foraging in cluttered habitats for small, fluttering insects, which generate "acoustic glints" upon a background of modulated echoes from the surroundings (Schnitzler et al. 1983; Henson et al. 1987).

Acoustic Stimulation↗

Coding of small sinusoidal frequency and amplitude modulations in the inferior colliculus of 'CF-FM' bat, Rhinolophus ferrumequinum.

Single neurons in the inferior colliculus of the Greater Horseshoe bat, Rhinolophus ferrumequinum, showed two broad categories of response patterns to sinusoidally frequency (SFM) or amplitude (SAM) modulated stimuli. Tonic responding cells (best excitatory frequency (BEF) between 10 and 90 kHz) showed a rough sinusoidal modulation of the discharge pattern to SFM. Transient responding neurons generally showing on- or off-responses to pure tones, (BEF between 65 and 88 kHz), displayed highly synchronized discharge patterns to SFM-cycles (Fig. 1). Modulation rates between 20 and 100 Hz were most effective and some neurons encoded modulation rates up to 350 Hz (Figs. 2 and 3). The SFM responses were best synchronized to the modulation envelope for center frequencies in the upper portion of the tuning curve (Figs. 4 and 5). Sharply tuned neurons with BEF around 80 kHz had the lowest threshold for modulation depth (+/- 10 Hz or 0.025%) (Fig. 6). In general, SAMs evoked the same type of response patterns and were encoded down to modulation index of 3% (Fig. 7). The fine frequency and amplitude discriminations for periodical modulations by collicular neurons is discussed as related to the detection and discrimination performance of bats, when preying on flying insects in clustered surroundings.

Acoustic Stimulation↗

Phase evaluation in hypothetical receivers simulating ranging in bats.

Echo delay discrimination by the bat Eptesicus fuscus had been investigated in an experiment with simulated targets jittering in range (Simmons 1979). The dip in the resulting psychometric curve was used by Simmons to suggest the neuronal implementation of a coherent cross-correlation receiver in the auditory system of bats. By computer simulation it is shown here that the dip may be even more pronounced and less susceptible to noise with alternative receiver configurations which not necessarily evaluate signal phase information coherently, e.g., a bank of neuronal filters with envelope-processing. New behavioral experiments are suggested to critically test such model hypotheses.

Acoustic Stimulation↗

Retinofugal projections in the rufous horseshoe bat, Rhinolophus rouxi.

The retinal projections in the horseshoe bat were studied with anterograde transport of wheat germ agglutinin conjugated to horseradish peroxidase. Retinal fibers clearly terminate bilaterally in the lateral geniculate nuclei, superior colliculus, pretectal area, and nucleus of the optic tract. The suprachiasmatic nucleus and the lateral terminal nucleus of the accessory optic tract receive extremely weak, through bilateral retinal input. No projections to medial and dorsal accessory optic nuclei were found. There was a limited retinal projection to the ipsilateral dorsal geniculate nucleus. The focus of the ipsilateral projection corresponded to a less densely labeled region on the contralateral side. In this study an ipsilateral retinal projection to the anterior superior colliculus is documented for the first time in a Microchiropteran bat. In the contralateral superior colliculus retinal fibers terminate in a patch-like pattern at caudal levels.

Animals↗

An immunohistochemical study on the innervation of SP-IR nerves in the cerebral arteries of the bent-winged bat.

The overall distribution of substance P (SP) immunoreactive (IR) nerves surrounding the cerebral arteries of the bent-winged bat were investigated immunohistochemically. In this microchiropteran species, the walls from the vertebral artery to the caudal part of the basilar artery have considerably well-developed plexuses of SP-IR nerves, whereas no demonstrable SP-IR fibers were found in the rostral part of the basilar artery, and in more rostrally located arteries the nerve supply was very sparse or occasionally lacking. This innervation pattern has not yet been established for the cerebral arterial systems of other mammals that have been studied under normal conditions, but it is very similar to the pattern of SP-IR innervation observed in the guinea pig and cat of which the trigeminal ganglia have been destroyed. From the combination of this and other immunohistochemical findings, it is suggested that SP-IR nerves innervating the vertebral and basilar arteries of the bent-winged bat originate from the upper cervical dorsal root ganglia (DRG) and enter the cranial cavity along the vertebral artery and through the meninges.

Animals↗

Serological analysis and pathogenic potentials of Wangiella dermatitidis isolated from bats.

Five isolates of Wangiella dermatitidis from macroscopically asymptomatic bats were analysed by the agar-gel immunodiffusion tests and the mouse inoculation studies on virulence. No cross-reactivity among the antigens and antisera prepared from the bat and human isolates was observed. The fungus was shown to be neurotropic, of low infectivity and not implicated as a potential opportunistic pathogen. Differences in the antigenic composition and pathogenic potentials between isolates of the same fungus of different origins may reflect adaptation of the fungus in different hosts.

Animals↗

The influence of the social thermoregulation on the cold-adaptive growth of BAT in hairless and furred mice.

When mice were living in groups they developed less brown adipose tissue (BAT) during cold adaptation as compared with single mice. This effect of social aggregation was more pronounced in genetically hairless mice than in furred mice. In both races of mice the most significant difference in BAT growth was found between single mice and pairs of mice, indicating that the formation of pairs causes the relatively most effective improvement of thermal balance.

Adaptation, Physiological↗

Coding of sinusoidally amplitude modulated acoustic stimuli in the inferior colliculus of the rufous horseshoe bat, Rhinolophus rouxi.

Single neuron responses to sinusoidally amplitude modulated (SAM) signals were studied in the inferior colliculus of the horseshoe bat, Rhinolophus rouxi. 57% of the neurons responded to SAM stimuli with periodical discharges synchronized to the modulation cycle. The proportion of cells driven by amplitude modulated signals was independent of the best frequency of the neurons. Best modulation frequencies were at or below 100 Hz in about 70% of the neurons. Synchronized activity could be elicited by modulation frequencies up to 400 Hz. Best SAM responses were observed at stimulus intensities 10 dB above threshold. Generally the BMF of a neuron did not change with intensity. The BMF decreased with decreasing modulation depth of the amplitude modulation. A trend for a topographical organization of neurons according to best modulation frequencies was detected. The results did not reveal any significant specialization of the bat's auditory system for coding of amplitude modulations as compared to other mammals.

Animals↗

Isolation of Lecythophora mutabilis and Wangiella dermatitidis from the fruit eating bat, Eidolon helvum.

Lecythophora mutabilis was isolated from the lungs of 3 and from the liver of 2 bats, Eidolon helvum a fruit eating species. Wangiella dermatitidis was recovered from the liver of 2 bats of the same species. The isolates were pathogenic for laboratory mice when injected by subcutaneous, intraperitoneal and intravenous routes. W. dermatitidis was neurotropic in the mice injected intravenously.

Animals↗

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↗

A comparative study of the physiological properties of the inner ear in Doppler shift compensating bats (Rhinolophus rouxi and Pteronotus parnellii).

Cochlear microphonic (CM) and evoked neural (N-1) potentials were studied in two species of Doppler shift compensating bats with the aid of electrodes chronically implanted in the scala tympani. Potentials were recorded from animals fully recovered from the effects of anesthesia and surgery. In Pteronotus p. parnellii and Rhinolophus rouxi the CM amplitude showed a narrow band, high amplitude peak at a frequency about 200 Hz above the resting frequency of each species. In Pteronotus the peak was 25-35 dB higher in amplitude than the general CM level below or above the frequency of the amplitude peak. In Rhinolophus the amplitude peak was only a few dB above the general CM level but it was prominent because of a sharp null in a narrow band of frequencies just below the peak. The amplitude peak and the null were markedly affected by body temperature and anesthesia. In Pteronotus high amplitude CM potentials were produced by resonance, and stimulated cochlear emissions were prominent in Pteronotus but they were not observed in Rhinolophus. In Pteronotus the resonance was indicated by a CM afterpotential that occurred after brief tone pulses. The resonance was not affected by the addition of a terminal FM to the stimulus and when the ear was stimulated with broadband noise it resulted in a continual state of resonance. Rapid, 180 degree phase shifts in the CM were observed when the stimulus frequency swept through the frequency of the CM amplitude peak in Pteronotus and the frequency of the CM null in Rhinolophus. These data indicate marked differences in the physiological properties of the cochlea and in the mechanisms responsible for sharp tuning in these two species of bats.

Anesthesia, General↗

The cochlear frequency map of the mustache bat, Pteronotus parnellii.

The frequency-place map of the cochlea of mustache bats was constructed by the analysis of HRP-transport patterns in spiral ganglion cells following iontophoretic tracer injections into cochlear nucleus regions responsive to different frequencies. The cochlea consists of 5 half turns (total length 14.3 mm) and the representation of certain frequency bands can be assigned to specific cochlear regions: The broad high frequency range between 70 and 111 kHz is represented in the most basal half turn within only 3.2 mm. This region is terminated apically by a distinct narrowing of the scala vestibuli that coincides with a pronounced increase in basilar membrane (BM) thickness. The narrow intermediate frequency range between 54 and 70 kHz is expanded onto 50% of cochlear length between 4.0 and 11.1 mm distance from apex. The frequency range around 60 kHz, where the tuning characteristics of the auditory system are exceptionally sharp, is located in the center of this expanded BM-region in the second half turn within a maximum of innervation density. These data can account for the vast overrepresentation of neurons sharply tuned to about 60 kHz at central stations of the auditory pathway. In the cochlear region just basal to the innervation maximum, where label from injections at 66 and 70 kHz was found, a number of morphological specializations coincide: the BM is maximally thickened, innervation density is low, the spiral ligament is locally enlarged, and the 'thick lining', a dense covering of the scala tympani throughout the basal halfturn, suddenly disappears. Low frequencies up to 54 kHz are represented within the apical half turns over a 4 mm span of the basilar membrane. The data are compared to the cochlea of horseshoe bats and the possible functional role of the morphological discontinuities for sharp tuning and the generation of otoacoustic emissions is discussed.

Animals↗

Ascorbic acid content of neotropical plant parts available to wild monkeys and bats.

The ascorbic acid content of foliage available to wild primates and bats in Panama (in transition between wet and dry seasons) was lower than that of temperate zone foliage but higher than that of most fruits and vegetables. Intakes of ascorbic acid (mg/kg b.wt/day) by wild primates and frugivorous bats in Panama are much greater than that of most human populations.

Animal Nutritional Physiological Phenomena↗

Effect of light adaptation on electrical responses of the retinas of four species of bats.

The levels of light adaptation at which the retinas of 4 species of microchiropteran bats became unable to generate electroretinograms were progressively ordered. The order correlated well with light preferences based on activity patterns of the 4 species. These results suggest that the ability of the retina to function in ambient light may govern some natural behaviors of these bats.

Adaptation, Ocular↗

GABAergic and glycinergic neural inhibition in excitatory frequency tuning of bat inferior collicular neurons.

This study examined the effect of GABAergic and glycinergic inhibition on excitatory frequency tuning curves (FTCs) of inferior collicular (IC) neurons of the big brown bat, Eptesicus fuscus. The excitatory FTCs of 70 IC neurons were either V-shaped (57, 81%), closed (11, 16%), or double-peaked (2, 3%). By means of a two-tone stimulation paradigm, inhibitory FTCs were obtained at one frequency flank only (low-frequency flank: 11, 16%; high-frequency flank: 7, 10%), at both frequency flanks (36, 51%) of excitatory FTCs, or between two excitatory FTCs (2, 3%). IC neurons that had inhibitory FTCs typically had larger Q(10) and Q(30) values (i.e., sharper excitatory FTCs) than neurons that did not have inhibitory FTCs. Neurons with inhibitory FTCs at both frequency flanks had larger Q(10) and Q(30) values than neurons with inhibitory FTCs at one frequency flank only. IC neurons with a small difference between excitatory and inhibitory best frequencies typically had sharper excitatory frequency tuning. Bicuculline (an antagonist for GABAA) application produced a greater degree of abolishing inhibitory FTCs than strychnine (an antagonist for glycine) application. Application of both drugs was most effective in abolishing the inhibitory FTCs of IC neurons. The implications of these findings for bat echolocation are discussed.

Acoustic Stimulation↗

Maximum likelihood analysis of the complete mitochondrial genomes of eutherians and a reevaluation of the phylogeny of bats and insectivores.

The complete mitochondrial genomes of two microbats, the horseshoe bat Rhinolophus pumilus, and the Japanese pipistrelle Pipistrellus abramus, and that of an insectivore, the long-clawed shrew Sorex unguiculatus, were sequenced and analyzed phylogenetically by a maximum likelihood method in an effort to enhance our understanding of mammalian evolution. Our analysis suggested that (1) a sister relationship exists between moles and shrews, which form an eulipotyphlan clade; (2) chiropterans have a sister-relationship with eulipotyphlans; and (3) the Eulipotyphla/Chiroptera clade is closely related to fereuungulates (Cetartiodactyla, Perissodactyla and Carnivora). Divergence times on the mammalian tree were estimated from consideration of a relaxed molecular clock, the amino acid sequences of 12 concatenated mitochondrial proteins and multiple reference criteria. Moles and shrews were estimated to have diverged approximately 48 MyrBP, and bats and eulipotyphlans to have diverged 68 MyrBP. Recent phylogenetic controversy over the polyphyly of microbats, the monophyly of rodents, and the position of hedgehogs is also examined.

Animals↗

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↗