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PubMed1997

Ontogeny of vocal signals in the little brown bat, Myotis lucifugus

This study reports developmental changes in the vocal signals produced by wild-caught pre-volant and volant Myotis lucifugusAudio recordings were made from young animals (1-33 days old) and adults (over 1 year of age). The animals were removed from an attic maternity roost and studied individually in a room below. To stimulate flight-related behaviours, animals were released from a launching platform via a hinged floor, and their vocalizations were recorded as they approached a soft foam pad below. When the hinged floor opened, the youngest animals (1-4 days old) typically dropped onto the pad, but older animals either flapped their wings to achieve some horizontal displacement (>4 days) or sustained horizontal flight (>17 days). Vocalizations recorded under these conditions showed frequency modulation characteristic of adult echolocation sounds, even in animals as young as 4 days. Ontogenetic trends showed an increase in sound frequency, an increase in sound repetition rate and a decrease in sound duration as the animals matured. These data are discussed in the context of the development of echolocation behaviour in bats.

Journal Article↗
PubMed1991

Parallel processing of afferent input by identified interneurones in the auditory pathway of the noctuid moth Noctua pronuba (L.).

1. Interneurones 501 and 504 are identified sound-sensitive interneurones in the pterothoracic ganglion of the noctuid moth Noctua pronuba. Both neurones receive monosynaptic input from the A1 afferent and experiments with current injection suggest that the synapse is chemical. The EPSPs evoked in either IN 501 or 504 by the A1 afferent do not facilitate. 2. Temporal integration in INs 501 and 504 was compared by presenting the moth with tones at repetition rates found in the search, approach and terminal phases of the echolocating call of a hunting bat. INs 501 and 504 differ in their capacity to resolve stimulus repetition rates because the mean decay times of their compound EPSPs differ by a factor of three, although both interneurones receive monosynaptic input from the A1 afferent. 3. The features extracted from the authentic, prerecorded, call of an echolocating bat at the level of the pterothoracic ganglion were examined by recording sequentially from a range of interneurones in the same preparation. The capacity of INs 501 and 504 to encode the various phases of the call was examined in the light of their measured mean decay times and related to the avoidance behaviour of the insect.

Acoustic Stimulation↗
PubMed1987

The nuclei of the lateral lemniscus in the rufous horseshoe bat, Rhinolophus rouxi. A neurophysiological approach.

In the rufous horseshoe bat, Rhinolophus rouxi, responses to pure tones and sinusoidally frequency modulated (SFM) signals were recorded from 289 single units and 241 multiunit clusters located in the nuclei of the lateral lemniscus (NLL). The distribution of best frequencies (BFs) of units in all three nuclei of the lateral lemniscus showed an overrepresentation in the range corresponding to the constant-frequency (CF) part of the echolocation signal ('filter frequency' range): in the ventral nucleus of the lateral lemniscus (VNLL) 'filter neurons' represented 43% of all units encountered, in the intermediate nucleus (INLL) 33%, and in dorsal nucleus (DNLL) 29% (Fig. 2a). Neurons with best frequencies in the filter frequency range had highest Q10dB-values (maxima up to 400, Fig. 2c) and only in low-frequency units were values comparable to those found in other mammals. On the average, filter neurons in ventral nucleus had higher Q10dB-values (about 220) than did those in intermediate and dorsal nucleus (both about 160, Fig 2d). Response patterns and tuning properties showed higher complexity in the dorsal and intermediate nucleus than in the ventral nucleus of the lateral lemniscus (Figs. 4 and 6). Multiple best frequencies were found in 12 neurons, nine of them with harmonically related excitation maxima (Fig. 5c, d). Best frequencies of six of these harmonically tuned units could not be correlated with any harmonic components of the echolocation signal. Half of all multiple tuned neurons were located in the caudal dorsal nucleus the other half in the caudal intermediate nucleus. Synchronization of responses to sinusoidally frequency modulated (SFM) signals occurred in VNLL-units in the average up to modulation frequencies of 515 Hz (maximum about 800 Hz) whereas in the intermediate and dorsal nucleus of the lateral lemniscus responses were synchronized in the average only up to modulation frequencies of about 300 Hz (maximum about 600 Hz) (Figs. 7 and 8). A tonotopic arrangement of units was found in the intermediate nucleus of the lateral lemniscus with units having high best frequencies located medially and those with low best frequencies laterally. In the dorsal nucleus the tonotopic distribution was found to be fairly similar to that in the intermediate nucleus but much less pronounced. In more rostral parts of the dorsal nucleus additionally higher best frequencies predominated whereas in caudal areas of that nucleus and also of the intermediate nucleus low BFs were found more regularly.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1985

An HRP-study of the frequency-place map of the horseshoe bat cochlea: morphological correlates of the sharp tuning to a narrow frequency band.

The frequency-place map of the horseshoe bat cochlea was studied with the horseradish peroxidase (HRP) technique involving focal injections into various, physiologically defined regions of cochlear nucleus (CN). The locations of labeled spiral ganglion cells and their termination sites on inner hair cells of the organ of Corti from injections into CN-regions responsive to different frequencies were analyzed in three dimensional reconstructions of the cochlea. Horseshoe bats from different geographical populations were investigated. They emit orientation cells with constant frequency (CF) components around 77 kHz (Rhinolophus rouxi from Ceylon) and 84 kHz (Rhinolophus rouxi from India) and their auditory systems are sharply tuned to the respective CF-components. The HRP-map shows that in both populations: the frequency range around the CF-component of the echolocation signal is processed in the second half-turn of the cochlea, where basilar membrane (BM) is not thickened, secondary spiral lamina (LSS) is still present and innervation density is maximal; frequencies more than 5 kHz above the CF-component are processed in the first half-turn, where the thickened BM is accompanied by LSS and innervation density is low; frequencies below the spectral content of the orientation call are represented in apical turns showing no morphological specializations. The data demonstrate that the cochlea of horseshoe bats is normalized to the frequency of the individual specific CF-component of the echolocation call. The HRP-map can account for the overrepresentation of neurons sharply tuned to the CF-signal found in the central auditory system. A comparison of the HRP-map with a map derived with the 'swollen nuclei technique' following loud sound exposure (Bruns 1976b) reveals that the latter is shifted towards cochlear base by about 4 mm. This discrepancy warrants a new interpretation of the functional role of specialized morphological structures of the cochlea within the mechanisms giving rise to the exceptionally high frequency selectivity of the auditory system.

Animals↗
PubMed1975

Lipids of freshwater Dolphin Sotalia fluviatilis: comparison of odontocete bioacoustic lipids and habitat.

The melon and jaw lipids of the freshwater dolphin Sotalia fluviatilis are composed mainly of isovaleroyl wax esters and diisovaleroyl triglycerides. The blubber fat contains only a trace of wax ester and is mostly tri-(long chain) and monoisovaleroyl triglycerides. Detailed gas liquid chromatographic analyses of the intact wax esters and triglycerides and of the derived fatty acids and fatty alcohols indicate common compositional patterns in the wax esters and triglycerides of the respective head lipids. Both odd and even long chain (C12-C16) isostructures are prominent in the melon and jaw lipids, but only higher odd chain length iso-acids are major components in the blubber. Sotalia fluviatilis (family Delphinidae) and Inia geoffrensis (family Platanistidae) share the same freshwater habitat in the upper Amazon River, and both utilize echolocation to navigate and to find food. Comparison of their respective bioacoustical lipid compositions show distinctive types of head fats, Sotalia being rich in iso-5:0 and Inia lacking iso-5:0. This indicates that iso-valeric acid per se has no obligatory role in dolphin echolocation.

Acoustics↗
PubMedDate not supplied

The receptor and neuron distribution in the cochlea of the bat, Taphozous kachhensis.

Cochlear receptors and spiral ganglion neurons in the inner ear of the bat, Taphozous kachhensis of the family Emballonuridae, were quantitatively analyzed on the basis of surface specimen technique, histological sections, and three-dimensional computer reconstruction. The length of the basilar membrane was 12.1 (+/- 0.4) mm. There were about 23,000 neurons in the spiral ganglion and about 1430 inner hair cells and 4635 outer hair cells in the organ of Corti. Cochlear neurons had two density maxima, one basal and one apical. Inner hair cells had basically the same distribution but less pronounced. Outer hair cells had only one density maximum in the apical region corresponding to the respective maxima of inner hair cells and cochlear neurons. Basal density maxima of inner hair cells and neurons are most probably involved in the analysis of echolocation signals, and presumably correlated with maximum sensitivity of the auditory system. A comparison with the bat, Nyctalus noctula of the family Vespertilionidae revealed a conspicuous similarity in some quantitative aspects of the cochlear structure. These particular similarities may be looked upon as convergent adaptations given by similar flight, hunting, and echolocation strategies in both bat species.

Animals↗
PubMed2003

A new phylogeny of swiftlets (Aves: Apodidae) based on cytochrome-b DNA.

Due to a lack of distinctive morphological characters, swift taxonomy and phylogeny has always been an area of disagreement. To shed more light on this subject, we reconstructed swift(let) phylogeny based on 1143 bp of mitochondrial cytochrome-b DNA sequence. Although this is not the first attempt to reconstruct swift phylogeny using molecular data, our results show higher support for many of the branches due to our much longer sequences. However, placement of Hydrochous is still unexpected. Implementation of more conservative genetic regions and sampling of more taxa could solve this problem. Most importantly, the Collocaliini resolve as a monophyletic group. The internal structure of the group shows that non-echolocating Collocalia and echolocating Aerodramus form two distinct clades. This is in congruence with earlier classifications based on morphological characters, but in contrast with more recent classifications.

Animals↗
PubMed2000

Sequence sensitivity of neurons in cat primary auditory cortex.

Properties of sequence-sensitive neurons in primary auditory cortex of cats were explored in detail. Stimuli were sequences of two tones, in which the frequency and intensity of the first tone and the temporal separation between the first and second, or probe, tone were parametrically varied. After presentation of the first tone, the responses of 32 single units and 48 multiunits to the probe tone were found to be enhanced up to 140-5270% (median 340%) above the response obtained in the single-tone condition. Probe tone enhancement was induced from a considerable number of sequence conditions and depended on the frequency and intensity of the first tone and on the temporal separation between the onsets of the first and the probe tone. On average, the maximally enhanced response occurred when the first tone was 1 octave below or above the probe tone and its intensity was 14 dB louder than the probe tone. The most effective temporal separation of the tones for an enhancement effect was approximately 100 ms. The range of enhancing tones was largely outside the excitatory tuning curve of a neuron. Results extend previous findings of properties of sequence-sensitive neurons in the auditory cortex of echolocating bats and non-echolocating mammals, and suggest that sequence-sensitive neurons are quite common and involved in the cortical representation of spectrotemporal patterns of acoustic signals.

Acoustic Stimulation↗
PubMed2004

Extinction of the acoustic startle response in moths endemic to a bat-free habitat.

Most moths use ears solely to detect the echolocation calls of hunting, insectivorous bats and evoke evasive flight manoeuvres. This singularity of purpose predicts that this sensoribehavioural network will regress if the selective force that originally maintained it is removed. We tested this with noctuid moths from the islands of Tahiti and Moorea, sites where bats have never existed and where an earlier study demonstrated that the ears of endemic species resemble those of adventives although partially reduced in sensitivity. To determine if these moths still express the anti-bat defensive behaviour of acoustic startle response (ASR) we compared the nocturnal flight times of six endemic to six adventive species in the presence and absence of artificial bat echolocation sounds. Whereas all of the adventive species reduced their flight times when exposed to ultrasound, only one of the six endemic species did so. These differences were significant when tested using a phylogenetically based pairwise comparison and when comparing effect sizes. We conclude that the absence of bats in this habitat has caused the neural circuitry that normally controls the ASR behaviour in bat-exposed moths to become decoupled from the functionally vestigial ears of endemic Tahitian moths.

Animals↗
PubMed2003

Postnatal development of cochlear function in the mustached bat, Pteronotus parnellii.

Postnatal development of the mustached bat's cochlea was studied by measuring cochlear microphonic and compound action potentials. In adults, a cochlear resonance is involved in enhanced tuning to the second harmonic constant frequency component (CF2) of their echolocation calls at approximately 61 kHz This resonance is present immediately after birth in bats that do not yet echolocate. Its frequency is lower (46 kHz) and the corresponding threshold minimum of cochlear microphonic potentials is broader than in adults. Long-lasting ringing of the cochlear microphonic potential after tone stimulus offset that characterizes the adult auditory response close to CF2 is absent in newborns. In the course of the first 5 postnatal weeks, there is a concomitant upward shift of CF2 and the frequency of cochlear threshold minima. Up to the end of the third postnatal week, sensitivity of auditory threshold minima and the Q value of the cochlear resonance increase at a fast rate. Between 2 and 4 wk of age, two cochlear microphonic threshold minima are found consistently in the CF2 range that differ in their level-dependent dynamic growth behavior and are 1.5-5.7 kHz apart from each other. In older animals, there is a single minimum that approaches adult tuning in its sharpness. The data provide evidence to show that during maturation of the cochlea, the frequency and the sensitivity of the threshold minimum associated with CF2 increases and that these increases are associated with the fusion of two resonances that are partly dissociated in developing animals.

Acoustic Stimulation↗
PubMed2006

Behavioral flexibility positively correlated with relative brain volume in predatory bats.

We investigated the potential relationships between foraging strategies and relative brain and brain region volumes in predatory (animal-eating) echolocating bats. The species we considered represent the ancestral state for the order and approximately 70% of living bat species. The two dominant foraging strategies used by echolocating predatory bats are substrate-gleaning (taking prey from surfaces) and aerial hawking (taking airborne prey). We used species-specific behavioral, morphological, and ecological data to classify each of 59 predatory species as one of the following: (1) ground gleaning, (2) behaviorally flexible (i.e., known to both glean and hawk prey), (3) clutter tolerant aerial hawking, or (4) open-space aerial hawking. In analyses using both species level data and phylogenetically independent contrasts, relative brain size was larger in behaviorally flexible species. Further, relative neocortex volume was significantly reduced in bats that aerially hawk prey primarily in open spaces. Conversely, our foraging behavior index did not account for variability in hippocampus and inferior colliculus volume and we discuss these results in the context of past research.

Adaptation, Psychological↗
PubMed1988

Peak density and distribution of ganglion cells in the retinae of microchiropteran bats: implications for visual acuity.

We have estimated the total number, distribution and peak density of retinal ganglion cells (RGCs) in retinal wholemounts of several species of microchiropteran (echolocating) bats. The estimates are based on counts of Nissl-stained, presumed RGCs. The total number of presumed RGCs varies among the species: from about 4,500 in Rhinolophus rouxi to about 120,000 in Macroderma gigas. In addition, in two species (Nyctophilus gouldi and M. gigas), the estimates are based on counts of positively identified RGCs retrogradely labelled with the enzyme horseradish peroxidase injected into the retinorecipient nuclei. In these two species, the numbers and distributions of retrogradely labelled RGCs and Nissl-stained presumed RGCs are very similar. In all six species studied, the peak-density regions of presumed (or positively identified) RGCs are located in the inferotemporal retinae, and the RGC isodensity lines tend to be horizontally elongated. However, the RGC densities in the high-density regions are only 2-4 times greater than those in the low-density regions in the superior retinae. The somal sizes of RGCs vary from 5 to 16 micron in diameter and are unimodally distributed. There is no indication of the existence of distinct morphological classes of RGCs. The axial lengths of microchiropteran eyes vary from 1.8 mm in R. rouxi to 7.0 mm in M. gigas. For all species the posterior nodal distance (PND) was assumed to be 0.52 of the axial length of the eye. This assumption is based on the analysis of published data concerning schematic eyes of nocturnal vertebrates. These derived values of the PNDs allowed us to calculate the retinal magnification factors and the number of RGCs per degree of visual angle. From these, the upper limits of visual acuity were derived on the basis of the assumptions of the sampling theorem. The estimated upper limits of visual acuity of the six species of echolocating bats vary from about 0.35 cycles/degree in R. rouxi to about 2 cycles/degree in M. gigas. This range is quite similar to the range of visual acuities in murid rodents.

Animals↗
PubMed2005

Evidence for the use of reflected self-generated seismic waves for spatial orientation in a blind subterranean mammal.

Subterranean mammals like the blind mole-rat (Rodentia: Spalax ehrenbergi) are functionally blind and possess poor auditory sensitivity, limited to low-frequency sounds. Nevertheless, the mole-rat demonstrates extremely efficient ability to orient spatially. A previous field study has revealed that the mole-rat can assess the location, size and density of an underground obstacle, and accordingly excavates the most efficient bypass tunnel to detour around the obstacles. In the present study we used a multidisciplinary approach to examine the possibility that the mole-rat estimates the location and physical properties of underground obstacles using reflected self-generated seismic waves (seismic 'echolocation'). Our field observations revealed that all the monitored mole-rats produced low-frequency seismic waves (250-300 Hz) at intervals of 8+/-5 s (range: 1-13 s) between head drums while digging a bypass to detour an obstacle. Using a computerized simulation model we demonstrated that it is possible for the mole-rat to determine its distance from an obstacle boundary (open ditch or stone) by evaluating the amplitude (intensity) of the seismic wave reflected back to it from the obstacle interface. By evaluating the polarity of the reflected wave the mole-rat could distinguish between air space and solid obstacles. Further, the model showed that the diffracted waves from the obstacle's corners could give the mole-rat precise information on the obstacle size and its relative spatial position. In a behavioural experiment using a special T-maze setup, we tested whether the mole-rat can perceive seismic waves through the somatosensory system and localize the source. The results revealed that the mole-rat is able to detect low frequency seismic waves using only its paws, and in most cases the mole-rats determined accurately the direction of the vibratory source. In a histological examination of the glabrous skin of the mole-rat's paws we identified lamellate corpuscle mechanoreceptors that might be used to detect low frequency seismic waves. The combined findings from these different approaches lead us to suggest that a specialized seismic 'echolocation' system could be used by subterranean mammals to determine the most energy-conserving strategy with which to bypass an obstacle, as well as to estimate their distance from the surface, keeping their tunnels at the optimal depth.

Adaptation, Physiological↗
PubMed2006

Steering by hearing: a bat's acoustic gaze is linked to its flight motor output by a delayed, adaptive linear law.

Adaptive behaviors require sensorimotor computations that convert information represented initially in sensory coordinates to commands for action in motor coordinates. Fundamental to these computations is the relationship between the region of the environment sensed by the animal (gaze) and the animal's locomotor plan. Studies of visually guided animals have revealed an anticipatory relationship between gaze direction and the locomotor plan during target-directed locomotion. Here, we study an acoustically guided animal, an echolocating bat, and relate acoustic gaze (direction of the sonar beam) to flight planning as the bat searches for and intercepts insect prey. We show differences in the relationship between gaze and locomotion as the bat progresses through different phases of insect pursuit. We define acoustic gaze angle, theta(gaze), to be the angle between the sonar beam axis and the bat's flight path. We show that there is a strong linear linkage between acoustic gaze angle at time t [theta(gaze)(t)] and flight turn rate at time t + tau into the future [theta(flight) (t + tau)], which can be expressed by the formula theta(flight) (t + tau) = ktheta(gaze)(t). The gain, k, of this linkage depends on the bat's behavioral state, which is indexed by its sonar pulse rate. For high pulse rates, associated with insect attacking behavior, k is twice as high compared with low pulse rates, associated with searching behavior. We suggest that this adjustable linkage between acoustic gaze and motor output in a flying echolocating bat simplifies the transformation of auditory information to flight motor commands.

Animals↗
PubMed2001

Iontophoresis in vivo demonstrates a key role for GABA(A) and glycinergic inhibition in shaping frequency response areas in the inferior colliculus of guinea pig.

The processing of biologically important sounds depends on the analysis of their frequency content by the cochlea and the CNS. GABAergic inhibition in the inferior colliculus shapes frequency response areas in echolocating bats, but a similar role in nonspecialized mammals has been questioned. We used the powerful combination of iontophoresis with detailed analysis of frequency response areas to test the hypothesis that GABAergic and glycinergic inhibition operating in the inferior colliculus of a nonspecialized mammal (guinea pig) shape the frequency responses of neurons in this nucleus. Our analysis reveals two groups of response areas in the inferior colliculus: V-shaped and non-V-shaped. The response as a function of level in neurons with V-shaped response areas can be either monotonic or nonmonotonic. Application of bicuculline or strychnine in these neurons, to block inhibition mediated by GABA(A) or glycinergic receptors, respectively, increases firing rate primarily within the boundaries of the control response area. In contrast, neurons in the non-V-shaped group have response areas that include narrow, closed, tilted, and double-peaked types. In this group, blockade of GABA(A) and glycine receptors increases firing rate but also changes response area shape, with most becoming more V-shaped. We conclude that (1) non-V-shaped response areas can be generated by GABA and glycinergic synapses within the inferior colliculus and do not simply reflect inhibition acting more peripherally in the pathway and (2) frequency-dependent inhibition is an important general feature of the mammalian inferior colliculus and not a specialization unique to echolocating bats.

Acoustic Stimulation↗
PubMed1985

[Diagnosis of aortic valve calcinosis with a view to the surgical treatment of aortic valve disease].

Combined roentgenologic/echocardiographic diagnosis of aortic valve calcinosis was made in 135 patients operated on for a rheumatic aortic heart disease. Four degrees of aortic valve calcinosis were identified: 1) isolated fine points of calcinosis, as revealed by echolocation and specific treatment of the removed valves; 2) small-focal calcinosis as revealed roentgenologically in 79.2% of cases and by echolocation, in 93%; 3) large-focal, and 4) wide-spread calcinosis, the latter two being detectable both roentgenologically and echocardiographically in all cases. The 4-degree classification of aortic valvular lesions allows a detailed assessment of the nature of the affection that is important for the choice of surgical procedure.

Aortic Valve↗
PubMed2000

An extralemniscal component of the mustached bat inferior colliculus selective for direction and rate of linear frequency modulations.

Frequency modulations (FMs) are prevalent in human speech, and are important acoustic cues for the categorical discrimination of phonetic contrasts. For bats, FM sweeps are also important for communication and are often the only component in echolocation calls. Auditory neurons tuned to the direction and rate of FM might underlie the encoding of rapid frequency transitions. In the mustached bat, we have discovered a population of such FM selective cells in an area interposed between the central nucleus of the inferior colliculus (ICC) and the nuclei of the lateral lemniscus (NLL). We believe this area to be the ventral extent of the external nucleus of the inferior colliculus (ICXv). To describe FM selectivity of neurons in the ICXv and to compare it to other midbrain nuclei, up- and down-sweeping linear FM stimuli were presented at different modulation rates. Extracellular recordings were made from 171 single units in the ICC, ICXv, and NLL of 10 mustached bats. In the ICXv, there was a much higher degree of FM selectivity than in ICC or NLL and a consistent preference for upward over downward FM sweeps. Anterograde and retrograde transport was examined following focal injections of wheatgerm agglutinin-horseradish peroxidase (WGA-HRP) into ICXv. The main targets of anterograde transport were the deep layers of the superior colliculus and the suprageniculate division of the medial geniculate body. The primary site of retrograde transport was the nucleus of the central acoustic tract in the brainstem. Thus, the ICXv appears to be a part of the central acoustic tract, an extralemniscal pathway linking the auditory brainstem directly to a multimodal nucleus of the thalamus.

Acoustic Stimulation↗
PubMed1981

The connections of the inferior colliculus and the organization of the brainstem auditory system in the greater horseshoe bat (Rhinolophus ferrumequinum).

The connections of the inferior colliculus, the mammalian mid-brain auditory center, were determined in the greater horseshoe bat (Rhinolophus ferrumequinum), using the horseradish peroxidase method. In order to localize the auditory centers of this bat, brains were investigated with the aid of cell and fiber-stained material. The results show that most auditory centers are highly developed in this echolocating bat. However, the organization of the central auditory system does not generally differ from the mammalian scheme. This holds also for the organization of the superior olivary complex where a well-developed medial superior olivary nucleus was found. In addition to the ventral and dorsal nuclei of the lateral lemniscus a third well-developed nucleus has been defined which projects ipsilaterally to the inferior colliculus and which was called the intermediate nucleus of the lateral leminiscus. All nuclei of the central auditory pathway project ipsi-, contra-, or bilaterally to the central nucleus of the inferior colliculus with the exception of the medial nucleus of the trapezoid body and the medial geniculate body. The tonotopic organization of these projections and their possible functions are discussed in context with neurophysiological investigations.

Animals↗