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Mysterious Mystacina: how the New Zealand short-tailed bat (Mystacina tuberculata) locates insect prey.

The New Zealand short-tailed bat Mystacina tuberculata evolved in the absence of terrestrial mammals and initially with few potential predators. Unusual among bats, it is well adapted for the capture of prey on the ground. Bats from Fiordland, New Zealand had relatively low wing loadings and aspect ratios adapted for flight in cluttered habitats. We predicted that M. tuberculata would locate prey in air (uncluttered space) by echolocation. Echolocation call sequences associated with prey capture (terminal buzzes) were heard in the field, and bats detected and localized prey suspended on fishing line by echolocation in a flight cage. The bats emitted brief, multiharmonic echolocation calls at low duty cycle during search phase, and 64% of calls contained most energy in the fundamental harmonic. Approach- and terminal-phase calls were also broadband and multiharmonic. We predicted that bats would not use echolocation to locate prey hidden on the ground in leaf litter (cluttered space). Bats seemed unable to locate hidden prey precisely from the air and instead hunted for such prey while crawling. Echolocation calls were emitted at a low repetition rate on the ground, suggesting that here echolocation was used for orientation and not for prey detection. We experimentally removed cues available to the bats and showed that bats located mealworms in leaf litter by listening for prey-generated noises and possibly by olfaction.

Animals↗

Timing of praying mantis evasive responses during simulated bat attack sequences.

Praying mantids perform evasive maneuvers that vary with the level of danger posed by their bat predators. The vocalization pattern of attacking bats provides cues that mantids can potentially use to decide how and when to respond. Using pulse trains simulating bat attack echolocation sequences, this study determines when in the attack sequence the mantis power dive (its response to high-level threat) occurs and predicts the parameters within the echolocation sequence that are important for eliciting the response. For sequences with a rapid transition from low to high pulse repetition rates (PRRs), the evasive response occurred close to the point during the simulated sequence when the bat would have contacted the mantis. However, the evasive response occurred earlier if the transition was gradual. Regardless of the transition type, the prediction data show that sequences trigger the response when PRRs reach 20-40 pulses s(-1). These results suggest that a bat gradually increasing its PRR could 'tip off' the mantis, enabling it to escape. Attack sequences contain gradual transitions when bats engage in strobing behavior, an echolocation phenomenon that may help the bat perceive the auditory scene. Conversely, bat attack sequences that contain rapid increases in PRR close to the point of capture could circumvent the mantid's auditory defense. Based on these findings, mantids as well as other insects could benefit from having a back-up defense response to offset any advantage the bat gains by rapidly switching from low to high PRRs.

Acoustic Stimulation↗

Effect of diet quality on carbon and nitrogen turnover and isotopic discrimination in blood of a New World nectarivorous bat.

Diet composition of carbon and nitrogen (C:N) could affect diet-tissue isotopic discrimination and elemental turnover rate in consumers but studies that test the nature of these changes are scarce. We compared carbon and nitrogen isotopic discrimination and turnover rates in individuals of Pallas' long-tongued bats Glossophaga soricina fed diets with protein soya isolate or amaranth grains as their main source of protein. Diets were of similar protein biological value but the soya diet had higher nitrogen content (2.2%N) and lower C:N ratio (39.6) than the amaranth diet (1.3%N, C:N=40.5). Most bats on the soya diet gained body mass whereas most bats on the amaranth diet lost body mass. Half-lives of carbon (24.3+/-3.8 days) and nitrogen (25.6+/-4.4 days) in bats switched to the soya diet were very similar. In contrast, in the bats switched to the amaranth diet, carbon half-life (39.7+/-3.4 days) was longer than that of nitrogen (25.0+/-6.0 days). The enrichment in 15N between diet and blood was higher when bats were fed the amaranth diet (4.4+/-0.2 per thousand) than when they were fed the soya diet (3.3+/-0.2 per thousand). Similarly, bats on the amaranth diet had higher 13C enrichment (2.0+/-0.2 per thousand) than bats on the soya diet (0.1+/-0.1 per thousand). Our results support recent hypotheses of the effect of nutrition on diet-tissue isotopic discrimination and turnover rate, and further shows that blood stable isotope analysis is an adequate approach to track seasonal dietary shifts in wild bats.

Amaranthus↗

Correlations between structure and function in the design of the bat lung: a morphometric study.

The lungs of five species of bat Pipistrellus pipistrellus, Miniopterus minor (Peters), Tadarida mops (De Blainville), Cynopterus brachyotis (Muller) and Cheiromeles torquatus (Horstield) have been analysed by morphometric techniques. The mean body weight (W) ranged from 5 g in Pipistrellus pipistrellus to 173 g in Cheiromeles torquatus; the lung volume (VL) ranged from 0.3 cm3 in Pipistrellus to 10 cm3 in Cheiromeles. The volume densities or the main components of the bat lung, namely the parenchyma [VV(p,L)] (the gas exchange region) and the non-parenchyma [VV(np,L)], were closely similar, the VV(p,L) constituting a mean value of 84.2% and the VV(np,L) 15.8% in the five species. The VL, the surface area of the blood--gas (tissue) barrier (St), the pulmonary capillary blood volume (Vc), and the total morphometric pulmonary diffusing capacity (DLO2) were all strongly correlated with body weight. The harmonic mean thickness of the blood--gas (tissue) barrier (tau ht) and the surface density of the blood--gas (tissue) barrier [SV(t,p)] were poorly correlated with W. The bats had a remarkably higher VL than either birds or terrestrial mammals. The Vc in the bat lung was similar to that in the bird lung but higher than that of the terrestrial mammals. The bats had a more extensive St than either the birds or the terrestrial mammals. In the bats the tau ht was thicker than in the birds but thinner than that of the terrestrial mammals. These pulmonary structural adaptations culminated in a higher DLO2 in the bat than either in the birds or in the terrestrial mammals. The superior morphometric properties of the bat lung coupled with the established physiological adaptations may help to explain how the bat lung is capable of providing the immense amount of oxygen demanded by flight.

Animals↗

Doppler-shift compensation behavior in horseshoe bats revisited: auditory feedback controls both a decrease and an increase in call frequency.

Among mammals, echolocation in bats illustrates the vital role of proper audio-vocal feedback control particularly well. Bats adjust the temporal, spectral and intensity parameters of their echolocation calls depending on the characteristics of the returning echo signal. The mechanism of audio-vocal integration in both mammals and birds is, however, still largely unknown. Here, we present behavioral evidence suggesting a novel audio-vocal control mechanism in echolocating horseshoe bats (Rhinolophus ferrumequinum). These bats compensate for even subtle frequency shifts in the echo caused by flight-induced Doppler effects by adjusting the frequency of their echolocation calls. Under natural conditions, when approaching background targets, the bats usually encounter only positive Doppler shifts. Hence, we commonly believed that, during this Doppler-shift compensation behavior, horseshoe bats use auditory feedback to compensate only for these increases in echo frequency (=positive shifts) by actively lowering their call frequency below the resting frequency (the call frequency emitted when not flying and not experiencing Doppler shifts). Re-investigation of the Doppler-shift compensation behavior, however, shows that decreasing echo frequencies (=negative shifts) are involved as well: auditory feedback from frequencies below the resting frequency, when presented at similar suprathreshold intensity levels as higher echo frequencies, cause the bat's call frequency to increase above the resting frequency. However, compensation for negative shifts is less complete than for positive shifts (22% versus 95%), probably because of biomechanical restrictions in the larynx of bats. Therefore, Doppler-shift compensation behavior involves a quite different neural substrate and audio-vocal control mechanism from those previously assumed. The behavioral results are no longer consistent with solely inhibitory feedback originating from frequencies above the resting frequency. Instead, we propose that auditory feedback follows an antagonistic push/pull principle, with inhibitory feedback lowering and excitatory feedback increasing call frequencies. While the behavioral significance of an active compensation for echo frequencies below RF remains unclear, these behavioral results are crucial for determining the neural implementation of audio-vocal feedback control in horseshoe bats and possibly in mammals in general.

Animals↗

Echolocating bats use a nearly time-optimal strategy to intercept prey.

Acquisition of food in many animal species depends on the pursuit and capture of moving prey. Among modern humans, the pursuit and interception of moving targets plays a central role in a variety of sports, such as tennis, football, Frisbee, and baseball. Studies of target pursuit in animals, ranging from dragonflies to fish and dogs to humans, have suggested that they all use a constant bearing (CB) strategy to pursue prey or other moving targets. CB is best known as the interception strategy employed by baseball outfielders to catch ballistic fly balls. CB is a time-optimal solution to catch targets moving along a straight line, or in a predictable fashion--such as a ballistic baseball, or a piece of food sinking in water. Many animals, however, have to capture prey that may make evasive and unpredictable maneuvers. Is CB an optimum solution to pursuing erratically moving targets? Do animals faced with such erratic prey also use CB? In this paper, we address these questions by studying prey capture in an insectivorous echolocating bat. Echolocating bats rely on sonar to pursue and capture flying insects. The bat's prey may emerge from foliage for a brief time, fly in erratic three-dimensional paths before returning to cover. Bats typically take less than one second to detect, localize and capture such insects. We used high speed stereo infra-red videography to study the three dimensional flight paths of the big brown bat, Eptesicus fuscus, as it chased erratically moving insects in a dark laboratory flight room. We quantified the bat's complex pursuit trajectories using a simple delay differential equation. Our analysis of the pursuit trajectories suggests that bats use a constant absolute target direction strategy during pursuit. We show mathematically that, unlike CB, this approach minimizes the time it takes for a pursuer to intercept an unpredictably moving target. Interestingly, the bat's behavior is similar to the interception strategy implemented in some guided missiles. We suggest that the time-optimal strategy adopted by the bat is in response to the evolutionary pressures of having to capture erratic and fast moving insects.

Animals↗

Representation of perceptual dimensions of insect prey during terminal pursuit by echolocating bats.

The echolocating big brown bat, Eptesicus fuscus, broadcasts brief frequency-modulated (FM) ultrasonic sounds and perceives objects from echoes of these sounds returning to its ears. Eptesicus is an insectivorous species that uses sonar to locate and track flying prey. Although the bat normally hunts in open areas, it nevertheless is capable of chasing insects into cluttered environments such as vegetation, where it completes interceptions in much the same manner as in the open except that it has to avoid the obstacles as well as catch the insect. During pursuit, the bat shortens its sonar signals and increases their rate of emission as it closes in to seize the target, and it keeps its head pointed at the insect throughout the maneuver. In the terminal stage of interception, the bat makes rapid adjustments in its flight-path and body posture to capture the insect, and these reactions occur whether the bat is pursuing its prey in the open or close to obstacles such as vegetation. Insects can be distinguished from other objects by the spectrum and phase of their echoes, and Eptesicus is very good at discriminating these acoustic features. To identify the insect in the open, but especially to distinguish which object is the insect in clutter, the bat must have some means for representing these features throughout the interception maneuver. Moreover, continuity for perception of these features is necessary to keep track of the prey in complex surroundings, so the nature of the auditory representations for the spectrum and phase of echoes has to be conserved across the approach, tracking, and terminal stages. The first problem is that representation of changes in the phase of echoes requires neural responses in the bat's auditory system to have temporal precision in the microsecond range, which seems implausible from conventional single-unit studies in the bat's inferior colliculus, where the temporal jitter of responses typically is hundreds of microseconds. Another problem is that echoes do not explicitly evoke neural responses in the inferior colliculus distinct from responses evoked by the broadcast during the terminal stage because the delay of echoes is too short for responsiveness to recover from the emissions. In contrast, each emission and each echo evokes its own responses during the approach and tracking stages of pursuit. How does the bat consistently represent the phase of echoes in spite of these evident limitations in neural responses? Local multiunit responses recorded from the inferior colliculus of Eptesicus reveal a novel format for encoding the phase of echoes at all stages of interception. Changes in echo phase (0 degree or 180 degrees) produce shifts in the latency of responses to the emission by hundreds of microseconds, an unexpected finding that demonstrates the existence of expanded time scales in neural responses representing the target at all stages of pursuit.

Animals↗

European bat lyssaviruses, The Netherlands.

To study European bat lyssavirus (EBLV) in bat reservoirs in the Netherlands, native bats have been tested for rabies since 1984. For all collected bats, data including species, age, sex, and date and location found were recorded. A total of 1,219 serotine bats, Eptesicus serotinus, were tested, and 251 (21%) were positive for lyssavirus antigen. Five (4%) of 129 specimens from the pond bat, Myotis dasycneme, were positive. Recently detected EBLV RNA segments encoding the nucleoprotein were sequenced and analyzed phylogenetically (45 specimens). All recent serotine bat specimens clustered with genotype 5 (EBLV1) sequences, and homologies within subgenotypes EBLV1a and EBLV1b were 99.0%-100% and 99.2%-100%, respectively. Our findings indicate that EBLVs of genotype 5 are endemic in the serotine bat in the Netherlands. Since EBLVs can cause fatal infections in humans, all serotine and pond bats involved in contact incidents should be tested to determine whether the victim was exposed to EBLVs.

Age Distribution↗

Potential force of infection of human rabies transmitted by vampire bats in the Amazonian region of Brazil.

Human rabies transmitted by bats has acquired greater epidemiologic relevance in various Latin American countries, just when cases transmitted by dogs have decreased. Concern has been heightened by reports of increased rates of bats biting humans in villages in the Amazonian region of Brazil. The aim of the present work was to estimate the potential force of infection (per capita rate at which susceptible individuals acquire infection) of human rabies transmitted by the common vampire bat if the rabies virus were to be introduced to a colony of bats close to a village with a high rate of human bites. The potential force of infection could be then used to anticipate the size of a rabies outbreak in control programs. We present an estimator of potential incidence, adapted from models for malaria. To obtain some of the parameters for the equation, a cross-sectional survey was conducted in Mina Nova, a village of gold prospectors in the Amazonian region of Brazil with high rates of bates biting humans. Bats were captured near dwellings and sent to The Rabies Diagnostic Laboratory at the Center for Control of Zoonoses (Sao Paulo, Brazil) to be examined. To estimate the force of infection, a hypothetical rabies outbreak among bats was simulated using the actual data obtained in the study area. Of 129 people interviewed, 23.33% had been attacked by a vampire bat during the year prior to the study, with an average of 2.8 bites per attacked person. Males (29.41%) were attacked more often than females (11.36%); also, adults (29.35%) were attacked more often than children (8.33%). None of the 12 bats captured in Mina Nova tested positive for rabies, but the force of infection for a hypothetical outbreak was estimated to be 0.0096 per person per year. This risk represents 0.96 cases per 100 area residents, giving an incidence of 1.54 cases of bat-transmitted human rabies per year in the village of Mina Nova (160 inhabitants). The estimated risk is comparable with what has been observed in similar Brazilian villages.

Adult↗

Could bats act as reservoir hosts for Rift Valley fever virus?

The inter-epizootic reservoir host of Rift Valley fever virus (RVFV) remains unknown, although the namaqua rock rat, Aethomys namaquensis, as well as bats have been implicated. Bats can be asymptomatically infected with rabies, as well as several arboviruses; the possibility that they can act as host for RVFV therefore exists. To examine this possibility, 350 different samples (brain, liver, salivary glands and brown fat) obtained from 150 bats (comprising seven species) were tested for RVFV antigen using an enzyme linked immunosorbent assay (ELISA). None of the samples tested positive, but the ELISA proved to have limited sensitivity (> or = 10(3) TCID50/ml). In order to determine whether bats could be infected with RVFV, one Miniopterus schreibersii and two Eptesicus capensis bats were inoculated by the oral or intramuscular route with 100 ml and 30 ml, respectively, of a RVFV suspension with a titre of 10(6) TCID50/ml. None of the bats developed any clinical signs. A low concentration of RVFV antigen was found in the liver and urine of M. schreibersii, but not in brain tissue. A third E. capensis bat was inoculated by the intramuscular route and sacrificed on day 18. A low level of antigen was detected in the brown fat. These results demonstrate that bats can be infected with RVFV, and that further studies should be done to determine the potential of different bat species to act as reservoir hosts for RVFV during inter-epizootic periods.

Animals↗

Observations on the epizootiology of vampire bat rabies.

Vampire bat populations were opportunely sampled before, during, and at varying intervals after outbreaks of bovine rabies. The captured bats were examined for rabies neutralizing antibody and virus. In all, sera from 1,024 vampire bats were tested for antibody, and tissues from these bats plus 83 others were tested for virus. Neutralizing antibody only rarely appeared in vampire serum samples taken before bovine rabies outbreaks, and only low percentages of samples positive for rabies antibody were obtained from bats captured during bovine outbreaks. In contrast, varying percentages of positive samples (including some high percentages) were taken from bats captured at various intervals after bovine outbreaks. Only eleven rabies virus isolations were obtained in the course of this study. In each case the virus came from a bat captured just before or during a bovine outbreak. The authors suggest that rabies virus behaves in vampire populations the way diverse infectious agents typically behave in other hosts. That is, the virus infects many individuals; some die and others survive to demonstrate their exposure through the appearance of anitbody. The disease disappears from the bat population in time and does not return until a sufficient number of susceptible bats have re-entered the population.

Animals↗

[Hemoparasites of bats in Madagascar].

This study aims to evaluate the prevalence and density of haemoparasites in wild malagasy bats. Among the 440 bats, belonging to 14 species sampled in 5 localities in different bio-climatic zones of the island, 93 (21%) showed at least 1 haemoparasite with, by order of frequency, Haemoproteidae (15.7% of 440 bats), microfilariae (7.0%) and Trypanosoma (0.7%). Among these 93 bats, 92 (99%) belonged to the family Vespertilionidae. Four bat species, all endemic to the Madagascar region (Madagascar and Comoros), were found to harbour parasites: Miniopterus manavi with Haemoproteidae (38% of 129 individuals), microfilariae (23%) and Trypanosoma (2%); Myotis goudoti with Haemoproteidae (24% of 68 individuals) and microfilariae (1%); Miniopterus gleni with Haemoproteidae (23% of 13 individuals); and Triaenops furculus with Haemoproteidae (4% of 28 individuals). The sex of bats was not linked to parasite prevalence. Within Miniopterus manavi, those individuals with greater weight also had a higher prevalence of microfilariae; and within the individuals harbouring microfilariae the greatest weights corresponded to the highest density of microfilariae. Ten bat species (with 202 individuals examined) were negative for any haemoparasite. This study is the first to provide evidence of haemoparasites in Malagasy bats; it provides interesting insights, especially concerning the parasite distribution per bat species and families, the pathogenicity of this type of parasitism and the parasite transmission by arthropod vectors.

Altitude↗

Isolation, characterization, and cDNA cloning of a vampire bat salivary plasminogen activator.

Vampire bat saliva contains a plasminogen activator that presumably assists these hematophagous animals during feeding. Here, we report that the vampire bat salivary plasminogen activator, Bat-PA, is homologous to tissue-type plasminogen activator (t-PA) but contains neither a kringle 2 domain nor a plasmin-sensitive processing site. Three Bat-PA species corresponding to full-length, finger-, and finger- epidermal growth factor homology domain- forms of t-PA have been isolated. Bat-PA(H), the full-length form, was purified and its activity has been characterized. Bat-PA(H) and t-PA are of similar efficacy when monitored for their abilities to catalyze plasminogen activation in the presence of a fibrin cofactor. Interestingly, Bat-PA activity toward plasminogen is stimulated 45,000-fold in the presence of fibrin I; the corresponding value for t-PA is only 205-fold. Bat-PA(H) is the only Bat-PA species which binds tightly to fibrin, although each of the three species exhibit remarkable stimulation by a fibrin cofactor.

Amino Acid Sequence↗

Neural computations for sound pattern recognition: evidence for summation of an array of frequency filters in an echolocating bat.

Microchiropteran bats use an auditory sonar system for orientation and prey capture. Many bats use highly structured constant-frequency (CF) and frequency-modulated (FM) sonar orientation signals. Mechanisms for sound pattern recognition are important for the perception of these and other types of auditory signals. The processing and recognition of FM sound components appears to be important for certain complex perceptual tasks, including target distance perception. I have conducted behavioral studies using artificial echoes to simulate the conditions of a bat flying toward a target. An innate vocalization response of the bat to the simulated approaching target was used to assess the ability of the bat to analyze the structure of and extract distance information from different types of synthetic FM sound patterns. The bat's performance depended on the structure of the artificial echo. The pattern recognition performance of the bats was similar when they were presented with either a naturally structured artificial CF/FM echo or an artificial CF/FM echo containing an FM component consisting of a series of pure tone steps. The ability of the bats to recognize appropriately the structure of an FM signal constructed from a sequence of pure tones depended on the number of pure tone steps in the series. Noctilio was able to recognize FM sound patterns containing 99 or greater pure tone steps. The minimum required number of pure tone steps could be distributed over different frequency ranges. The bats were able to resolve individual tone steps in the series that were separated by at least 100 Hz.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Comparative Genomics Reveals Convergent Evolution Between Avivorous Bats (Ia io and Nyctalus aviator).

Investigating the genetic basis of dietary specialization can provide insights into the evolution of niche breadth. In this study, we employed comparative genomics to investigate the adaptive mechanisms enabling two bat species (Nyctalus aviator and Ia io) to shift from insectivory to seasonal bird consumption (avivorous bats). Our findings revealed adaptation related to immune response and lipid metabolism in avivorous bat species. Avivorous bats exhibit strong positive selection and convergent evolution in immune-related genes, which are under heightened selective pressure compared to those of non-avivorous bats. These species also display significantly fewer endogenous retroviral elements. These findings emphasized the significance of immune-driven adaptive evolution in avivory. Additionally, our results showed that the dietary evolution of avivorous bats is accompanied by convergent evolution associated with the lipid metabolism. Notably, CEPT1, the upstream gene required for the activation of the PPAR pathway, underwent positive selection and convergence, which may have affected lipid metabolism. These adaptations may enable avivorous bat species to face the challenge of immune response and nutrition during dietary niche expansion. These findings not only provide comprehensive insights into the adaptive evolution driving the unique diet of avivorous bats but also offered novel perspectives on the molecular mechanisms underlying ecological niche evolution in a dietary context.

Animals↗

Bats as a continuing source of emerging infections in humans.

Amongst the 60 viral species reported to be associated with bats, 59 are RNA viruses, which are potentially important in the generation of emerging and re-emerging infections in humans. The prime examples of these are the lyssaviruses and Henipavirus. The transmission of Nipah, Hendra and perhaps SARS coronavirus and Ebola virus to humans may involve intermediate amplification hosts such as pigs, horses, civets and primates, respectively. Understanding of the natural reservoir or introductory host, the amplifying host, the epidemic centre and at-risk human populations are crucial in the control of emerging zoonosis. The association between the bat coronaviruses and certain lyssaviruses with particular bat species implies co-evolution between specific viruses and bat hosts. Cross-infection between the huge number of bat species may generate new viruses which are able to jump the trans-mammalian species barrier more efficiently. The currently known viruses that have been found in bats are reviewed and the risks of transmission to humans are highlighted. Certain families of bats including the Pteropodidae, Molossidae, Phyllostomidae, and Vespertilionidae are most frequently associated with known human pathogens. A systematic survey of bats is warranted to better understand the ecology of these viruses.

Animals↗

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

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

Animals↗

Complex sound analysis in the FM bat Eptesicus fuscus, correlated with structural parameters of frequency modulated signals.

Big brown bats, Eptesicus fuscus, were presented with artificial frequency modulated (FM) echoes that simulated an object becoming progressively closer to the bat. A stereotyped approach phase behavioral response of the bat to the virtual approaching target was used to determine the ability of the bat to analyze FM signals for target distance information. The degree to which the bats responded with approach phase behavior to a virtual approaching target was similar when they were presented with either a naturally structured artificial FM echo or an artificial FM echo constructed from a series of brief pure tone steps. The ability of the bats to respond to an FM signal structured from a sequence of pure tone elements depended on the number of pure tone steps in the series; the bats required the presentation of tone-step FM signals containing about 83 or greater pure tone elements. Moreover, the duration of the individual tone steps of the tone-step FM signals could not exceed a specific upper limit of about 0.05 ms. Finally, it appears that the bats were able to independently resolve individual tone steps within the tone-step FM signals that were separated by about 450 Hz or more.

Animals↗