Neurophysiological analysis of echolocation in bats

NASA Technical Reports Server (NTRS)

Suga, N.

1972-01-01

An analysis of echolocation and signal processing in brown bats is presented. Data cover echo detection, echo ranging, echolocalization, and echo analysis. Efforts were also made to identify the part of the brain that carries out the most essential processing function for echolocation. Results indicate the inferior colliculus and the auditory nuclei function together to process this information.

Different auditory feedback control for echolocation and communication in horseshoe bats.

PubMed

Liu, Ying; Feng, Jiang; Metzner, Walter

2013-01-01

Auditory feedback from the animal's own voice is essential during bat echolocation: to optimize signal detection, bats continuously adjust various call parameters in response to changing echo signals. Auditory feedback seems also necessary for controlling many bat communication calls, although it remains unclear how auditory feedback control differs in echolocation and communication. We tackled this question by analyzing echolocation and communication in greater horseshoe bats, whose echolocation pulses are dominated by a constant frequency component that matches the frequency range they hear best. To maintain echoes within this "auditory fovea", horseshoe bats constantly adjust their echolocation call frequency depending on the frequency of the returning echo signal. This Doppler-shift compensation (DSC) behavior represents one of the most precise forms of sensory-motor feedback known. We examined the variability of echolocation pulses emitted at rest (resting frequencies, RFs) and one type of communication signal which resembles an echolocation pulse but is much shorter (short constant frequency communication calls, SCFs) and produced only during social interactions. We found that while RFs varied from day to day, corroborating earlier studies in other constant frequency bats, SCF-frequencies remained unchanged. In addition, RFs overlapped for some bats whereas SCF-frequencies were always distinctly different. This indicates that auditory feedback during echolocation changed with varying RFs but remained constant or may have been absent during emission of SCF calls for communication. This fundamentally different feedback mechanism for echolocation and communication may have enabled these bats to use SCF calls for individual recognition whereas they adjusted RF calls to accommodate the daily shifts of their auditory fovea.

Different Auditory Feedback Control for Echolocation and Communication in Horseshoe Bats

PubMed Central

Liu, Ying; Feng, Jiang; Metzner, Walter

2013-01-01

Auditory feedback from the animal's own voice is essential during bat echolocation: to optimize signal detection, bats continuously adjust various call parameters in response to changing echo signals. Auditory feedback seems also necessary for controlling many bat communication calls, although it remains unclear how auditory feedback control differs in echolocation and communication. We tackled this question by analyzing echolocation and communication in greater horseshoe bats, whose echolocation pulses are dominated by a constant frequency component that matches the frequency range they hear best. To maintain echoes within this “auditory fovea”, horseshoe bats constantly adjust their echolocation call frequency depending on the frequency of the returning echo signal. This Doppler-shift compensation (DSC) behavior represents one of the most precise forms of sensory-motor feedback known. We examined the variability of echolocation pulses emitted at rest (resting frequencies, RFs) and one type of communication signal which resembles an echolocation pulse but is much shorter (short constant frequency communication calls, SCFs) and produced only during social interactions. We found that while RFs varied from day to day, corroborating earlier studies in other constant frequency bats, SCF-frequencies remained unchanged. In addition, RFs overlapped for some bats whereas SCF-frequencies were always distinctly different. This indicates that auditory feedback during echolocation changed with varying RFs but remained constant or may have been absent during emission of SCF calls for communication. This fundamentally different feedback mechanism for echolocation and communication may have enabled these bats to use SCF calls for individual recognition whereas they adjusted RF calls to accommodate the daily shifts of their auditory fovea. PMID:23638137

Echolocation in Oilbirds and swiftlets

PubMed Central

Brinkløv, Signe; Fenton, M. Brock; Ratcliffe, John M.

2013-01-01

The discovery of ultrasonic bat echolocation prompted a wide search for other animal biosonar systems, which yielded, among few others, two avian groups. One, the South American Oilbird (Steatornis caripensis: Caprimulgiformes), is nocturnal and eats fruit. The other is a selection of diurnal, insect-eating swiftlets (species in the genera Aerodramus and Collocalia: Apodidae) from across the Indo-Pacific. Bird echolocation is restricted to lower frequencies audible to humans, implying a system of poorer resolution than the ultrasonic (>20 kHz) biosonar of most bats and toothed whales. As such, bird echolocation has been labeled crude or rudimentary. Yet, echolocation is found in at least 16 extant bird species and has evolved several times in avian lineages. Birds use their syringes to produce broadband click-type biosonar signals that allow them to nest in dark caves and tunnels, probably with less predation pressure. There are ongoing discrepancies about several details of bird echolocation, from signal design to the question about whether echolocation is used during foraging. It remains to be seen if bird echolocation is as sophisticated as that of tongue-clicking rousette bats. Bird echolocation performance appears to be superior to that of blind humans using signals of notable similarity. However, no apparent specializations have been found so far in the birds' auditory system (from middle ear to higher processing centers). The advent of light-weight recording equipment and custom software for examining signals and reconstructing flight paths now provides the potential to study the echolocation behavior of birds in more detail and resolve such issues. PMID:23755019

High duty cycle echolocation and prey detection by bats.

PubMed

Lazure, Louis; Fenton, M Brock

2011-04-01

There are two very different approaches to laryngeal echolocation in bats. Although most bats separate pulse and echo in time by signalling at low duty cycles (LDCs), almost 20% of species produce calls at high duty cycles (HDCs) and separate pulse and echo in frequency. HDC echolocators are sensitive to Doppler shifts. HDC echolocation is well suited to detecting fluttering targets such as flying insects against a cluttered background. We used two complementary experiments to evaluate the relative effectiveness of LDC and HDC echolocation for detecting fluttering prey. We measured echoes from fluttering targets by broadcasting artificial bat calls, and found that echo amplitude was greatest for sounds similar to those used in HDC echolocation. We also collected field recordings of syntopic LDC and HDC bats approaching an insect-like fluttering target and found that HDC bats approached the target more often (18.6% of passes) than LDC bats (1.2% of passes). Our results suggest that some echolocation call characteristics, particularly duty cycle and pulse duration, translate into improved ability to detect fluttering targets in clutter, and that HDC echolocation confers a superior ability to detect fluttering prey in the forest understory compared with LDC echolocation. The prevalence of moths in the diets of HDC bats, which is often used as support for the allotonic frequency hypothesis, can therefore be partly explained by the better flutter detection ability of HDC bats.

Evolution of high duty cycle echolocation in bats.

PubMed

Fenton, M Brock; Faure, Paul A; Ratcliffe, John M

2012-09-01

Duty cycle describes the relative 'on time' of a periodic signal. In bats, we argue that high duty cycle (HDC) echolocation was selected for and evolved from low duty cycle (LDC) echolocation because increasing call duty cycle enhanced the ability of echolocating bats to detect, lock onto and track fluttering insects. Most echolocators (most bats and all birds and odontocete cetaceans) use LDC echolocation, separating pulse and echo in time to avoid forward masking. They emit short duration, broadband, downward frequency modulated (FM) signals separated by relatively long periods of silence. In contrast, bats using HDC echolocation emit long duration, narrowband calls dominated by a single constant frequency (CF) separated by relatively short periods of silence. HDC bats separate pulse and echo in frequency by exploiting information contained in Doppler-shifted echoes arising from their movements relative to background objects and their prey. HDC echolocators are particularly sensitive to amplitude and frequency glints generated by the wings of fluttering insects. We hypothesize that narrowband/CF calls produced at high duty cycle, and combined with neurobiological specializations for processing Doppler-shifted echoes, were essential to the evolution of HDC echolocation because they allowed bats to detect, lock onto and track fluttering targets. This advantage was especially important in habitats with dense vegetation that produce overlapping, time-smeared echoes (i.e. background acoustic clutter). We make four specific, testable predictions arising from this hypothesis.

Inconspicuous echolocation in hoary bats (Lasiurus cinereus).

PubMed

Corcoran, Aaron J; Weller, Theodore J

2018-05-16

Echolocation allows bats to occupy diverse nocturnal niches. Bats almost always use echolocation, even when other sensory stimuli are available to guide navigation. Here, using arrays of calibrated infrared cameras and ultrasonic microphones, we demonstrate that hoary bats ( Lasiurus cinereus ) use previously unknown echolocation behaviours that challenge our current understanding of echolocation. We describe a novel call type ('micro' calls) that has three orders of magnitude less sound energy than other bat calls used in open habitats. We also document bats flying close to microphones (less than 3 m) without producing detectable echolocation calls. Acoustic modelling indicates that bats are not producing calls that exceed 70-75 dB at 0.1 m, a level that would have little or no known use for a bat flying in the open at speeds exceeding 7 m s -1 This indicates that hoary bats sometimes fly without echolocation. We speculate that bats reduce echolocation output to avoid eavesdropping by conspecifics during the mating season. These findings might partly explain why tens of thousands of hoary bats are killed by wind turbines each year. They also challenge the long-standing assumption that bats-model organisms for sensory specialization-are reliant on sonar for nocturnal navigation. © 2018 The Author(s).

Unknown beaked whale echolocation signals recorded off eastern New Zealand.

PubMed

Giorli, Giacomo; Goetz, Kimberly T; Delarue, Julien; Maxner, Emily; Kowarski, Katie A; Bruce Martin, Steven; McPherson, Craig

2018-04-01

The echolocation signals of most beaked whale species are still unknown. In fact, out of the 22 species comprising the family Ziphiidae, only the echolocation pulses for 7 species have been clearly described. This study describes two distinct beaked whale echolocation signals recorded in the Cook Strait region using passive acoustic technology. These signals differ from previously described Ziphiid species clicks. A description of the time-frequency characteristics of the two signals is provided. Understanding the characteristics of these signals is necessary to correctly identify species from their echolocation signals and enables future monitoring of beaked whales using passive acoustics techniques.

The physics of bat echolocation: Signal processing techniques

NASA Astrophysics Data System (ADS)

Denny, Mark

2004-12-01

The physical principles and signal processing techniques underlying bat echolocation are investigated. It is shown, by calculation and simulation, how the measured echolocation performance of bats can be achieved.

Human Exploration of Enclosed Spaces through Echolocation.

PubMed

Flanagin, Virginia L; Schörnich, Sven; Schranner, Michael; Hummel, Nadine; Wallmeier, Ludwig; Wahlberg, Magnus; Stephan, Thomas; Wiegrebe, Lutz

2017-02-08

Some blind humans have developed echolocation, as a method of navigation in space. Echolocation is a truly active sense because subjects analyze echoes of dedicated, self-generated sounds to assess space around them. Using a special virtual space technique, we assess how humans perceive enclosed spaces through echolocation, thereby revealing the interplay between sensory and vocal-motor neural activity while humans perform this task. Sighted subjects were trained to detect small changes in virtual-room size analyzing real-time generated echoes of their vocalizations. Individual differences in performance were related to the type and number of vocalizations produced. We then asked subjects to estimate virtual-room size with either active or passive sounds while measuring their brain activity with fMRI. Subjects were better at estimating room size when actively vocalizing. This was reflected in the hemodynamic activity of vocal-motor cortices, even after individual motor and sensory components were removed. Activity in these areas also varied with perceived room size, although the vocal-motor output was unchanged. In addition, thalamic and auditory-midbrain activity was correlated with perceived room size; a likely result of top-down auditory pathways for human echolocation, comparable with those described in echolocating bats. Our data provide evidence that human echolocation is supported by active sensing, both behaviorally and in terms of brain activity. The neural sensory-motor coupling complements the fundamental acoustic motor-sensory coupling via the environment in echolocation. SIGNIFICANCE STATEMENT Passive listening is the predominant method for examining brain activity during echolocation, the auditory analysis of self-generated sounds. We show that sighted humans perform better when they actively vocalize than during passive listening. Correspondingly, vocal motor and cerebellar activity is greater during active echolocation than vocalization alone. Motor and subcortical auditory brain activity covaries with the auditory percept, although motor output is unchanged. Our results reveal behaviorally relevant neural sensory-motor coupling during echolocation. Copyright © 2017 the authors 0270-6474/17/371614-14$15.00/0.

'No cost of echolocation for flying bats' revisited.

PubMed

Voigt, Christian C; Lewanzik, Daniel

2012-08-01

Echolocation is energetically costly for resting bats, but previous experiments suggested echolocation to come at no costs for flying bats. Yet, previous studies did not investigate the relationship between echolocation, flight speed, aerial manoeuvres and metabolism. We re-evaluated the 'no-cost' hypothesis, by quantifying the echolocation pulse rate, the number of aerial manoeuvres (landings and U-turns), and the costs of transport in the 5-g insectivorous bat Rhogeessa io (Vespertilionidae). On average, bats (n = 15) travelled at 1.76 ± 0.36 m s⁻¹ and performed 11.2 ± 6.1 U-turns and 2.8 ± 2.9 ground landings when flying in an octagonal flight cage. Bats made more U-turns with decreasing wing loading (body weight divided by wing area). At flight, bats emitted 19.7 ± 2.7 echolocation pulses s⁻¹ (range 15.3-25.8 pulses s⁻¹), and metabolic rate averaged 2.84 ± 0.95 ml CO₂ min⁻¹, which was more than 16 times higher than at rest. Bats did not echolocate while not engaged in flight. Costs of transport were not related to the rate of echolocation pulse emission or the number of U-turns, but increased with increasing number of landings; probably as a consequence of slower travel speed when staying briefly on ground. Metabolic power of flight was lower than predicted for R. io under the assumption that energetic costs of echolocation call production is additive to the aerodynamic costs of flight. Results of our experiment are consistent with the notion that echolocation does not add large energetic costs to the aerodynamic power requirements of flight in bats.

A blind human expert echolocator shows size constancy for objects perceived by echoes.

PubMed

Milne, Jennifer L; Anello, Mimma; Goodale, Melvyn A; Thaler, Lore

2015-01-01

Some blind humans make clicking noises with their mouth and use the reflected echoes to perceive objects and surfaces. This technique can operate as a crude substitute for vision, allowing human echolocators to perceive silent, distal objects. Here, we tested if echolocation would, like vision, show size constancy. To investigate this, we asked a blind expert echolocator (EE) to echolocate objects of different physical sizes presented at different distances. The EE consistently identified the true physical size of the objects independent of distance. In contrast, blind and blindfolded sighted controls did not show size constancy, even when encouraged to use mouth clicks, claps, or other signals. These findings suggest that size constancy is not a purely visual phenomenon, but that it can operate via an auditory-based substitute for vision, such as human echolocation.

Application of the Biosonar Measurement Tool (BMT) and Instrumented Mine Simulators (IMS) to Exploration of Dolphin Echolocation During Free-Swimming, Bottom-Object Searches

DTIC Science & Technology

2003-09-01

0-933957-31-9 311 Application of the Biosonar Measurement Tool (BMT) and Instrumented...dolphin biosonar (echolocation). Research work conducted by the Navy has addressed the characteristics of echolocation clicks, mechanisms of...information on dolphin echolocation that can be data mined for biosonar search strategies under real-world conditions. Results can be applied to the

Vocalization of echolocation-like pulses for interindividual interaction in horseshoe bats (Rhinolophus ferrumequinum).

PubMed

Kobayasi, Kohta I; Hiryu, Shizuko; Shimozawa, Ryota; Riquimaroux, Hiroshi

2012-11-01

Although much is known about the echolocation of horseshoe bats (Rhinolophus spp.), little is known about the characteristics and function of their communication calls. This study focused on a stereotyped behavior of a bat approaching a companion animal in the colony, and examined their interaction and vocalization during this behavior. The bats emit echolocation-like vocalizations when approaching each other and these vocalizations contain a "buildup" pulse sequence, in which the frequency of the pulse increases gradually to normal echolocation pulse frequencies. The results suggest that the echolocation-like pulses serve an important role in communication within the colony.

Depth dependent variation of the echolocation pulse rate of bottlenose dolphins (Tursiops truncatus).

PubMed

Simard, Peter; Hibbard, Ashley L; McCallister, Kimberly A; Frankel, Adam S; Zeddies, David G; Sisson, Geoffrey M; Gowans, Shannon; Forys, Elizabeth A; Mann, David A

2010-01-01

Trained odontocetes appear to have good control over the timing (pulse rate) of their echolocation clicks; however, there is comparatively little information about how free-ranging odontocetes modify their echolocation in relation to their environment. This study investigates echolocation pulse rate in 14 groups of free-ranging bottlenose dolphins (Tursiops truncatus) at a variety of depths (2.4-30.1 m) in the Gulf of Mexico. Linear regression models indicated a significant decrease in mean pulse rate with mean water depth. Pulse rates for most groups were multi-modal. Distance to target estimates were as high as 91.8 m, assuming that echolocation was produced at a maximal rate for the target distance. A 5.29-ms processing lag time was necessary to explain the pulse rate modes observed. Although echolocation is likely reverberation limited, these results support the hypotheses that free-ranging bottlenose dolphins in this area are adapting their echolocation signals for a variety of target detection and ranging purposes, and that the target distance is a function of water depth.

Instrumenting free-swimming dolphins echolocating in open water.

PubMed

Martin, Stephen W; Phillips, Michael; Bauer, Eric J; Moore, Patrick W; Houser, Dorian S

2005-04-01

Dolphins within the Navy Marine Mammal Program use echolocation to effectively locate underwater mines. They currently outperform manmade systems at similar tasks, particularly in cluttered environments and on buried targets. In hopes of improving manmade mine-hunting sonar systems, two instrumentation packages were developed to monitor free-swimming dolphin motion and echolocation during open-water target detection tasks. The biosonar measurement tool (BMT) is carried by a dolphin and monitors underwater position and attitude while simultaneously recording echolocation clicks and returning echoes through high-gain binaural receivers. The instrumented mine simulator (IMS) is a modified bottom target that monitors echolocation signals arriving at the target during ensonification. Dolphin subjects were trained to carry the BMT in open-bay bottom-object target searches in which the IMS could serve as a bottom object. The instrumentation provides detailed data that reveal hereto-unavailable information on the search strategies of free-swimming dolphins conducting open-water, bottom-object search tasks with echolocation.

Inferring echolocation in ancient bats.

PubMed

Simmons, Nancy B; Seymour, Kevin L; Habersetzer, Jörg; Gunnell, Gregg F

2010-08-19

Laryngeal echolocation, used by most living bats to form images of their surroundings and to detect and capture flying prey, is considered to be a key innovation for the evolutionary success of bats, and palaeontologists have long sought osteological correlates of echolocation that can be used to infer the behaviour of fossil bats. Veselka et al. argued that the most reliable trait indicating echolocation capabilities in bats is an articulation between the stylohyal bone (part of the hyoid apparatus that supports the throat and larynx) and the tympanic bone, which forms the floor of the middle ear. They examined the oldest and most primitive known bat, Onychonycteris finneyi (early Eocene, USA), and argued that it showed evidence of this stylohyal-tympanic articulation, from which they concluded that O. finneyi may have been capable of echolocation. We disagree with their interpretation of key fossil data and instead argue that O. finneyi was probably not an echolocating bat.

Instrumenting free-swimming dolphins echolocating in open water

NASA Astrophysics Data System (ADS)

Martin, Stephen W.; Phillips, Michael; Bauer, Eric J.; Moore, Patrick W.; Houser, Dorian S.

2005-04-01

Dolphins within the Navy Marine Mammal Program use echolocation to effectively locate underwater mines. They currently outperform manmade systems at similar tasks, particularly in cluttered environments and on buried targets. In hopes of improving manmade mine-hunting sonar systems, two instrumentation packages were developed to monitor free-swimming dolphin motion and echolocation during open-water target detection tasks. The biosonar measurement tool (BMT) is carried by a dolphin and monitors underwater position and attitude while simultaneously recording echolocation clicks and returning echoes through high-gain binaural receivers. The instrumented mine simulator (IMS) is a modified bottom target that monitors echolocation signals arriving at the target during ensonification. Dolphin subjects were trained to carry the BMT in open-bay bottom-object target searches in which the IMS could serve as a bottom object. The instrumentation provides detailed data that reveal hereto-unavailable information on the search strategies of free-swimming dolphins conducting open-water, bottom-object search tasks with echolocation. .

Testing Convergent Evolution in Auditory Processing Genes between Echolocating Mammals and the Aye-Aye, a Percussive-Foraging Primate

PubMed Central

Jerjos, Michael; Hohman, Baily; Lauterbur, M. Elise; Kistler, Logan

2017-01-01

Abstract Several taxonomically distinct mammalian groups—certain microbats and cetaceans (e.g., dolphins)—share both morphological adaptations related to echolocation behavior and strong signatures of convergent evolution at the amino acid level across seven genes related to auditory processing. Aye-ayes (Daubentonia madagascariensis) are nocturnal lemurs with a specialized auditory processing system. Aye-ayes tap rapidly along the surfaces of trees, listening to reverberations to identify the mines of wood-boring insect larvae; this behavior has been hypothesized to functionally mimic echolocation. Here we investigated whether there are signals of convergence in auditory processing genes between aye-ayes and known mammalian echolocators. We developed a computational pipeline (Basic Exon Assembly Tool) that produces consensus sequences for regions of interest from shotgun genomic sequencing data for nonmodel organisms without requiring de novo genome assembly. We reconstructed complete coding region sequences for the seven convergent echolocating bat–dolphin genes for aye-ayes and another lemur. We compared sequences from these two lemurs in a phylogenetic framework with those of bat and dolphin echolocators and appropriate nonecholocating outgroups. Our analysis reaffirms the existence of amino acid convergence at these loci among echolocating bats and dolphins; some methods also detected signals of convergence between echolocating bats and both mice and elephants. However, we observed no significant signal of amino acid convergence between aye-ayes and echolocating bats and dolphins, suggesting that aye-aye tap-foraging auditory adaptations represent distinct evolutionary innovations. These results are also consistent with a developing consensus that convergent behavioral ecology does not reliably predict convergent molecular evolution. PMID:28810710

When echolocating bats do not echolocate

PubMed Central

Chiu, Chen

2008-01-01

Echolocating bats are known to continuously generate high frequency sonar pulses and listen to the reflecting echoes to localize objects and orient in the environment. However, silent behavior has been reported in a recent paper, which demonstrated that the big brown bat (Eptesicus fuscus) can fly a relative long distant (0.6 to 8 m) without echolocating when flying with another conspecific in a large flight room.1 Methodology and conclusion developed in this study have the potential for further experimental design to answer the question of how millions of bats navigate and orient in cohesive groups. In addition, the discovery of silent behavior suggests the possible use of cooperative sonar in echolocating animals. PMID:19704880

Allen's big-eared bat (Idionycteris phyllotis) documented in colorado based on recordings of its distinctive echolocation call

USGS Publications Warehouse

Hayes, M.A.; Navo, K.W.; Bonewell, L.; Mosch, C.J.; Adams, Rick A.

2009-01-01

Allen's big-eared bat (Idionycteris phyllotis) inhabits much of the southwestern USA, but has not been documented in Colorado. We recorded echolocation calls consistent with I. phyllotis near La Sal Creek, Montrose County, Colorado. Based on characteristics of echolocation calls and flight behavior, we conclude that the echolocation calls described here were emitted by I. phyllotis and that they represent the first documentation of this species in Colorado.

Marine Mammals: Hearing and Echolocation at Coconut Island

DTIC Science & Technology

2012-09-30

1 DISTRIBUTION STATEMENT A. Approved for public release; distribution is unlimited. Marine Mammals: Hearing and Echolocation at Coconut ...REPORT DATE 2012 2. REPORT TYPE N/A 3. DATES COVERED - 4. TITLE AND SUBTITLE Marine Mammals: Hearing and Echolocation at Coconut Island

Driving factors for the evolution of species-specific echolocation call design in new world free-tailed bats (molossidae).

PubMed

Jung, Kirsten; Molinari, Jesús; Kalko, Elisabeth K V

2014-01-01

Phylogeny, ecology, and sensorial constraints are thought to be the most important factors influencing echolocation call design in bats. The Molossidae is a diverse bat family with a majority of species restricted to tropical and subtropical regions. Most molossids are specialized to forage for insects in open space, and thus share similar navigational challenges. We use an unprecedented dataset on the echolocation calls of 8 genera and 18 species of New World molossids to explore how habitat, phylogenetic relatedness, body mass, and prey perception contribute to echolocation call design. Our results confirm that, with the exception of the genus Molossops, echolocation calls of these bats show a typical design for open space foraging. Two lines of evidence point to echolocation call structure of molossids reflecting phylogenetic relatedness. First, such structure is significantly more similar within than among genera. Second, except for allometric scaling, such structure is nearly the same in congeneric species. Despite contrasting body masses, 12 of 18 species call within a relatively narrow frequency range of 20 to 35 kHz, a finding that we explain by using a modeling approach whose results suggest this frequency range to be an adaptation optimizing prey perception in open space. To conclude, we argue that the high variability in echolocation call design of molossids is an advanced evolutionary trait allowing the flexible adjustment of echolocation systems to various sensorial challenges, while conserving sender identity for social communication. Unraveling evolutionary drivers for echolocation call design in bats has so far been hampered by the lack of adequate model organisms sharing a phylogenetic origin and facing similar sensorial challenges. We thus believe that knowledge of the echolocation call diversity of New World molossid bats may prove to be landmark to understand the evolution and functionality of species-specific signal design in bats.

Driving Factors for the Evolution of Species-Specific Echolocation Call Design in New World Free-Tailed Bats (Molossidae)

PubMed Central

Jung, Kirsten; Molinari, Jesús

2014-01-01

Phylogeny, ecology, and sensorial constraints are thought to be the most important factors influencing echolocation call design in bats. The Molossidae is a diverse bat family with a majority of species restricted to tropical and subtropical regions. Most molossids are specialized to forage for insects in open space, and thus share similar navigational challenges. We use an unprecedented dataset on the echolocation calls of 8 genera and 18 species of New World molossids to explore how habitat, phylogenetic relatedness, body mass, and prey perception contribute to echolocation call design. Our results confirm that, with the exception of the genus Molossops, echolocation calls of these bats show a typical design for open space foraging. Two lines of evidence point to echolocation call structure of molossids reflecting phylogenetic relatedness. First, such structure is significantly more similar within than among genera. Second, except for allometric scaling, such structure is nearly the same in congeneric species. Despite contrasting body masses, 12 of 18 species call within a relatively narrow frequency range of 20 to 35 kHz, a finding that we explain by using a modeling approach whose results suggest this frequency range to be an adaptation optimizing prey perception in open space. To conclude, we argue that the high variability in echolocation call design of molossids is an advanced evolutionary trait allowing the flexible adjustment of echolocation systems to various sensorial challenges, while conserving sender identity for social communication. Unraveling evolutionary drivers for echolocation call design in bats has so far been hampered by the lack of adequate model organisms sharing a phylogenetic origin and facing similar sensorial challenges. We thus believe that knowledge of the echolocation call diversity of New World molossid bats may prove to be landmark to understand the evolution and functionality of species-specific signal design in bats. PMID:24454833

A false killer whale adjusts its hearing when it echolocates.

PubMed

Nachtigall, Paul E; Supin, Alexander Y

2008-06-01

The use of auditory evoked potential (AEP) measurements has added considerably to knowledge of the hearing mechanisms of marine mammals. We have recently measured the hearing of a stranded infant Risso's dolphin, the audiograms of white-beaked dolphins temporarily caught and released, and the hearing of anaesthetized polar bears. Most small toothed whales echolocate and hear very high frequency sounds underwater. While much has previously been learned about the echolocation performance and characteristics of the outgoing signals of echolocating dolphins and small whales, the hearing processes occurring while these animals actively echolocate have not previously been examined. Working with a well-trained echolocating false killer whale (Pseudorca crassidens) wearing latex surface suction cup electrodes, we have measured echolocation hearing AEPs in response to outgoing echolocation clicks, returning echoes, and comparable simulated whale clicks and echoes in a variety of situations. We have found that: (1) the whale may hear her loud outgoing clicks and much quieter returning echoes at comparable levels, (2) the whale has protective mechanisms that dampen the intensity of her outgoing signals - she hears her outgoing signals at a level about 40 dB lower than similar signals presented directly in front of her, (3) when echo return levels are lowered either by making the targets smaller or by placing the targets farther away - without changing the levels of her outgoing signals - the hearing of these echoes remains at almost the same level, (4) if targets are made much smaller and harder to echolocate, the animal will modify what she hears of her outgoing signal - as if to heighten overall hearing sensitivity to keep the echo level hearable, (5) the animal has an active 'automatic gain control' mechanism in her hearing based on both forward masking that balances outgoing pulse intensity and time between pulse and echo, and active hearing control. Overall, hearing during echolocation appears to be a very active process.

Flight and echolocation behaviour of three vespertilionid bat species while commuting on flyways.

PubMed

Schaub, Andrea; Schnitzler, Hans-Ulrich

2007-12-01

This study compares the flight and echolocation behaviour of three vespertilionid bat species while they commute on flyways. We measured the bats' spatial position relative to vertical background contours and relative to the ground while recording their echolocation behaviour. In Myotis daubentonii, we found a significant influence of spatial context on the position and dimensions of flyways as well as on echolocation behaviour. In gap situations, flyways tended to be narrower and located closer to background structures, flight speeds were lower and the bandwidth of echolocation signals was larger than in edge situations. Differences in background structure did not affect flight and echolocation behaviour. When commuting in the same gap situation flyway positions and dimensions for M. daubentonii and Myotis brandtii were similar but differed from those of Pipistrellus pipistrellus, which were slightly higher and further out than those used by the Myotis species. In M. brandtii, flyway positions and dimensions remained constant over 3 years. We found species-dependent differences in signal structure, but pulse interval and flight speed were similar across all species. The influence of available space on the position of flyways, on flight speed and on echolocation behaviour is discussed.

Echolocation in humans: an overview.

PubMed

Thaler, Lore; Goodale, Melvyn A

2016-11-01

Bats and dolphins are known for their ability to use echolocation. They emit bursts of sounds and listen to the echoes that bounce back to detect the objects in their environment. What is not as well-known is that some blind people have learned to do the same thing, making mouth clicks, for example, and using the returning echoes from those clicks to sense obstacles and objects of interest in their surroundings. The current review explores some of the research that has examined human echolocation and the changes that have been observed in the brains of echolocation experts. We also discuss potential applications and assistive technology based on echolocation. Blind echolocation experts can sense small differences in the location of objects, differentiate between objects of various sizes and shapes, and even between objects made of different materials, just by listening to the reflected echoes from mouth clicks. It is clear that echolocation may enable some blind people to do things that are otherwise thought to be impossible without vision, potentially providing them with a high degree of independence in their daily lives and demonstrating that echolocation can serve as an effective mobility strategy in the blind. Neuroimaging has shown that the processing of echoes activates brain regions in blind echolocators that would normally support vision in the sighted brain, and that the patterns of these activations are modulated by the information carried by the echoes. This work is shedding new light on just how plastic the human brain is. WIREs Cogn Sci 2016, 7:382-393. doi: 10.1002/wcs.1408 For further resources related to this article, please visit the WIREs website. © 2016 Wiley Periodicals, Inc.

Testing Convergent Evolution in Auditory Processing Genes between Echolocating Mammals and the Aye-Aye, a Percussive-Foraging Primate.

PubMed

Bankoff, Richard J; Jerjos, Michael; Hohman, Baily; Lauterbur, M Elise; Kistler, Logan; Perry, George H

2017-07-01

Several taxonomically distinct mammalian groups-certain microbats and cetaceans (e.g., dolphins)-share both morphological adaptations related to echolocation behavior and strong signatures of convergent evolution at the amino acid level across seven genes related to auditory processing. Aye-ayes (Daubentonia madagascariensis) are nocturnal lemurs with a specialized auditory processing system. Aye-ayes tap rapidly along the surfaces of trees, listening to reverberations to identify the mines of wood-boring insect larvae; this behavior has been hypothesized to functionally mimic echolocation. Here we investigated whether there are signals of convergence in auditory processing genes between aye-ayes and known mammalian echolocators. We developed a computational pipeline (Basic Exon Assembly Tool) that produces consensus sequences for regions of interest from shotgun genomic sequencing data for nonmodel organisms without requiring de novo genome assembly. We reconstructed complete coding region sequences for the seven convergent echolocating bat-dolphin genes for aye-ayes and another lemur. We compared sequences from these two lemurs in a phylogenetic framework with those of bat and dolphin echolocators and appropriate nonecholocating outgroups. Our analysis reaffirms the existence of amino acid convergence at these loci among echolocating bats and dolphins; some methods also detected signals of convergence between echolocating bats and both mice and elephants. However, we observed no significant signal of amino acid convergence between aye-ayes and echolocating bats and dolphins, suggesting that aye-aye tap-foraging auditory adaptations represent distinct evolutionary innovations. These results are also consistent with a developing consensus that convergent behavioral ecology does not reliably predict convergent molecular evolution. © The Author 2017. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution.

Single-lobed frequency-dependent beam shape in an echolocating false killer whale (Pseudorca crassidens).

PubMed

Kloepper, Laura N; Nachtigall, Paul E; Quintos, Christopher; Vlachos, Stephanie A

2012-01-01

Recent studies indicate some odontocetes may produce echolocation beams with a dual-lobed vertical structure. The shape of the odontocete echolocation beam was further investigated in a false killer whale performing an echolocation discrimination task. Clicks were recorded with an array of 16 hydrophones and frequency-dependent amplitude plots were constructed to assess beam shape. The majority of the echolocation clicks were single-lobed in structure with most energy located between 20 and 80 kHz. These data indicate the false killer whale does not produce a dual-lobed structure, as has been shown in bottlenose dolphins, which may be a function of lowered frequencies in the emitted signal due to hearing loss. © 2012 Acoustical Society of America.

Sensory biology: echolocation from click to call, mouth to wing.

PubMed

Fenton, M Brock; Ratcliffe, John M

2014-12-15

Echolocators use echoes of sounds they produce, clicks or calls, to detect objects. Usually, these signals originate from the head. New work reveals that three species of bats use their wings to generate echolocation signals. Copyright © 2014 Elsevier Ltd. All rights reserved.

Patterns and causes of geographic variation in bat echolocation pulses.

PubMed

Jiang, Tinglei; Wu, Hui; Feng, Jiang

2015-05-01

Evolutionary biologists have a long-standing interest in how acoustic signals in animals vary geographically, because divergent ecology and sensory perception play an important role in speciation. Geographic comparisons are valuable in determining the factors that influence divergence of acoustic signals. Bats are social mammals and they depend mainly on echolocation pulses to locate prey, to navigate and to communicate. Mounting evidence shows that geographic variation of bat echolocation pulses is common, with a mean 5-10 kHz differences in peak frequency, and a high level of individual variation may be nested in this geographical variation. However, understanding the geographic variation of echolocation pulses in bats is very difficult, because of differences in sample and statistical analysis techniques as well as the variety of factors shaping the vocal geographic evolution. Geographic differences in echolocation pulses of bats generally lack latitudinal, longitudinal and elevational patterns, and little is known about vocal dialects. Evidence is accumulating to support the fact that geographic variation in echolocation pulses of bats may be caused by genetic drift, cultural drift, ecological selection, sexual selection and social selection. Future studies could relate geographic differences in echolocation pulses to social adaptation, vocal learning strategies and patterns of dispersal. In addition, new statistical techniques and acoustic playback experiments may help to illustrate the causes and consequences of the geographic evolution of echolocation pulse in bats. © 2015 International Society of Zoological Sciences, Institute of Zoology/Chinese Academy of Sciences and Wiley Publishing Asia Pty Ltd.

Independent Losses of Visual Perception Genes Gja10 and Rbp3 in Echolocating Bats (Order: Chiroptera)

PubMed Central

Shen, Bin; Fang, Tao; Dai, Mengyao; Jones, Gareth; Zhang, Shuyi

2013-01-01

A trade-off between the sensory modalities of vision and hearing is likely to have occurred in echolocating bats as the sophisticated mechanism of laryngeal echolocation requires considerable neural processing and has reduced the reliance of echolocating bats on vision for perceiving the environment. If such a trade-off exists, it is reasonable to hypothesize that some genes involved in visual function may have undergone relaxed selection or even functional loss in echolocating bats. The Gap junction protein, alpha 10 (Gja10, encoded by Gja10 gene) is expressed abundantly in mammal retinal horizontal cells and plays an important role in horizontal cell coupling. The interphotoreceptor retinoid-binding protein (Irbp, encoded by the Rbp3 gene) is mainly expressed in interphotoreceptor matrix and is known to be critical for normal functioning of the visual cycle. We sequenced Gja10 and Rbp3 genes in a taxonomically wide range of bats with divergent auditory characteristics (35 and 18 species for Gja10 and Rbp3, respectively). Both genes have became pseudogenes in species from the families Hipposideridae and Rhinolophidae that emit constant frequency echolocation calls with Doppler shift compensation at high-duty-cycles (the most sophisticated form of biosonar known), and in some bat species that emit echolocation calls at low-duty-cycles. Our study thus provides further evidence for the hypothesis that a trade-off occurs at the genetic level between vision and echolocation in bats. PMID:23874796

Independent losses of visual perception genes Gja10 and Rbp3 in echolocating bats (Order: Chiroptera).

PubMed

Shen, Bin; Fang, Tao; Dai, Mengyao; Jones, Gareth; Zhang, Shuyi

2013-01-01

A trade-off between the sensory modalities of vision and hearing is likely to have occurred in echolocating bats as the sophisticated mechanism of laryngeal echolocation requires considerable neural processing and has reduced the reliance of echolocating bats on vision for perceiving the environment. If such a trade-off exists, it is reasonable to hypothesize that some genes involved in visual function may have undergone relaxed selection or even functional loss in echolocating bats. The Gap junction protein, alpha 10 (Gja10, encoded by Gja10 gene) is expressed abundantly in mammal retinal horizontal cells and plays an important role in horizontal cell coupling. The interphotoreceptor retinoid-binding protein (Irbp, encoded by the Rbp3 gene) is mainly expressed in interphotoreceptor matrix and is known to be critical for normal functioning of the visual cycle. We sequenced Gja10 and Rbp3 genes in a taxonomically wide range of bats with divergent auditory characteristics (35 and 18 species for Gja10 and Rbp3, respectively). Both genes have became pseudogenes in species from the families Hipposideridae and Rhinolophidae that emit constant frequency echolocation calls with Doppler shift compensation at high-duty-cycles (the most sophisticated form of biosonar known), and in some bat species that emit echolocation calls at low-duty-cycles. Our study thus provides further evidence for the hypothesis that a trade-off occurs at the genetic level between vision and echolocation in bats.

Bats aloft: Variation in echolocation call structure at high altitudes

USDA-ARS?s Scientific Manuscript database

Bats alter their echolocation calls in response to changes in ecological and behavioral conditions, but little is known about how they adjust their call structure in response to changes in altitude. This study examines altitudinal variation in the echolocation calls of Brazilian free-tailed bats, T...

Discovery Learning in Autonomous Agents Using Genetic Algorithms

DTIC Science & Technology

1993-12-01

Meyer and Wilson (47). 65. Roitblat , H. L., et al. "Biomimetic Sonar Processing: Prom Dolphin Echoloc-Ation to Artificial Neural Networks." In Meyer and...34 In Meyer and Wilson (47). 65. Roitblat , H. L., et al. "Biomimetic Sonar Processing: From Dolphin Echolocation to Artificial Neural Networks." In

Time-frequency and advanced frequency estimation techniques for the investigation of bat echolocation calls.

PubMed

Kopsinis, Yannis; Aboutanios, Elias; Waters, Dean A; McLaughlin, Steve

2010-02-01

In this paper, techniques for time-frequency analysis and investigation of bat echolocation calls are studied. Particularly, enhanced resolution techniques are developed and/or used in this specific context for the first time. When compared to traditional time-frequency representation methods, the proposed techniques are more capable of showing previously unseen features in the structure of bat echolocation calls. It should be emphasized that although the study is focused on bat echolocation recordings, the results are more general and applicable to many other types of signal.

A new fossil species supports an early origin for toothed whale echolocation.

PubMed

Geisler, Jonathan H; Colbert, Matthew W; Carew, James L

2014-04-17

Odontocetes (toothed whales, dolphins and porpoises) hunt and navigate through dark and turbid aquatic environments using echolocation; a key adaptation that relies on the same principles as sonar. Among echolocating vertebrates, odontocetes are unique in producing high-frequency vocalizations at the phonic lips, a constriction in the nasal passages just beneath the blowhole, and then using air sinuses and the melon to modulate their transmission. All extant odontocetes seem to echolocate; however, exactly when and how this complex behaviour--and its underlying anatomy--evolved is largely unknown. Here we report an odontocete fossil, Oligocene in age (approximately 28 Myr ago), from South Carolina (Cotylocara macei, gen. et sp. nov.) that has several features suggestive of echolocation: a dense, thick and downturned rostrum; air sac fossae; cranial asymmetry; and exceptionally broad maxillae. Our phylogenetic analysis places Cotylocara in a basal clade of odontocetes, leading us to infer that a rudimentary form of echolocation evolved in the early Oligocene, shortly after odontocetes diverged from the ancestors of filter-feeding whales (mysticetes). This was followed by enlargement of the facial muscles that modulate echolocation calls, which in turn led to marked, convergent changes in skull shape in the ancestors of Cotylocara, and in the lineage leading to extant odontocetes.

It's not black or white—on the range of vision and echolocation in echolocating bats

PubMed Central

Boonman, Arjan; Bar-On, Yinon; Cvikel, Noam; Yovel, Yossi

2013-01-01

Around 1000 species of bats in the world use echolocation to navigate, orient, and detect insect prey. Many of these bats emerge from their roost at dusk and start foraging when there is still light available. It is however unclear in what way and to which extent navigation, or even prey detection in these bats is aided by vision. Here we compare the echolocation and visual detection ranges of two such species of bats which rely on different foraging strategies (Rhinopoma microphyllum and Pipistrellus kuhlii). We find that echolocation is better than vision for detecting small insects even in intermediate light levels (1–10 lux), while vision is advantageous for monitoring far-away landscape elements in both species. We thus hypothesize that, bats constantly integrate information acquired by the two sensory modalities. We suggest that during evolution, echolocation was refined to detect increasingly small targets in conjunction with using vision. To do so, the ability to hear ultrasonic sound is a prerequisite which was readily available in small mammals, but absent in many other animal groups. The ability to exploit ultrasound to detect very small targets, such as insects, has opened up a large nocturnal niche to bats and may have spurred diversification in both echolocation and foraging tactics. PMID:24065924

Behavioral evidence for community-wide species discrimination from echolocation calls in bats.

PubMed

Schuchmann, Maike; Siemers, Björn M

2010-07-01

Recognizing species identity is crucial for many aspects of animal life and is often mediated by acoustic signals. Although most animals are able to distinguish acoustic signals of their own species from other sympatrically occurring species, it is yet unknown whether animals can distinguish among acoustic signals of different closely related sympatric species. In this context, echolocating bats are a particularly interesting model system: their echolocation system evolved primarily for spatial orientation and foraging, but recent studies indicate that echolocation also has an important communicative function. Yet, the role of echolocation calls for species discrimination and thus potentially for interspecific communication has not been investigated. Using a behavioral discrimination assay, we found that two species of wild horseshoe bats could discriminate calls of their own species from those of three sympatric congeneric species. We further show that the bats were able to discriminate between echolocation calls of different congeneric species from the local community. In both cases, discrimination ability was high despite strong overlap of species' call frequency bands. This study provides the first experimental evidence for species discrimination based on echolocation calls. On a more general level, it shows for the first time that animals can distinguish among acoustic signals of different closely related and ecologically similar species from their local community.

Perception of silent and motionless prey on vegetation by echolocation in the gleaning bat Micronycteris microtis.

PubMed

Geipel, Inga; Jung, Kirsten; Kalko, Elisabeth K V

2013-03-07

Gleaning insectivorous bats that forage by using echolocation within dense forest vegetation face the sensorial challenge of acoustic masking effects. Active perception of silent and motionless prey in acoustically cluttered environments by echolocation alone has thus been regarded impossible. The gleaning insectivorous bat Micronycteris microtis however, forages in dense understory vegetation and preys on insects, including dragonflies, which rest silent and motionless on vegetation. From behavioural experiments, we show that M. microtis uses echolocation as the sole sensorial modality for successful prey perception within a complex acoustic environment. All individuals performed a stereotypical three-dimensional hovering flight in front of prey items, while continuously emitting short, multi-harmonic, broadband echolocation calls. We observed a high precision in target localization which suggests that M. microtis perceives a detailed acoustic image of the prey based on shape, surface structure and material. Our experiments provide, to our knowledge, the first evidence that a gleaning bat uses echolocation alone for successful detection, classification and precise localization of silent and motionless prey in acoustic clutter. Overall, we conclude that the three-dimensional hovering flight of M. microtis in combination with a frequent emission of short, high-frequency echolocation calls is the key for active prey perception in acoustically highly cluttered environments.

Perception of silent and motionless prey on vegetation by echolocation in the gleaning bat Micronycteris microtis

PubMed Central

Geipel, Inga; Jung, Kirsten; Kalko, Elisabeth K. V.

2013-01-01

Gleaning insectivorous bats that forage by using echolocation within dense forest vegetation face the sensorial challenge of acoustic masking effects. Active perception of silent and motionless prey in acoustically cluttered environments by echolocation alone has thus been regarded impossible. The gleaning insectivorous bat Micronycteris microtis however, forages in dense understory vegetation and preys on insects, including dragonflies, which rest silent and motionless on vegetation. From behavioural experiments, we show that M. microtis uses echolocation as the sole sensorial modality for successful prey perception within a complex acoustic environment. All individuals performed a stereotypical three-dimensional hovering flight in front of prey items, while continuously emitting short, multi-harmonic, broadband echolocation calls. We observed a high precision in target localization which suggests that M. microtis perceives a detailed acoustic image of the prey based on shape, surface structure and material. Our experiments provide, to our knowledge, the first evidence that a gleaning bat uses echolocation alone for successful detection, classification and precise localization of silent and motionless prey in acoustic clutter. Overall, we conclude that the three-dimensional hovering flight of M. microtis in combination with a frequent emission of short, high-frequency echolocation calls is the key for active prey perception in acoustically highly cluttered environments. PMID:23325775

Neural Correlates of Natural Human Echolocation in Early and Late Blind Echolocation Experts

PubMed Central

Thaler, Lore; Arnott, Stephen R.; Goodale, Melvyn A.

2011-01-01

Background A small number of blind people are adept at echolocating silent objects simply by producing mouth clicks and listening to the returning echoes. Yet the neural architecture underlying this type of aid-free human echolocation has not been investigated. To tackle this question, we recruited echolocation experts, one early- and one late-blind, and measured functional brain activity in each of them while they listened to their own echolocation sounds. Results When we compared brain activity for sounds that contained both clicks and the returning echoes with brain activity for control sounds that did not contain the echoes, but were otherwise acoustically matched, we found activity in calcarine cortex in both individuals. Importantly, for the same comparison, we did not observe a difference in activity in auditory cortex. In the early-blind, but not the late-blind participant, we also found that the calcarine activity was greater for echoes reflected from surfaces located in contralateral space. Finally, in both individuals, we found activation in middle temporal and nearby cortical regions when they listened to echoes reflected from moving targets. Conclusions These findings suggest that processing of click-echoes recruits brain regions typically devoted to vision rather than audition in both early and late blind echolocation experts. PMID:21633496

Dolphins can maintain vigilant behavior through echolocation for 15 days without interruption or cognitive impairment.

PubMed

Branstetter, Brian K; Finneran, James J; Fletcher, Elizabeth A; Weisman, Brian C; Ridgway, Sam H

2012-01-01

In dolphins, natural selection has developed unihemispheric sleep where alternating hemispheres of their brain stay awake. This allows dolphins to maintain consciousness in response to respiratory demands of the ocean. Unihemispheric sleep may also allow dolphins to maintain vigilant states over long periods of time. Because of the relatively poor visibility in the ocean, dolphins use echolocation to interrogate their environment. During echolocation, dolphin produce clicks and listen to returning echoes to determine the location and identity of objects. The extent to which individual dolphins are able to maintain continuous vigilance through this active sense is unknown. Here we show that dolphins may continuously echolocate and accurately report the presence of targets for at least 15 days without interruption. During a total of three sessions, each lasting five days, two dolphins maintained echolocation behaviors while successfully detecting and reporting targets. Overall performance was between 75 to 86% correct for one dolphin and 97 to 99% correct for a second dolphin. Both animals demonstrated diel patterns in echolocation behavior. A 15-day testing session with one dolphin resulted in near perfect performance with no significant decrement over time. Our results demonstrate that dolphins can continuously monitor their environment and maintain long-term vigilant behavior through echolocation.

Evolutionary origins of ultrasonic hearing and laryngeal echolocation in bats inferred from morphological analyses of the inner ear

PubMed Central

2013-01-01

Introduction Many mammals have evolved highly adapted hearing associated with ecological specialisation. Of these, bats possess the widest frequency range of vocalisations and associated hearing sensitivities, with frequencies of above 200 kHz in some lineages that use laryngeal echolocation. High frequency hearing in bats appears to have evolved via structural modifications of the inner ear, however, studying these minute features presents considerable challenges and hitherto few such attempts have been made. To understand these adaptations more fully, as well as gain insights into the evolutionary origins of ultrasonic hearing and echolocation in bats, we undertook micro-computed tomography (μCT) scans of the cochleae of representative bat species from 16 families, encompassing their broad range of ecological diversity. To characterise cochlear gross morphology, we measured the relative basilar membrane length and number of turns, and compared these values between echolocating and non-echolocating bats, as well as other mammals. Results We found that hearing and echolocation call frequencies in bats correlated with both measures of cochlear morphology. In particular, relative basilar membrane length was typically longer in echolocating species, and also correlated positively with the number of cochlear turns. Ancestral reconstructions of these parameters suggested that the common ancestor of all extant bats was probably capable of ultrasonic hearing; however, we also found evidence of a significant decrease in the rate of morphological evolution of the basilar membrane in multiple ancestral branches within the Yangochiroptera suborder. Within the echolocating Yinpterochiroptera, there was some evidence of an increase in the rate of basilar membrane evolution in some tips of the tree, possibly associated with reported shifts in call frequency associated with recent speciation events. Conclusions The two main groups of echolocating bat were found to display highly variable inner ear morphologies. Ancestral reconstructions and rate shift analyses of ear morphology point to a complex evolutionary history, with the former supporting ultrasonic hearing in the common bat ancestor but the latter suggesting that morphological changes associated with echolocation might have occurred later. These findings are consistent with theories that sophisticated laryngeal echolocation, as seen in modern lineages, evolved following the divergence of the two main suborders. PMID:23360746

People's Ability to Detect Objects Using Click-Based Echolocation: A Direct Comparison between Mouth-Clicks and Clicks Made by a Loudspeaker.

PubMed

Thaler, Lore; Castillo-Serrano, Josefina

2016-01-01

Echolocation is the ability to use reflected sound to obtain information about the spatial environment. Echolocation is an active process that requires both the production of the emission as well as the sensory processing of the resultant sound. Appreciating the general usefulness of echo-acoustic cues for people, in particular those with vision impairments, various devices have been built that exploit the principle of echolocation to obtain and provide information about the environment. It is common to all these devices that they do not require the person to make a sound. Instead, the device produces the emission autonomously and feeds a resultant sound back to the user. Here we tested if echolocation performance in a simple object detection task was affected by the use of a head-mounted loudspeaker as compared to active clicking. We found that 27 sighted participants new to echolocation did generally better when they used a loudspeaker as compared to mouth-clicks, and that two blind participants with experience in echolocation did equally well with mouth clicks and the speaker. Importantly, performance of sighted participants' was not statistically different from performance of blind experts when they used the speaker. Based on acoustic click data collected from a subset of our participants, those participants whose mouth clicks were more similar to the speaker clicks, and thus had higher peak frequencies and sound intensity, did better. We conclude that our results are encouraging for the consideration and development of assistive devices that exploit the principle of echolocation.

People’s Ability to Detect Objects Using Click-Based Echolocation: A Direct Comparison between Mouth-Clicks and Clicks Made by a Loudspeaker

PubMed Central

Thaler, Lore; Castillo-Serrano, Josefina

2016-01-01

Echolocation is the ability to use reflected sound to obtain information about the spatial environment. Echolocation is an active process that requires both the production of the emission as well as the sensory processing of the resultant sound. Appreciating the general usefulness of echo-acoustic cues for people, in particular those with vision impairments, various devices have been built that exploit the principle of echolocation to obtain and provide information about the environment. It is common to all these devices that they do not require the person to make a sound. Instead, the device produces the emission autonomously and feeds a resultant sound back to the user. Here we tested if echolocation performance in a simple object detection task was affected by the use of a head-mounted loudspeaker as compared to active clicking. We found that 27 sighted participants new to echolocation did generally better when they used a loudspeaker as compared to mouth-clicks, and that two blind participants with experience in echolocation did equally well with mouth clicks and the speaker. Importantly, performance of sighted participants’ was not statistically different from performance of blind experts when they used the speaker. Based on acoustic click data collected from a subset of our participants, those participants whose mouth clicks were more similar to the speaker clicks, and thus had higher peak frequencies and sound intensity, did better. We conclude that our results are encouraging for the consideration and development of assistive devices that exploit the principle of echolocation. PMID:27135407

Female Mate Choice Can Drive the Evolution of High Frequency Echolocation in Bats: A Case Study with Rhinolophus mehelyi

PubMed Central

Puechmaille, Sébastien J.; Borissov, Ivailo M.; Zsebok, Sándor; Allegrini, Benjamin; Hizem, Mohammed; Kuenzel, Sven; Schuchmann, Maike; Teeling, Emma C.

2014-01-01

Animals employ an array of signals (i.e. visual, acoustic, olfactory) for communication. Natural selection favours signals, receptors, and signalling behaviour that optimise the received signal relative to background noise. When the signal is used for more than one function, antagonisms amongst the different signalling functions may constrain the optimisation of the signal for any one function. Sexual selection through mate choice can strongly modify the effects of natural selection on signalling systems ultimately causing maladaptive signals to evolve. Echolocating bats represent a fascinating group in which to study the evolution of signalling systems as unlike bird songs or frog calls, echolocation has a dual role in foraging and communication. The function of bat echolocation is to generate echoes that the calling bat uses for orientation and food detection with call characteristics being directly related to the exploitation of particular ecological niches. Therefore, it is commonly assumed that echolocation has been shaped by ecology via natural selection. Here we demonstrate for the first time using a novel combined behavioural, ecological and genetic approach that in a bat species, Rhinolophus mehelyi: (1) echolocation peak frequency is an honest signal of body size; (2) females preferentially select males with high frequency calls during the mating season; (3) high frequency males sire more off-spring, providing evidence that echolocation calls may play a role in female mate choice. Our data refute the sole role of ecology in the evolution of echolocation and highlight the antagonistic interplay between natural and sexual selection in shaping acoustic signals. PMID:25075972

Female mate choice can drive the evolution of high frequency echolocation in bats: a case study with Rhinolophus mehelyi.

PubMed

Puechmaille, Sébastien J; Borissov, Ivailo M; Zsebok, Sándor; Allegrini, Benjamin; Hizem, Mohammed; Kuenzel, Sven; Schuchmann, Maike; Teeling, Emma C; Siemers, Björn M

2014-01-01

Animals employ an array of signals (i.e. visual, acoustic, olfactory) for communication. Natural selection favours signals, receptors, and signalling behaviour that optimise the received signal relative to background noise. When the signal is used for more than one function, antagonisms amongst the different signalling functions may constrain the optimisation of the signal for any one function. Sexual selection through mate choice can strongly modify the effects of natural selection on signalling systems ultimately causing maladaptive signals to evolve. Echolocating bats represent a fascinating group in which to study the evolution of signalling systems as unlike bird songs or frog calls, echolocation has a dual role in foraging and communication. The function of bat echolocation is to generate echoes that the calling bat uses for orientation and food detection with call characteristics being directly related to the exploitation of particular ecological niches. Therefore, it is commonly assumed that echolocation has been shaped by ecology via natural selection. Here we demonstrate for the first time using a novel combined behavioural, ecological and genetic approach that in a bat species, Rhinolophus mehelyi: (1) echolocation peak frequency is an honest signal of body size; (2) females preferentially select males with high frequency calls during the mating season; (3) high frequency males sire more off-spring, providing evidence that echolocation calls may play a role in female mate choice. Our data refute the sole role of ecology in the evolution of echolocation and highlight the antagonistic interplay between natural and sexual selection in shaping acoustic signals.

Assessing bat detectability and occupancy with multiple automated echolocation detectors

Treesearch

Marcos P. Gorresen; Adam C. Miles; Christopher M. Todd; Frank J. Bonaccorso; Theodore J. Weller

2008-01-01

Occupancy analysis and its ability to account for differential detection probabilities is important for studies in which detecting echolocation calls is used as a measure of bat occurrence and activity. We examined the feasibility of remotely acquiring bat encounter histories to estimate detection probability and occupancy. We used echolocation detectors coupled o...

Interaction of vestibular, echolocation, and visual modalities guiding flight by the big brown bat, Eptesicus fuscus.

PubMed

Horowitz, Seth S; Cheney, Cheryl A; Simmons, James A

2004-01-01

The big brown bat (Eptesicus fuscus) is an aerial-feeding insectivorous species that relies on echolocation to avoid obstacles and to detect flying insects. Spatial perception in the dark using echolocation challenges the vestibular system to function without substantial visual input for orientation. IR thermal video recordings show the complexity of bat flights in the field and suggest a highly dynamic role for the vestibular system in orientation and flight control. To examine this role, we carried out laboratory studies of flight behavior under illuminated and dark conditions in both static and rotating obstacle tests while administering heavy water (D2O) to impair vestibular inputs. Eptesicus carried out complex maneuvers through both fixed arrays of wires and a rotating obstacle array using both vision and echolocation, or when guided by echolocation alone. When treated with D2O in combination with lack of visual cues, bats showed considerable decrements in performance. These data indicate that big brown bats use both vision and echolocation to provide spatial registration for head position information generated by the vestibular system.

The evolution of bat vestibular systems in the face of potential antagonistic selection pressures for flight and echolocation.

PubMed

Davies, Kalina T J; Bates, Paul J J; Maryanto, Ibnu; Cotton, James A; Rossiter, Stephen J

2013-01-01

The vestibular system maintains the body's sense of balance and, therefore, was probably subject to strong selection during evolutionary transitions in locomotion. Among mammals, bats possess unique traits that place unusual demands on their vestibular systems. First, bats are capable of powered flight, which in birds is associated with enlarged semicircular canals. Second, many bats have enlarged cochleae associated with echolocation, and both cochleae and semicircular canals share a space within the petrosal bone. To determine how bat vestibular systems have evolved in the face of these pressures, we used micro-CT scans to compare canal morphology across species with contrasting flight and echolocation capabilities. We found no increase in canal radius in bats associated with the acquisition of powered flight, but canal radius did correlate with body mass in bat species from the suborder Yangochiroptera, and also in non-echolocating Old World fruit bats from the suborder Yinpterochiroptera. No such trend was seen in members of the Yinpterochiroptera that use laryngeal echolocation, although canal radius was associated with wing-tip roundedness in this group. We also found that the vestibular system scaled with cochlea size, although the relationship differed in species that use constant frequency echolocation. Across all bats, the shape of the anterior and lateral canals was associated with large cochlea size and small body size respectively, suggesting differential spatial constraints on each canal depending on its orientation within the skull. Thus in many echolocating bats, it seems that the combination of small body size and enlarged cochlea together act as a principal force on the vestibular system. The two main groups of echolocating bats displayed different canal morphologies, in terms of size and shape in relation to body mass and cochlear size, thus suggesting independent evolutionary pathways and offering tentative support for multiple acquisitions of echolocation.

The Evolution of Bat Vestibular Systems in the Face of Potential Antagonistic Selection Pressures for Flight and Echolocation

PubMed Central

Davies, Kalina T. J.; Bates, Paul J. J.; Maryanto, Ibnu; Cotton, James A.; Rossiter, Stephen J.

2013-01-01

The vestibular system maintains the body’s sense of balance and, therefore, was probably subject to strong selection during evolutionary transitions in locomotion. Among mammals, bats possess unique traits that place unusual demands on their vestibular systems. First, bats are capable of powered flight, which in birds is associated with enlarged semicircular canals. Second, many bats have enlarged cochleae associated with echolocation, and both cochleae and semicircular canals share a space within the petrosal bone. To determine how bat vestibular systems have evolved in the face of these pressures, we used micro-CT scans to compare canal morphology across species with contrasting flight and echolocation capabilities. We found no increase in canal radius in bats associated with the acquisition of powered flight, but canal radius did correlate with body mass in bat species from the suborder Yangochiroptera, and also in non-echolocating Old World fruit bats from the suborder Yinpterochiroptera. No such trend was seen in members of the Yinpterochiroptera that use laryngeal echolocation, although canal radius was associated with wing-tip roundedness in this group. We also found that the vestibular system scaled with cochlea size, although the relationship differed in species that use constant frequency echolocation. Across all bats, the shape of the anterior and lateral canals was associated with large cochlea size and small body size respectively, suggesting differential spatial constraints on each canal depending on its orientation within the skull. Thus in many echolocating bats, it seems that the combination of small body size and enlarged cochlea together act as a principal force on the vestibular system. The two main groups of echolocating bats displayed different canal morphologies, in terms of size and shape in relation to body mass and cochlear size, thus suggesting independent evolutionary pathways and offering tentative support for multiple acquisitions of echolocation. PMID:23637943

Early milestones in the understanding of echolocation in bats.

PubMed

Grinnell, Alan D

2018-04-23

Almost 80 years ago, Griffin and Galambos discovered the phenomenon of echolocation in bats. Since then, the field has grown exponentially as new generations of investigators have joined the chase and technological advances have revolutionized working with ultrasound in the laboratory and in the field. Today our understanding of the diversity of behavioral and neural adaptations for echolocation constitutes one of the paramount triumphs of neuroethology. At the invitation of the editor in chief, I here review some of the important milestones in the discovery and early understanding of echolocation in bats through about the mid-1980s.

Postnatal ontogeny of the cochlea and flight ability in Jamaican fruit bats (Phyllostomidae) with implications for the evolution of echolocation.

PubMed

Carter, Richard T; Adams, Rick A

2015-04-01

Recent evidence has shown that the developmental emergence of echolocation calls in young bats follow an independent developmental pathway from other vocalizations and that adult-like echolocation call structure significantly precedes flight ability. These data in combination with new insights into the echolocation ability of some shrews suggest that the evolution of echolocation in bats may involve inheritance of a primitive sonar system that was modified to its current state, rather than the ad hoc evolution of echolocation in the earliest bats. Because the cochlea is crucial in the sensation of echoes returning from sonar pulses, we tracked changes in cochlear morphology during development that included the basilar membrane (BM) and secondary spiral lamina (SSL) along the length of the cochlea in relation to stages of flight ability in young bats. Our data show that the morphological prerequisite for sonar sensitivity of the cochlea significantly precedes the onset of flight in young bats and, in fact, development of this prerequisite is complete before parturition. In addition, there were no discernible changes in cochlear morphology with stages of flight development, demonstrating temporal asymmetry between the development of morphology associated with echo-pulse return sensitivity and volancy. These data further corroborate and support the hypothesis that adaptations for sonar and echolocation evolved before flight in mammals. © 2015 Anatomical Society.

Echolocation versus echo suppression in humans

PubMed Central

Wallmeier, Ludwig; Geßele, Nikodemus; Wiegrebe, Lutz

2013-01-01

Several studies have shown that blind humans can gather spatial information through echolocation. However, when localizing sound sources, the precedence effect suppresses spatial information of echoes, and thereby conflicts with effective echolocation. This study investigates the interaction of echolocation and echo suppression in terms of discrimination suppression in virtual acoustic space. In the ‘Listening’ experiment, sighted subjects discriminated between positions of a single sound source, the leading or the lagging of two sources, respectively. In the ‘Echolocation’ experiment, the sources were replaced by reflectors. Here, the same subjects evaluated echoes generated in real time from self-produced vocalizations and thereby discriminated between positions of a single reflector, the leading or the lagging of two reflectors, respectively. Two key results were observed. First, sighted subjects can learn to discriminate positions of reflective surfaces echo-acoustically with accuracy comparable to sound source discrimination. Second, in the Listening experiment, the presence of the leading source affected discrimination of lagging sources much more than vice versa. In the Echolocation experiment, however, the presence of both the lead and the lag strongly affected discrimination. These data show that the classically described asymmetry in the perception of leading and lagging sounds is strongly diminished in an echolocation task. Additional control experiments showed that the effect is owing to both the direct sound of the vocalization that precedes the echoes and owing to the fact that the subjects actively vocalize in the echolocation task. PMID:23986105

The Effects of Attenuating Returning Echolocation Signals at the Lower Jaw of a Dolphin (Tursiops Truncatus)

DTIC Science & Technology

1991-06-01

echolocation signals are guided to the inner ear of odonto - relatively little loss of energy. The biochemical composition cete cetaceans via areas of fatty...pool, on the range of useful al. (1986), and others have provided evidence that odonto - energy in the emitted clicks. cete cetaceans emit echolocation

Parallel evolution of auditory genes for echolocation in bats and toothed whales.

PubMed

Shen, Yong-Yi; Liang, Lu; Li, Gui-Sheng; Murphy, Robert W; Zhang, Ya-Ping

2012-06-01

The ability of bats and toothed whales to echolocate is a remarkable case of convergent evolution. Previous genetic studies have documented parallel evolution of nucleotide sequences in Prestin and KCNQ4, both of which are associated with voltage motility during the cochlear amplification of signals. Echolocation involves complex mechanisms. The most important factors include cochlear amplification, nerve transmission, and signal re-coding. Herein, we screen three genes that play different roles in this auditory system. Cadherin 23 (Cdh23) and its ligand, protocadherin 15 (Pcdh15), are essential for bundling motility in the sensory hair. Otoferlin (Otof) responds to nerve signal transmission in the auditory inner hair cell. Signals of parallel evolution occur in all three genes in the three groups of echolocators--two groups of bats (Yangochiroptera and Rhinolophoidea) plus the dolphin. Significant signals of positive selection also occur in Cdh23 in the Rhinolophoidea and dolphin, and Pcdh15 in Yangochiroptera. In addition, adult echolocating bats have higher levels of Otof expression in the auditory cortex than do their embryos and non-echolocation bats. Cdh23 and Pcdh15 encode the upper and lower parts of tip-links, and both genes show signals of convergent evolution and positive selection in echolocators, implying that they may co-evolve to optimize cochlear amplification. Convergent evolution and expression patterns of Otof suggest the potential role of nerve and brain in echolocation. Our synthesis of gene sequence and gene expression analyses reveals that positive selection, parallel evolution, and perhaps co-evolution and gene expression affect multiple hearing genes that play different roles in audition, including voltage and bundle motility in cochlear amplification, nerve transmission, and brain function.

The influence of feeding on the evolution of sensory signals: a comparative test of an evolutionary trade-off between masticatory and sensory functions of skulls in southern African horseshoe bats (Rhinolophidae).

PubMed

Jacobs, D S; Bastian, A; Bam, L

2014-12-01

The skulls of animals have to perform many functions. Optimization for one function may mean another function is less optimized, resulting in evolutionary trade-offs. Here, we investigate whether a trade-off exists between the masticatory and sensory functions of animal skulls using echolocating bats as model species. Several species of rhinolophid bats deviate from the allometric relationship between body size and echolocation frequency. Such deviation may be the result of selection for increased bite force, resulting in a decrease in snout length which could in turn lead to higher echolocation frequencies. If so, there should be a positive relationship between bite force and echolocation frequency. We investigated this relationship in several species of southern African rhinolophids using phylogenetically informed analyses of the allometry of their bite force and echolocation frequency and of the three-dimensional shape of their skulls. As predicted, echolocation frequency was positively correlated with bite force, suggesting that its evolution is influenced by a trade-off between the masticatory and sensory functions of the skull. In support of this, variation in skull shape was explained by both echolocation frequency (80%) and bite force (20%). Furthermore, it appears that selection has acted on the nasal capsules, which have a frequency-specific impedance matching function during vocalization. There was a negative correlation between echolocation frequency and capsule volume across species. Optimization of the masticatory function of the skull may have been achieved through changes in the shape of the mandible and associated musculature, elements not considered in this study. © 2014 European Society For Evolutionary Biology. Journal of Evolutionary Biology © 2014 European Society For Evolutionary Biology.

Active echolocation beam focusing in the false killer whale, Pseudorca crassidens.

PubMed

Kloepper, Laura N; Nachtigall, Paul E; Donahue, Megan J; Breese, Marlee

2012-04-15

The odontocete sound production system is highly complex and produces intense, directional signals that are thought to be focused by the melon and the air sacs. Because odontocete echolocation signals are variable and the emitted click frequency greatly affects the echolocation beam shape, investigations of beam focusing must account for frequency-related beam changes. In this study we tested whether the echolocation beam of a false killer whale changed depending on target difficulty and distance while also accounting for frequency-related changes in the echolocation beam. The data indicate that the false killer whale changes its beam size according to target distance and difficulty, which may be a strategy of maximizing the energy of the target echo. We propose that the animal is using a strategy of changing the focal region according to target distance and that this strategy is under active control.

Echolocation behavior of franciscana dolphins (Pontoporia blainvillei) in the wild.

PubMed

Melcón, Mariana L; Failla, Mauricio; Iñíguez, Miguel A

2012-06-01

Franciscana dolphins are small odontocetes hard to study in the field. In particular, little is known on their echolocation behavior in the wild. In this study we recorded 357 min and analyzed 1019 echolocation signals in the Rio Negro Estuary, Argentina. The clicks had a peak frequency at 139 kHz, and a bandwidth of 19 kHz, ranging from 130 to 149 kHz. This is the first study describing echolocation signals of franciscana dolphins in the wild, showing the presence of narrow-band high frequency signals in these dolphins. Whether they use other vocalizations to communicate or not remains uncertain.

The size-weight illusion induced through human echolocation.

PubMed

Buckingham, Gavin; Milne, Jennifer L; Byrne, Caitlin M; Goodale, Melvyn A

2015-02-01

Certain blind individuals have learned to interpret the echoes of self-generated sounds to perceive the structure of objects in their environment. The current work examined how far the influence of this unique form of sensory substitution extends by testing whether echolocation-induced representations of object size could influence weight perception. A small group of echolocation experts made tongue clicks or finger snaps toward cubes of varying sizes and weights before lifting them. These echolocators experienced a robust size-weight illusion. This experiment provides the first demonstration of a sensory substitution technique whereby the substituted sense influences the conscious perception through an intact sense. © The Author(s) 2014.

An aerial-hawking bat uses stealth echolocation to counter moth hearing.

PubMed

Goerlitz, Holger R; ter Hofstede, Hannah M; Zeale, Matt R K; Jones, Gareth; Holderied, Marc W

2010-09-14

Ears evolved in many nocturnal insects, including some moths, to detect bat echolocation calls and evade capture [1, 2]. Although there is evidence that some bats emit echolocation calls that are inconspicuous to eared moths, it is difficult to determine whether this was an adaptation to moth hearing or originally evolved for a different purpose [2, 3]. Aerial-hawking bats generally emit high-amplitude echolocation calls to maximize detection range [4, 5]. Here we present the first example of an echolocation counterstrategy to overcome prey hearing at the cost of reduced detection distance. We combined comparative bat flight-path tracking and moth neurophysiology with fecal DNA analysis to show that the barbastelle, Barbastella barbastellus, emits calls that are 10 to 100 times lower in amplitude than those of other aerial-hawking bats, remains undetected by moths until close, and captures mainly eared moths. Model calculations demonstrate that only bats emitting such low-amplitude calls hear moth echoes before their calls are conspicuous to moths. This stealth echolocation allows the barbastelle to exploit food resources that are difficult to catch for other aerial-hawking bats emitting calls of greater amplitude. Copyright © 2010 Elsevier Ltd. All rights reserved.

Nonecholocating fruit bats produce biosonar clicks with their wings.

PubMed

Boonman, Arjan; Bumrungsri, Sara; Yovel, Yossi

2014-12-15

Because evolution mostly acts over millions of years, the intermediate steps leading to a functional sensory system remain enigmatic. Accordingly, there is an ongoing debate regarding the evolution of bat echolocation. In search of the origin of bat echolocation, we studied how Old World fruit bats, which have always been classified as nonecholocating, orient in complete darkness. We found that two of these nonecholocating species used click-like sounds to detect and discriminate objects in complete darkness. However, we discovered that this click-based echo sensing is rudimentary and does not allow these bats to estimate distance accurately as all other echolocating bats can. Moreover, unlike all other echolocating bats, which generate pulses using the larynx or the tongue, these bats generated clicks with their wings. We provide evidence suggesting that all Old World fruit bats can click with their wings. Although this click-based echo sensing used by Old World fruit bats may not represent the ancestral form of current (laryngeal) bat echolocation, we argue that clicking fruit bats could be considered behavioral fossils, opening a window to study the evolution of echolocation. Copyright © 2014 Elsevier Ltd. All rights reserved.

Postnatal development of echolocation abilities in a bottlenose dolphin (Tursiops truncatus): temporal organization.

PubMed

Favaro, Livio; Gnone, Guido; Pessani, Daniela

2013-03-01

In spite of all the information available on adult bottlenose dolphin (Tursiops truncatus) biosonar, the ontogeny of its echolocation abilities has been investigated very little. Earlier studies have reported that neonatal dolphins can produce both whistles and burst-pulsed sounds just after birth and that early-pulsed sounds are probably a precursor of echolocation click trains. The aim of this research is to investigate the development of echolocation signals in a captive calf, born in the facilities of the Acquario di Genova. A set of 81 impulsive sounds were collected from birth to the seventh postnatal week and six additional echolocation click trains were recorded when the dolphin was 1 year old. Moreover, behavioral observations, concurring with sound production, were carried out by means of a video camera. For each sound we measured five acoustic parameters: click train duration (CTD), number of clicks per train, minimum, maximum, and mean click repetition rate (CRR). CTD and number of clicks per train were found to increase with age. Maximum and mean CRR followed a decreasing trend with dolphin growth starting from the second postnatal week. The calf's first head scanning movement was recorded 21 days after birth. Our data suggest that in the bottlenose dolphin the early postnatal weeks are essential for the development of echolocation abilities and that the temporal features of the echolocation click trains remain relatively stable from the seventh postnatal week up to the first year of life. © 2013 Wiley Periodicals, Inc.

Parahippocampal cortex is involved in material processing via echoes in blind echolocation experts.

PubMed

Milne, Jennifer L; Arnott, Stephen R; Kish, Daniel; Goodale, Melvyn A; Thaler, Lore

2015-04-01

Some blind humans use sound to navigate by emitting mouth-clicks and listening to the echoes that reflect from silent objects and surfaces in their surroundings. These echoes contain information about the size, shape, location, and material properties of objects. Here we present results from an fMRI experiment that investigated the neural activity underlying the processing of materials through echolocation. Three blind echolocation experts (as well as three blind and three sighted non-echolocating control participants) took part in the experiment. First, we made binaural sound recordings in the ears of each echolocator while he produced clicks in the presence of one of three different materials (fleece, synthetic foliage, or whiteboard), or while he made clicks in an empty room. During fMRI scanning these recordings were played back to participants. Remarkably, all participants were able to identify each of the three materials reliably, as well as the empty room. Furthermore, a whole brain analysis, in which we isolated the processing of just the reflected echoes, revealed a material-related increase in BOLD activation in a region of left parahippocampal cortex in the echolocating participants, but not in the blind or sighted control participants. Our results, in combination with previous findings about brain areas involved in material processing, are consistent with the idea that material processing by means of echolocation relies on a multi-modal material processing area in parahippocampal cortex. Copyright © 2014 Elsevier Ltd. All rights reserved.

Neurophysiological findings relevant to echolocation in marine animals

NASA Technical Reports Server (NTRS)

Bullock, T. H.; Ridgway, S. H.

1972-01-01

A review of echolocation mechanisms in marine mammals, chiefly porpoises, is given. Data cover peripheral auditory and central neurophysiological specializations favorable to the analysis of echolocating clicks and their echoes. Conclusions show (1) signals are received from 50 up to at least 135 kHz, (2) sound is received through the mandible skin, and (3) the midbrain sites are insensitive to low frequencies (below 6 kHz).

Blindness enhances auditory obstacle circumvention: Assessing echolocation, sensory substitution, and visual-based navigation

PubMed Central

Scarfe, Amy C.; Moore, Brian C. J.; Pardhan, Shahina

2017-01-01

Performance for an obstacle circumvention task was assessed under conditions of visual, auditory only (using echolocation) and tactile (using a sensory substitution device, SSD) guidance. A Vicon motion capture system was used to measure human movement kinematics objectively. Ten normally sighted participants, 8 blind non-echolocators, and 1 blind expert echolocator navigated around a 0.6 x 2 m obstacle that was varied in position across trials, at the midline of the participant or 25 cm to the right or left. Although visual guidance was the most effective, participants successfully circumvented the obstacle in the majority of trials under auditory or SSD guidance. Using audition, blind non-echolocators navigated more effectively than blindfolded sighted individuals with fewer collisions, lower movement times, fewer velocity corrections and greater obstacle detection ranges. The blind expert echolocator displayed performance similar to or better than that for the other groups using audition, but was comparable to that for the other groups using the SSD. The generally better performance of blind than of sighted participants is consistent with the perceptual enhancement hypothesis that individuals with severe visual deficits develop improved auditory abilities to compensate for visual loss, here shown by faster, more fluid, and more accurate navigation around obstacles using sound. PMID:28407000

A summary of research investigating echolocation abilities of blind and sighted humans.

PubMed

Kolarik, Andrew J; Cirstea, Silvia; Pardhan, Shahina; Moore, Brian C J

2014-04-01

There is currently considerable interest in the consequences of loss in one sensory modality on the remaining senses. Much of this work has focused on the development of enhanced auditory abilities among blind individuals, who are often able to use sound to navigate through space. It has now been established that many blind individuals produce sound emissions and use the returning echoes to provide them with information about objects in their surroundings, in a similar manner to bats navigating in the dark. In this review, we summarize current knowledge regarding human echolocation. Some blind individuals develop remarkable echolocation abilities, and are able to assess the position, size, distance, shape, and material of objects using reflected sound waves. After training, normally sighted people are also able to use echolocation to perceive objects, and can develop abilities comparable to, but typically somewhat poorer than, those of blind people. The underlying cues and mechanisms, operable range, spatial acuity and neurological underpinnings of echolocation are described. Echolocation can result in functional real life benefits. It is possible that these benefits can be optimized via suitable training, especially among those with recently acquired blindness, but this requires further study. Areas for further research are identified. Copyright © 2014 Elsevier B.V. All rights reserved.

Echolocation signals of foraging killer whales (Orcinus orca)

NASA Astrophysics Data System (ADS)

Au, Whitlow W. L.; Ford, John K. B.; Allman, Kelly A.

2002-05-01

Fish eating resident killer whales that frequent the coastal waters of Vancouver Island, Canada have a strong preference for chinook salmon. The whales in Johnston Strait often forage along the steep cliffs that extend into the water, echolocating their prey. Echolocation signals were measured with a four hydrophone symmetrical star array and the signals were simultaneous digitized at a sample rate of 500 kHz using a lunch-box PC. A portable VCR recorded the images from an underwater camera located close to the array center. Only signals emanated from close to the beam axis (1185 total) were chosen for a detailed analysis. Killer whales project very broad band echolocation signals (Q 1.3 to 1.5) that tend to have a bimodal frequency structure. Ninety seven percent of the signals had center frequencies between 45 and 80 kHz with a band-width between 35 and 50 kHz. The peak-to-peak source level of the echolocation signal decreased as a function of the one way transmission loss to the array. Source levels varied between 200 and 225 dB re 1 μPa. Using a model of target strength for chinook salmons, the echo levels from the echolocation signals are estimated for different ranges between whale and salmon.

Blindness enhances auditory obstacle circumvention: Assessing echolocation, sensory substitution, and visual-based navigation.

PubMed

Kolarik, Andrew J; Scarfe, Amy C; Moore, Brian C J; Pardhan, Shahina

2017-01-01

Performance for an obstacle circumvention task was assessed under conditions of visual, auditory only (using echolocation) and tactile (using a sensory substitution device, SSD) guidance. A Vicon motion capture system was used to measure human movement kinematics objectively. Ten normally sighted participants, 8 blind non-echolocators, and 1 blind expert echolocator navigated around a 0.6 x 2 m obstacle that was varied in position across trials, at the midline of the participant or 25 cm to the right or left. Although visual guidance was the most effective, participants successfully circumvented the obstacle in the majority of trials under auditory or SSD guidance. Using audition, blind non-echolocators navigated more effectively than blindfolded sighted individuals with fewer collisions, lower movement times, fewer velocity corrections and greater obstacle detection ranges. The blind expert echolocator displayed performance similar to or better than that for the other groups using audition, but was comparable to that for the other groups using the SSD. The generally better performance of blind than of sighted participants is consistent with the perceptual enhancement hypothesis that individuals with severe visual deficits develop improved auditory abilities to compensate for visual loss, here shown by faster, more fluid, and more accurate navigation around obstacles using sound.

Echolocation call intensity and directionality in flying short-tailed fruit bats, Carollia perspicillata (Phyllostomidae).

PubMed

Brinkløv, Signe; Jakobsen, Lasse; Ratcliffe, John M; Kalko, Elisabeth K V; Surlykke, Annemarie

2011-01-01

The directionality of bat echolocation calls defines the width of bats' sonar "view," while call intensity directly influences detection range since adequate sound energy must impinge upon objects to return audible echoes. Both are thus crucial parameters for understanding biosonar signal design. Phyllostomid bats have been classified as low intensity or "whispering bats," but recent data indicate that this designation may be inaccurate. Echolocation beam directionality in phyllostomids has only been measured through electrode brain-stimulation of restrained bats, presumably excluding active beam control via the noseleaf. Here, a 12-microphone array was used to measure echolocation call intensity and beam directionality in the frugivorous phyllostomid, Carollia perspicillata, echolocating in flight. The results showed a considerably narrower beam shape (half-amplitude beam angles of approximately 16° horizontally and 14° vertically) and louder echolocation calls [source levels averaging 99 dB sound pressure level (SPL) root mean square] for C. perspicillata than was found for this species when stationary. This suggests that naturally behaving phyllostomids shape their sound beam to achieve a longer and narrower sonar range than previously thought. C. perspicillata orient and forage in the forest interior and the narrow beam might be adaptive in clutter, by reducing the number and intensity of off-axis echoes.

Processing of Natural Echolocation Sequences in the Inferior Colliculus of Seba’s Fruit Eating Bat, Carollia perspicillata

PubMed Central

Kordes, Sebastian; Kössl, Manfred

2017-01-01

Abstract For the purpose of orientation, echolocating bats emit highly repetitive and spatially directed sonar calls. Echoes arising from call reflections are used to create an acoustic image of the environment. The inferior colliculus (IC) represents an important auditory stage for initial processing of echolocation signals. The present study addresses the following questions: (1) how does the temporal context of an echolocation sequence mimicking an approach flight of an animal affect neuronal processing of distance information to echo delays? (2) how does the IC process complex echolocation sequences containing echo information from multiple objects (multiobject sequence)? Here, we conducted neurophysiological recordings from the IC of ketamine-anaesthetized bats of the species Carollia perspicillata and compared the results from the IC with the ones from the auditory cortex (AC). Neuronal responses to an echolocation sequence was suppressed when compared to the responses to temporally isolated and randomized segments of the sequence. The neuronal suppression was weaker in the IC than in the AC. In contrast to the cortex, the time course of the acoustic events is reflected by IC activity. In the IC, suppression sharpens the neuronal tuning to specific call-echo elements and increases the signal-to-noise ratio in the units’ responses. When presenting multiple-object sequences, despite collicular suppression, the neurons responded to each object-specific echo. The latter allows parallel processing of multiple echolocation streams at the IC level. Altogether, our data suggests that temporally-precise neuronal responses in the IC could allow fast and parallel processing of multiple acoustic streams. PMID:29242823

Environmental acoustic cues guide the biosonar attention of a highly specialised echolocator.

PubMed

Lattenkamp, Ella Z; Kaiser, Samuel; Kaučič, Rožle; Großmann, Martina; Koselj, Klemen; Goerlitz, Holger R

2018-04-23

Sensory systems experience a trade-off between maximizing the detail and amount of sampled information. This trade-off is particularly pronounced in sensory systems that are highly specialised for a single task and thus experience limitations in other tasks. We hypothesised that combining sensory input from multiple streams of information may resolve this trade-off and improve detection and sensing reliability. Specifically, we predicted that perceptive limitations experienced by animals reliant on specialised active echolocation can be compensated for by the phylogenetically older and less specialised process of passive hearing. We tested this hypothesis in greater horseshoe bats, which possess morphological and neural specialisations allowing them to identify fluttering prey in dense vegetation using echolocation only. At the same time, their echolocation system is both spatially and temporally severely limited. Here, we show that greater horseshoe bats employ passive hearing to initially detect and localise prey-generated and other environmental sounds, and then raise vocalisation level and concentrate the scanning movements of their sonar beam on the sound source for further investigation with echolocation. These specialised echolocators thus supplement echo-acoustic information with environmental acoustic cues, enlarging perceived space beyond their biosonar range. Contrary to our predictions, we did not find consistent preferences for prey-related acoustic stimuli, indicating the use of passive acoustic cues also for detection of non-prey objects. Our findings suggest that even specialised echolocators exploit a wide range of environmental information, and that phylogenetically older sensory systems can support the evolution of sensory specialisations by compensating for their limitations. © 2018. Published by The Company of Biologists Ltd.

Processing of Natural Echolocation Sequences in the Inferior Colliculus of Seba's Fruit Eating Bat, Carollia perspicillata.

PubMed

Beetz, M Jerome; Kordes, Sebastian; García-Rosales, Francisco; Kössl, Manfred; Hechavarría, Julio C

2017-01-01

For the purpose of orientation, echolocating bats emit highly repetitive and spatially directed sonar calls. Echoes arising from call reflections are used to create an acoustic image of the environment. The inferior colliculus (IC) represents an important auditory stage for initial processing of echolocation signals. The present study addresses the following questions: (1) how does the temporal context of an echolocation sequence mimicking an approach flight of an animal affect neuronal processing of distance information to echo delays? (2) how does the IC process complex echolocation sequences containing echo information from multiple objects (multiobject sequence)? Here, we conducted neurophysiological recordings from the IC of ketamine-anaesthetized bats of the species Carollia perspicillata and compared the results from the IC with the ones from the auditory cortex (AC). Neuronal responses to an echolocation sequence was suppressed when compared to the responses to temporally isolated and randomized segments of the sequence. The neuronal suppression was weaker in the IC than in the AC. In contrast to the cortex, the time course of the acoustic events is reflected by IC activity. In the IC, suppression sharpens the neuronal tuning to specific call-echo elements and increases the signal-to-noise ratio in the units' responses. When presenting multiple-object sequences, despite collicular suppression, the neurons responded to each object-specific echo. The latter allows parallel processing of multiple echolocation streams at the IC level. Altogether, our data suggests that temporally-precise neuronal responses in the IC could allow fast and parallel processing of multiple acoustic streams.

Finding flowers in the dark: nectar-feeding bats integrate olfaction and echolocation while foraging for nectar

PubMed Central

Gonzalez-Terrazas, Tania P.; Martel, Carlos; Milet-Pinheiro, Paulo; Ayasse, Manfred; Kalko, Elisabeth K. V.; Tschapka, Marco

2016-01-01

Nectar-feeding bats depend mainly on floral nectar to fulfil their energetic requirements. Chiropterophilous flowers generally present strong floral scents and provide conspicuous acoustic echoes to attract bats. While floral scents are assumed to attract bats over long distances, acoustic properties of flower structures may provide detailed information, thus supporting the localization of a flower at close ranges. So far, to our knowledge, there is no study trying to understand the relative importance as well as the combination of these generally coupled cues for detection (presence) and localization (exact position) of open flowers in nature. For a better comprehension of the significance of olfaction and echolocation in the foraging behaviour of nectar-feeding bats, we conducted two-choice experiments with Leptonycteris yerbabuenae. We tested the bats' behaviour in three experimental scenarios with different cues: (i) olfaction versus echolocation, (ii) echolocation versus echolocation and olfaction, and (iii) olfaction versus echolocation and olfaction. We used the floral scent of the bat-pollinated cactus Pachycereus pringlei as olfactory cue and an acrylic paraboloid as acoustic cue. Additionally, we recorded the echolocation behaviour of the bats and analysed the floral scent of P. pringlei. When decoupled cues were offered, bats displayed no preference in choice for any of the two cues. However, bats reacted first to and chose more often the coupled cues. All bats echolocated continuously and broadcast a long terminal group before a successful visit. The floral scent bouquet of P. pringlei is composed of 20 compounds, some of which (e.g. methyl benzoate) were already reported from chiropterophilous plants. Our investigation demonstrates for the first time to our knowledge, that nectar-feeding bats integrate over different sensory modes for detection and precise localization of open flowers. The combined information from olfactory and acoustic cues allows bats to forage more efficiently. PMID:27853595

Spike Neuromorphic VLSI-Based Bat Echolocation for Micro-Aerial Vehicle Guidance

DTIC Science & Technology

2007-03-31

IFinal 03/01/04 - 02/28/07 4. TITLE AND SUBTITLE 5a. CONTRACT NUMBER Neuromorphic VLSI-based Bat Echolocation for Micro-aerial 5b.GRANTNUMBER Vehicle...uncovered interesting new issues in our choice for representing the intensity of signals. We have just finished testing the first chip version of an echo...timing-based algorithm (’openspace’) for sonar-guided navigation amidst multiple obstacles. 15. SUBJECT TERMS Neuromorphic VLSI, bat echolocation

Single-click beam patterns suggest dynamic changes to the field of view of echolocating Atlantic spotted dolphins (Stenella frontalis) in the wild.

PubMed

Jensen, Frants H; Wahlberg, Magnus; Beedholm, Kristian; Johnson, Mark; de Soto, Natacha Aguilar; Madsen, Peter T

2015-05-01

Echolocating animals exercise an extensive control over the spectral and temporal properties of their biosonar signals to facilitate perception of their actively generated auditory scene when homing in on prey. The intensity and directionality of the biosonar beam defines the field of view of echolocating animals by affecting the acoustic detection range and angular coverage. However, the spatial relationship between an echolocating predator and its prey changes rapidly, resulting in different biosonar requirements throughout prey pursuit and capture. Here, we measured single-click beam patterns using a parametric fit procedure to test whether free-ranging Atlantic spotted dolphins (Stenella frontalis) modify their biosonar beam width. We recorded echolocation clicks using a linear array of receivers and estimated the beam width of individual clicks using a parametric spectral fit, cross-validated with well-established composite beam pattern estimates. The dolphins apparently increased the biosonar beam width, to a large degree without changing the signal frequency, when they approached the recording array. This is comparable to bats that also expand their field of view during prey capture, but achieve this by decreasing biosonar frequency. This behaviour may serve to decrease the risk that rapid escape movements of prey take them outside the biosonar beam of the predator. It is likely that shared sensory requirements have resulted in bats and toothed whales expanding their acoustic field of view at close range to increase the likelihood of successfully acquiring prey using echolocation, representing a case of convergent evolution of echolocation behaviour between these two taxa. © 2015. Published by The Company of Biologists Ltd.

Oilbirds produce echolocation signals beyond their best hearing range and adjust signal design to natural light conditions

PubMed Central

Brinkløv, Signe; Elemans, Coen P. H.

2017-01-01

Oilbirds are active at night, foraging for fruits using keen olfaction and extremely light-sensitive eyes, and echolocate as they leave and return to their cavernous roosts. We recorded the echolocation behaviour of wild oilbirds using a multi-microphone array as they entered and exited their roosts under different natural light conditions. During echolocation, the birds produced click bursts (CBs) lasting less than 10 ms and consisting of a variable number (2–8) of clicks at 2–3 ms intervals. The CBs have a bandwidth of 7–23 kHz at −6 dB from signal peak frequency. We report on two unique characteristics of this avian echolocation system. First, oilbirds reduce both the energy and number of clicks in their CBs under conditions of clear, moonlit skies, compared with dark, moonless nights. Second, we document a frequency mismatch between the reported best frequency of oilbird hearing (approx. 2 kHz) and the bandwidth of their echolocation CBs. This unusual signal-to-sensory system mismatch probably reflects avian constraints on high-frequency hearing but may still allow oilbirds fine-scale, close-range detail resolution at the upper extreme (approx. 10 kHz) of their presumed hearing range. Alternatively, oilbirds, by an as-yet unknown mechanism, are able to hear frequencies higher than currently appreciated. PMID:28573036

Directionality of nose-emitted echolocation calls from bats without a nose leaf (Plecotus auritus).

PubMed

Jakobsen, Lasse; Hallam, John; Moss, Cynthia F; Hedenström, Anders

2018-02-13

All echolocating bats and whales measured to date emit a directional bio-sonar beam that affords them a number of advantages over an omni-directional beam, i.e. reduced clutter, increased source level and inherent directional information. In this study, we investigated the importance of directional sound emission for navigation through echolocation by measuring the sonar beam of brown long-eared bats, Plecotus auritus Plecotus auritus emits sound through the nostrils but has no external appendages to readily facilitate a directional sound emission as found in most nose emitters. The study shows that P. auritus , despite lacking an external focusing apparatus, emits a directional echolocation beam (directivity index=13 dB) and that the beam is more directional vertically (-6 dB angle at 22 deg) than horizontally (-6 dB angle at 35 deg). Using a simple numerical model, we found that the recorded emission pattern is achievable if P. auritus emits sound through the nostrils as well as the mouth. The study thus supports the hypothesis that a directional echolocation beam is important for perception through echolocation and we propose that animals with similarly non-directional emitter characteristics may facilitate a directional sound emission by emitting sound through both the nostrils and the mouth. © 2018. Published by The Company of Biologists Ltd.

'Compromise' in Echolocation Calls between Different Colonies of the Intermediate Leaf-Nosed Bat (Hipposideros larvatus).

PubMed

Chen, Yi; Liu, Qi; Su, Qianqian; Sun, Yunxiao; Peng, Xingwen; He, Xiangyang; Zhang, Libiao

2016-01-01

Each animal population has its own acoustic signature which facilitates identification, communication and reproduction. The sonar signals of bats can convey social information, such as species identity and contextual information. The goal of this study was to determine whether bats adjust their echolocation call structures to mutually recognize and communicate when they encounter the bats from different colonies. We used the intermediate leaf-nosed bats (Hipposideros larvatus) as a case study to investigate the variations of echolocation calls when bats from one colony were introduced singly into the home cage of a new colony or two bats from different colonies were cohabitated together for one month. Our experiments showed that the single bat individual altered its peak frequency of echolocation calls to approach the call of new colony members and two bats from different colonies adjusted their call frequencies toward each other to a similar frequency after being chronically cohabitated. These results indicate that the 'compromise' in echolocation calls might be used to ensure effective mutual communication among bats.

‘Compromise’ in Echolocation Calls between Different Colonies of the Intermediate Leaf-Nosed Bat (Hipposideros larvatus)

PubMed Central

Chen, Yi; Liu, Qi; Su, Qianqian; Sun, Yunxiao; Peng, Xingwen; He, Xiangyang; Zhang, Libiao

2016-01-01

Each animal population has its own acoustic signature which facilitates identification, communication and reproduction. The sonar signals of bats can convey social information, such as species identity and contextual information. The goal of this study was to determine whether bats adjust their echolocation call structures to mutually recognize and communicate when they encounter the bats from different colonies. We used the intermediate leaf-nosed bats (Hipposideros larvatus) as a case study to investigate the variations of echolocation calls when bats from one colony were introduced singly into the home cage of a new colony or two bats from different colonies were cohabitated together for one month. Our experiments showed that the single bat individual altered its peak frequency of echolocation calls to approach the call of new colony members and two bats from different colonies adjusted their call frequencies toward each other to a similar frequency after being chronically cohabitated. These results indicate that the ‘compromise’ in echolocation calls might be used to ensure effective mutual communication among bats. PMID:27029005

Automatic gain control in the echolocation system of dolphins

NASA Astrophysics Data System (ADS)

Au, Whitlow W. L.; Benoit-Bird, Kelly J.

2003-06-01

In bats and technological sonars, the gain of the receiver is progressively increased with time after the transmission of a signal to compensate for acoustic propagation loss. The current understanding of dolphin echolocation indicates that automatic gain control is not a part of their sonar system. In order to test this understanding, we have performed field measurements of free-ranging echolocating dolphins. Here we show that dolphins do possess an automatic gain control mechanism, but that it is implemented in the transmission phase rather than the receiving phase of a sonar cycle. We find that the amplitude of the dolphins' echolocation signals are highly range dependent; this amplitude increases with increasing target range, R, in a 20log(R) fashion to compensate for propagation loss. If the echolocation target is a fish school with many sound scatterers, the echoes from the school will remain nearly constant with range as the dolphin closes in on it. This characteristic has the same effect as time-varying gain in bats and technological sonar when considered from a sonar system perspective.

Change in echolocation signals with hearing loss in a false killer whale (Pseudorca crassidens).

PubMed

Kloepper, Laura N; Nachtigall, Paul E; Breese, Marlee

2010-10-01

The echolocation signals of a false killer whale (Pseudorca crassidens) were collected during a wall thickness discrimination task and compared to clicks recorded during an identical experiment in 1992. During the sixteen year time period, the subject demonstrated a loss of high frequency hearing of about 70 kHz. Clicks between the two experiments were compared to investigate the effect of hearing loss on echolocation signals. There was a significant reduction in the peak frequency, center frequency and source level of clicks between the two time periods. Additionally, the subject currently produces more signals with low frequency peaks and fewer signals with high frequency peaks than she did in 1992. These results indicate the subject changed its echolocation signals to match its range of best hearing.

Echolocation system of the bottlenose dolphin

NASA Astrophysics Data System (ADS)

Dubrovsky, N. A.

2004-05-01

The hypothesis put forward by Vel’min and Dubrovsky [1] is discussed. The hypothesis suggests that bottlenose dolphins possess two functionally separate auditory subsystems: one of them serves for analyzing extraneous sounds, as in nonecholocating terrestrial animals, and the other performs the analysis of echoes caused by the echolocation clicks of the animal itself. The first subsystem is called passive hearing, and the second, active hearing. The results of experimental studies of dolphin’s echolocation system are discussed to confirm the proposed hypothesis. For the active hearing of dolphins, the notion of a critical interval is considered as the interval of time within which the formation of a merged auditory image of the echolocation object is formed when all echo highlights of the echo from this object fall within the critical interval.

Mouth-clicks used by blind expert human echolocators - signal description and model based signal synthesis.

PubMed

Thaler, Lore; Reich, Galen M; Zhang, Xinyu; Wang, Dinghe; Smith, Graeme E; Tao, Zeng; Abdullah, Raja Syamsul Azmir Bin Raja; Cherniakov, Mikhail; Baker, Christopher J; Kish, Daniel; Antoniou, Michail

2017-08-01

Echolocation is the ability to use sound-echoes to infer spatial information about the environment. Some blind people have developed extraordinary proficiency in echolocation using mouth-clicks. The first step of human biosonar is the transmission (mouth click) and subsequent reception of the resultant sound through the ear. Existing head-related transfer function (HRTF) data bases provide descriptions of reception of the resultant sound. For the current report, we collected a large database of click emissions with three blind people expertly trained in echolocation, which allowed us to perform unprecedented analyses. Specifically, the current report provides the first ever description of the spatial distribution (i.e. beam pattern) of human expert echolocation transmissions, as well as spectro-temporal descriptions at a level of detail not available before. Our data show that transmission levels are fairly constant within a 60° cone emanating from the mouth, but levels drop gradually at further angles, more than for speech. In terms of spectro-temporal features, our data show that emissions are consistently very brief (~3ms duration) with peak frequencies 2-4kHz, but with energy also at 10kHz. This differs from previous reports of durations 3-15ms and peak frequencies 2-8kHz, which were based on less detailed measurements. Based on our measurements we propose to model transmissions as sum of monotones modulated by a decaying exponential, with angular attenuation by a modified cardioid. We provide model parameters for each echolocator. These results are a step towards developing computational models of human biosonar. For example, in bats, spatial and spectro-temporal features of emissions have been used to derive and test model based hypotheses about behaviour. The data we present here suggest similar research opportunities within the context of human echolocation. Relatedly, the data are a basis to develop synthetic models of human echolocation that could be virtual (i.e. simulated) or real (i.e. loudspeaker, microphones), and which will help understanding the link between physical principles and human behaviour.

Mouth-clicks used by blind expert human echolocators – signal description and model based signal synthesis

PubMed Central

Zhang, Xinyu; Wang, Dinghe; Tao, Zeng; Abdullah, Raja Syamsul Azmir Bin. Raja; Cherniakov, Mikhail; Kish, Daniel

2017-01-01

Echolocation is the ability to use sound-echoes to infer spatial information about the environment. Some blind people have developed extraordinary proficiency in echolocation using mouth-clicks. The first step of human biosonar is the transmission (mouth click) and subsequent reception of the resultant sound through the ear. Existing head-related transfer function (HRTF) data bases provide descriptions of reception of the resultant sound. For the current report, we collected a large database of click emissions with three blind people expertly trained in echolocation, which allowed us to perform unprecedented analyses. Specifically, the current report provides the first ever description of the spatial distribution (i.e. beam pattern) of human expert echolocation transmissions, as well as spectro-temporal descriptions at a level of detail not available before. Our data show that transmission levels are fairly constant within a 60° cone emanating from the mouth, but levels drop gradually at further angles, more than for speech. In terms of spectro-temporal features, our data show that emissions are consistently very brief (~3ms duration) with peak frequencies 2-4kHz, but with energy also at 10kHz. This differs from previous reports of durations 3-15ms and peak frequencies 2-8kHz, which were based on less detailed measurements. Based on our measurements we propose to model transmissions as sum of monotones modulated by a decaying exponential, with angular attenuation by a modified cardioid. We provide model parameters for each echolocator. These results are a step towards developing computational models of human biosonar. For example, in bats, spatial and spectro-temporal features of emissions have been used to derive and test model based hypotheses about behaviour. The data we present here suggest similar research opportunities within the context of human echolocation. Relatedly, the data are a basis to develop synthetic models of human echolocation that could be virtual (i.e. simulated) or real (i.e. loudspeaker, microphones), and which will help understanding the link between physical principles and human behaviour. PMID:28859082

Decreased echolocation performance following high-frequency hearing loss in the false killer whale (Pseudorca crassidens).

PubMed

Kloepper, L N; Nachtigall, P E; Gisiner, R; Breese, M

2010-11-01

Toothed whales and dolphins possess a hypertrophied auditory system that allows for the production and hearing of ultrasonic signals. Although the fossil record provides information on the evolution of the auditory structures found in extant odontocetes, it cannot provide information on the evolutionary pressures leading to the hypertrophied auditory system. Investigating the effect of hearing loss may provide evidence for the reason for the development of high-frequency hearing in echolocating animals by demonstrating how high-frequency hearing assists in the functioning echolocation system. The discrimination abilities of a false killer whale (Pseudorca crassidens) were measured prior to and after documented high-frequency hearing loss. In 1992, the subject had good hearing and could hear at frequencies up to 100 kHz. In 2008, the subject had lost hearing at frequencies above 40 kHz. First in 1992, and then again in 2008, the subject performed an identical echolocation task, discriminating between machined hollow aluminum cylinder targets of differing wall thickness. Performances were recorded for individual target differences and compared between both experimental years. Performances on individual targets dropped between 1992 and 2008, with a maximum performance reduction of 36.1%. These data indicate that, with a loss in high-frequency hearing, there was a concomitant reduction in echolocation discrimination ability, and suggest that the development of a hypertrophied auditory system capable of hearing at ultrasonic frequencies evolved in response to pressures for fine-scale echolocation discrimination.

Dolphin hearing during echolocation: evoked potential responses in an Atlantic bottlenose dolphin (Tursiops truncatus).

PubMed

Li, Songhai; Nachtigall, Paul E; Breese, Marlee

2011-06-15

Auditory evoked potential (AEP) responses were recorded during echolocation in an Atlantic bottlenose dolphin (Tursiops truncatus) trained to accept suction-cup EEG electrodes and detect targets by echolocation. AEP recording was triggered by the echolocation clicks of the animal. Three targets with target strengths of -34, -28 and -22 dB were used at a target distance of 2 to 6.5 m for each target. The results demonstrated that the AEP appeared to both outgoing echolocation clicks and echoes during echolocation, with AEP complexes consisting of alternative positive and negative waves. The echo-related AEP amplitudes were obviously lower than the outgoing click-related AEP amplitudes for all the targets at the investigated target distances. However, for targets with target strengths of -22 and -28 dB, the peak-to-peak amplitudes of the echo-related AEPs were dependent on the target distances. The echo-related AEP response amplitudes increased at further target distances, demonstrating an overcompensation of echo attenuation with target distance in the echo-perception system of the dolphin biosonar. Measurement and analysis of outgoing click intensities showed that the click levels increased with target distance (R) by a factor of approximately 10 to 17.5 logR depending on target strength. The results demonstrated that a dual-component biosonar control system formed by intensity compensation behavior in both the transmission and receiving phases of a biosonar cycle exists synchronously in the dolphin biosonar system.

Morphological correlates of echolocation frequency in the endemic Cape horseshoe bat, Rhinolophus capensis (Chiroptera: Rhinolophidae).

PubMed

Odendaal, Lizelle J; Jacobs, David S

2011-05-01

We investigated intraspecific variation in echolocation calls of the Cape horseshoe bat, Rhinolophus capensis, by comparing echolocation and associated morphological parameters among individuals from three populations of this species. The populations were situated in the center and at the western and eastern limits of the distribution of R. capensis. The latter two populations were situated in ecotones between vegetation biomes. Ecotone populations deviated slightly from the allometric relationship between body size and peak frequency for the genus, and there was no relationship between these variables within R. capensis. Nasal chamber length was the best predictor of peak frequency but not correlated with body size. The evolution of echolocation thus appears to have been uncoupled from body size in R. capensis. Furthermore, females used higher frequencies than males, which imply a potential social role for peak frequency. The differences in peak frequency may have originated from random founder effects and then compounded by genetic drift and/or natural selection. The latter may have acted directly on peak frequency altering skull parameters involved in echolocation independently of body size, resulting in the evolution of local acoustic signatures.

Dolphin "packet" use during long-range echolocation tasks.

PubMed

Finneran, James J

2013-03-01

When echolocating, dolphins typically emit a single broadband "click," then wait to receive the echo before emitting another click. However, previous studies have shown that during long-range echolocation tasks, they may instead emit a burst, or "packet," of several clicks, then wait for the packet of echoes to return before emitting another packet of clicks. The reasons for the use of packets are unknown. In this study, packet use was examined by having trained bottlenose dolphins perform long-range echolocation tasks. The tasks featured "phantom" echoes produced by capturing the dolphin's outgoing echolocation clicks, convolving the clicks with an impulse response to create an echo waveform, and then broadcasting the delayed, scaled echo to the dolphin. Dolphins were trained to report the presence of phantom echoes or a change in phantom echoes. Target range varied from 25 to 800 m. At ranges below 75 m, the dolphins rarely used packets. As the range increased beyond 75 m, two of the three dolphins increasingly produced packets, while the third dolphin instead utilized very high click repetition rates. The use of click packets appeared to be governed more by echo delay (target range) than echo amplitude.

Modeling perspectives on echolocation strategies inspired by bats flying in groups.

PubMed

Lin, Yuan; Abaid, Nicole

2015-12-21

Bats navigating with echolocation - which is a type of active sensing achieved by interpreting echoes resulting from self-generated ultrasonic pulses - exhibit unique behaviors during group flight. While bats may benefit from eavesdropping on their peers׳ echolocation, they also potentially suffer from confusion between their own and peers׳ pulses, caused by an effect called frequency jamming. This hardship of group flight is supported by experimental observations of bats simplifying their sound-scape by shifting their pulse frequencies or suppressing echolocation altogether. Here, we investigate eavesdropping and varying pulse emission rate from a modeling perspective to understand these behaviors׳ potential benefits and detriments. We define an agent-based model of echolocating bats avoiding collisions in a three-dimensional tunnel. Through simulation, we show that bats with reasonably accurate eavesdropping can reduce collisions compared to those neglecting information from peers. In large populations, bats minimize frequency jamming by decreasing pulse emission rate, while collision risk increases; conversely, increasing pulse emission rate minimizes collisions by allowing more sensing information generated per bat. These strategies offer benefits for both biological and engineered systems, since frequency jamming is a concern in systems using active sensing. Copyright © 2015 Elsevier Ltd. All rights reserved.

Evoked potential application to study of echolocation in cetaceans

NASA Astrophysics Data System (ADS)

Supin, Alexander Ya.; Nactigall, Paul E.; Pawloski, Jeffrey; Au, Whitlow W. L.

2002-05-01

The evoked-potential (EP) method is effective in studies of hearing capabilities of cetaceans. However, until now EP studies in cetaceans were performed only in conditions of passive hearing by recording EP to external stimuli. Can this method be applied to study active echolocation in odontocetes? To answer this question, auditory brainstem evoked responses (ABR) were recorded in a false killer whale while the animal echolocated a target within an experiment in which the animal reported the target present or absent. The ABR collection was triggered by echolocation clicks. In these conditions, the recorded ABR pattern contained a duplicate set of waves. A comparison of ABR wave delays recorded during echolocation with those recorded during regular external stimulation has shown that the first set of waves is a response to the emitted click whereas the second one is a response to the echo. Both responses, to the emitted click and to the echo, were of comparable amplitude in spite of the intensity difference of these two sounds of more than 40 dB near the animal's head. This finding indicates some mechanisms releasing responses to echoes from masking by loud emitted clicks. The evoked-potential method may be productive to investigate these mechanisms.

Finite element simulation of broadband biosonar signal propagation in the near- and far-field of an echolocating Atlantic bottlenose dolphin (Tursiops truncatus).

PubMed

Wei, Chong; Au, Whitlow W L; Ketten, Darlene R; Zhang, Yu

2018-05-01

Bottlenose dolphins project broadband echolocation signals for detecting and locating prey and predators, and for spatial orientation. There are many unknowns concerning the specifics of biosonar signal production and propagation in the head of dolphins and this manuscript represents an effort to address this topic. A two-dimensional finite element model was constructed using high resolution CT scan data. The model simulated the acoustic processes in the vertical plane of the biosonar signal emitted from the phonic lips and propagated into the water through the animal's head. The acoustic field on the animal's forehead and the farfield transmission beam pattern of the echolocating dolphin were determined. The simulation results and prior acoustic measurements were qualitatively extremely consistent. The role of the main structures on the sound propagation pathway such as the air sacs, melon, and connective tissue was investigated. Furthermore, an investigation of the driving force at the phonic lips for dolphins that emit broadband echolocation signals and porpoises that emit narrowband echolocation signals suggested that the driving force is different for the two types of biosonar. Finally, the results provide a visual understanding of the sound transmission in dolphin's biosonar.

Biosonar performance of foraging Blainvilles beaked whales (Mesoplodon densirostris)

NASA Astrophysics Data System (ADS)

Madsen, Peter T.; Johnson, Mark; Tyack, Peter L.; Aguilar de Soto, Natacha; Zimmer, Walter M. X.

2004-05-01

Echolocating animals like bats and toothed whales navigate and locate food by means of echoes from sounds transmitted by the animals themselves. Toothed whale echolocation has been studied intensively in captivity, but little information exists on how echolocation is used by wild animals for orientation and prey location. To expand on this issue, a noninvasive, acoustic Dtag (96-kHz sampling, 16-bit resolution) was deployed on two Blainvilles beaked whales. The tagged whales only clicked at depths below 200 m during deep foraging dives. The echolocation clicks are directional, 250-ms transients with peak energy in the 30-40-kHz band. Echoes from the seafloor and from prey items were recorded. The regular click rate is not adjusted to the decreasing echo delay from incoming prey until the target is within an approximate body length of the whale after which the click rate is increased rapidly akin to the buzz phase of echolocating bats. This suggests that the whales use different sonar strategies for operating in near versus far field modes. Changes in received echo intensities from prey targets during approaches are compared to the active gain control in the receiving system of bats and in the transmitting system of dolphins.

A dolphin lower jaw is a hydroacoustic antenna of the traveling wave

NASA Astrophysics Data System (ADS)

Ryabov, Vyacheslav A.

2003-10-01

The purpose of the work is the analysis of a possible function of mental foramens as channels through which the echo passes in the lower jaw fat body and the determination of a role of channels and a skull in formation of the directivity of the dolphin echolocation hearing. Concrete problems were studying of the lower jaw morphology, modeling and calculation of a dolphin, tursiops truncatus p., echolocation hearing beam pattern. The outcomes of the work indicate those morphological structures of the lower jaw; the left and right half represents two hydroacoustic receiving antennas of the traveling wave type, TWA farther. The mental foramens of a dolphin lower jaw represent nonequidistant array of waveguide delay lines, and determine the phase and amplitude distribution of each of the antenna's array. The beam pattern of the echolocation hearing was calculated with the usage of the TWA model, and the allowance of flat sound wave diffraction. The beam pattern shape is naturally determined by the echolocation hearing functionality. It is equally well adapted both for echolocation and for pulses echo detection. A steepness of the bearing characteristic is estimated; it reaches 0.7 dB per degree.

Vocal reporting of echolocation targets: dolphins often report before click trains end.

PubMed

Ridgway, S H; Elsberry, W R; Blackwood, D J; Kamolnick, T; Todd, M; Carder, D A; Chaplin, Monica; Cranford, T W

2012-01-01

Bottlenose dolphins (Tursiops truncatus) wore opaque suction cups over their eyes while stationing behind an acoustically opaque door. This put the dolphins in a known position and orientation. When the door opened, the dolphin clicked to detect targets. Trainers specified that Dolphin S emit a whistle if the target was a 7.5 cm water filled sphere, or a pulse burst if the target was a rock. S remained quiet if there was no target. Dolphin B whistled for the sphere. She remained quiet for rock and for no target. Thus, S had to choose between three different responses, whistle, pulse burst, or remain quiet. B had to choose between two different responses, whistle or remain quiet. S gave correct vocal responses averaging 114 ms after her last echolocation click (range 182 ms before and 219 ms after the last click). Average response for B was 21 ms before her last echolocation click (range 250 ms before and 95 ms after the last click in the train). More often than not, B began her whistle response before her echolocation train ended. The findings suggest separate neural pathways for generation of response vocalizations as opposed to echolocation clicks. © 2012 Acoustical Society of America.

A novel biomimetic sonarhead using beamforming technology to mimic bat echolocation.

PubMed

Steckel, Jan; Peremans, Herbert

2012-07-01

A novel biomimetic sonarhead has been developed to allow researchers of bat echolocation behavior and biomimetic sonar to perform experiments with a system similar to the bat¿s sensory system. The bat's echolocation-related transfer function (ERTF) is implemented using an array of receivers to implement the head-related transfer function (HRTF), and an array of emitters mounted on a cylindrical manifold to implement the emission pattern of the bat. The complete system is controlled by a field-programmable gate array (FPGA) based embedded system connected through a USB interface.

Ranging in Human Sonar: Effects of Additional Early Reflections and Exploratory Head Movements

PubMed Central

Wallmeier, Ludwig; Wiegrebe, Lutz

2014-01-01

Many blind people rely on echoes from self-produced sounds to assess their environment. It has been shown that human subjects can use echolocation for directional localization and orientation in a room, but echo-acoustic distance perception - e.g. to determine one's position in a room - has received little scientific attention, and systematic studies on the influence of additional early reflections and exploratory head movements are lacking. This study investigates echo-acoustic distance discrimination in virtual echo-acoustic space, using the impulse responses of a real corridor. Six blindfolded sighted subjects and a blind echolocation expert had to discriminate between two positions in the virtual corridor, which differed by their distance to the front wall, but not to the lateral walls. To solve this task, participants evaluated echoes that were generated in real time from self-produced vocalizations. Across experimental conditions, we systematically varied the restrictions for head rotations, the subjects' orientation in virtual space and the reference position. Three key results were observed. First, all participants successfully solved the task with discrimination thresholds below 1 m for all reference distances (0.75–4 m). Performance was best for the smallest reference distance of 0.75 m, with thresholds around 20 cm. Second, distance discrimination performance was relatively robust against additional early reflections, compared to other echolocation tasks like directional localization. Third, free head rotations during echolocation can improve distance discrimination performance in complex environmental settings. However, head movements do not necessarily provide a benefit over static echolocation from an optimal single orientation. These results show that accurate distance discrimination through echolocation is possible over a wide range of reference distances and environmental conditions. This is an important functional benefit of human echolocation, which may also play a major role in the calibration of auditory space representations. PMID:25551226

Developing a Passive Acoustic Monitoring Network for Harbor Porpoise in California

NASA Astrophysics Data System (ADS)

Jacobson, Eiren Kate

Assessing the abundance of and trends in whale, dolphin, and porpoise (cetacean) populations using traditional visual methods can be challenging due primarily to their limited availability at the surface of the ocean. As a result, researchers are increasingly interested in incorporating non-visual and remote observations to improve cetacean population assessments. Passive acoustic monitoring (PAM) can complement or replace visual surveys for cetaceans that produce echolocation clicks, whistles, and other vocalizations. My doctoral dissertation is focused on developing methods to improve PAM of cetaceans. I used the Monterey Bay population of harbor porpoise (Phocoena phocoena ) as a case study for methods development. In Chapter 2, I used passive acoustic data to document that harbor porpoises avoid bottlenose dolphins (Tursiops truncatus) in nearshore Monterey Bay. In Chapter 3, I investigated whether different passive acoustic instruments could be used to monitor harbor porpoise. I recorded harbor porpoise echolocation clicks simultaneously on two different passive acoustic instruments and compared the number and peak frequency of echolocation signals recorded on the two instruments. I found that the number of echolocation clicks was highly correlated between instruments but that the peak frequency of echolocation clicks was not well-correlated, suggesting that some instruments may not be capable of discriminating harbor porpoise echolocation clicks in regions where multiple species with similar echolocation signals are present. In Chapter 4, I used paired visual and passive acoustic surveys to estimate the effective detection area of the passive acoustic sensors in a Bayesian framework. This approach resulted in a posterior distribution of the effective detection area that was consistent with previously published values. In Chapter 5, I used aerial survey and passive acoustic data in a simulation framework to investigate the statistical power of different passive acoustic network designs and hypothetical changes in harbor porpoise abundance. As a whole, this dissertation used an applied approach to methods development to advance the use of PAM for cetaceans.

Temporal tuning in the bat auditory cortex is sharper when studied with natural echolocation sequences.

PubMed

Beetz, M Jerome; Hechavarría, Julio C; Kössl, Manfred

2016-06-30

Precise temporal coding is necessary for proper acoustic analysis. However, at cortical level, forward suppression appears to limit the ability of neurons to extract temporal information from natural sound sequences. Here we studied how temporal processing can be maintained in the bats' cortex in the presence of suppression evoked by natural echolocation streams that are relevant to the bats' behavior. We show that cortical neurons tuned to target-distance actually profit from forward suppression induced by natural echolocation sequences. These neurons can more precisely extract target distance information when they are stimulated with natural echolocation sequences than during stimulation with isolated call-echo pairs. We conclude that forward suppression does for time domain tuning what lateral inhibition does for selectivity forms such as auditory frequency tuning and visual orientation tuning. When talking about cortical processing, suppression should be seen as a mechanistic tool rather than a limiting element.

Auditory cortex of newborn bats is prewired for echolocation.

PubMed

Kössl, Manfred; Voss, Cornelia; Mora, Emanuel C; Macias, Silvio; Foeller, Elisabeth; Vater, Marianne

2012-04-10

Neuronal computation of object distance from echo delay is an essential task that echolocating bats must master for spatial orientation and the capture of prey. In the dorsal auditory cortex of bats, neurons specifically respond to combinations of short frequency-modulated components of emitted call and delayed echo. These delay-tuned neurons are thought to serve in target range calculation. It is unknown whether neuronal correlates of active space perception are established by experience-dependent plasticity or by innate mechanisms. Here we demonstrate that in the first postnatal week, before onset of echolocation and flight, dorsal auditory cortex already contains functional circuits that calculate distance from the temporal separation of a simulated pulse and echo. This innate cortical implementation of a purely computational processing mechanism for sonar ranging should enhance survival of juvenile bats when they first engage in active echolocation behaviour and flight.

Evoked potential recording during echolocation in a false killer whale Pseudorca crassidens (L)

NASA Astrophysics Data System (ADS)

Supin, Alexander Ya.; Nachtigall, Paul E.; Pawloski, Jeffrey; Au, Whitlow W. L.

2003-05-01

Auditory brainstem responses (ABRs) were recorded in a false killer whale while the animal echolocated a target. The ABR collection was triggered by echolocation clicks of the animal. In these conditions, the recorded ABR pattern contained a duplicate set of waves. A comparison of ABR wave delays recorded during echolocation with those recorded during regular external stimulation with experimenter generated clicks showed that the first set of waves may be a response to the emitted click whereas the second one may be a response to the echo. Both responses, to the emitted click and to the echo, were of comparable amplitude in spite of the intensity difference of these two sounds that may differ by more than 40 dB near the animal's head. This finding indicates the presence of some mechanism of releasing responses to echoes from masking by loud emitted clicks. The evoked-potential method may be productive to investigate these mechanisms.

Robustness of cortical and subcortical processing in the presence of natural masking sounds.

PubMed

Beetz, M Jerome; García-Rosales, Francisco; Kössl, Manfred; Hechavarría, Julio C

2018-05-01

Processing of ethologically relevant stimuli could be interfered by non-relevant stimuli. Animals have behavioral adaptations to reduce signal interference. It is largely unexplored whether the behavioral adaptations facilitate neuronal processing of relevant stimuli. Here, we characterize behavioral adaptations in the presence of biotic noise in the echolocating bat Carollia perspicillata and we show that the behavioral adaptations could facilitate neuronal processing of biosonar information. According to the echolocation behavior, bats need to extract their own signals in the presence of vocalizations from conspecifics. With playback experiments, we demonstrate that C. perspicillata increases the sensory acquisition rate by emitting groups of echolocation calls when flying in noisy environments. Our neurophysiological results from the auditory midbrain and cortex show that the high sensory acquisition rate does not vastly increase neuronal suppression and that the response to an echolocation sequence is partially preserved in the presence of biosonar signals from conspecifics.

Self-motion facilitates echo-acoustic orientation in humans

PubMed Central

Wallmeier, Ludwig; Wiegrebe, Lutz

2014-01-01

The ability of blind humans to navigate complex environments through echolocation has received rapidly increasing scientific interest. However, technical limitations have precluded a formal quantification of the interplay between echolocation and self-motion. Here, we use a novel virtual echo-acoustic space technique to formally quantify the influence of self-motion on echo-acoustic orientation. We show that both the vestibular and proprioceptive components of self-motion contribute significantly to successful echo-acoustic orientation in humans: specifically, our results show that vestibular input induced by whole-body self-motion resolves orientation-dependent biases in echo-acoustic cues. Fast head motions, relative to the body, provide additional proprioceptive cues which allow subjects to effectively assess echo-acoustic space referenced against the body orientation. These psychophysical findings clearly demonstrate that human echolocation is well suited to drive precise locomotor adjustments. Our data shed new light on the sensory–motor interactions, and on possible optimization strategies underlying echolocation in humans. PMID:26064556

Matching-to-sample by an echolocating dolphin (Tursiops truncatus).

PubMed

Roitblat, H L; Penner, R H; Nachtigall, P E

1990-01-01

An adult male dolphin was trained to perform a three-alternative delayed matching-to-sample task while wearing eyecups to occlude its vision. Sample and comparison stimuli consisted of a small and a large PVC plastic tube, a water-filled stainless steel sphere, and a solid aluminum cone. Stimuli were presented under water and the dolphin was allowed to identify the stimuli through echolocation. The echolocation clicks emitted by the dolphin to each sample and each comparison stimulus were recorded and analyzed. Over 48 sessions of testing, choice accuracy averaged 94.5% correct. This high level of accuracy was apparently achieved by varying the number of echolocation clicks emitted to various stimuli. Performance appeared to reflect a preexperimental stereotyped search pattern that dictated the order in which comparison items were examined and a complex sequential-sampling decision process. A model for the dolphin's decision-making processes is described.

Active acoustic interference elicits echolocation changes in heterospecific bats.

PubMed

Jones, Te K; Wohlgemuth, Melville J; Conner, William E

2018-06-27

Echolocating bats often forage in the presence of both conspecific and heterospecific individuals who have the potential to produce acoustic interference. Recent studies have shown that at least one bat species, the Brazilian free-tailed bat ( Tadarida brasiliensis ), produces specialized social signals that disrupt the sonar of conspecific competitors. We herein discuss the differences between passive and active jamming signals and test whether heterospecific jamming occurs in species overlapping spatiotemporally as well as whether such interference elicits a jamming avoidance response (JAR). We compare the capture rates of tethered moths and the echolocation parameters of big brown bats ( Eptesicus fuscus ) challenged with the playback of the jamming signal normally produced by Brazilian free-tailed bats and playback of deconstructed versions of this signal. There were no differences in the capture rates of targets with and without the jamming signal although significant changes in both spectral and temporal features of the bats' echolocation were observed. These changes are consistent with improvements of the signal-to-noise ratio in the presence of acoustic interference. Accordingly, we propose to expand the traditional definition of the JAR, stating that echolocation changes in response to interference should decrease similarity between the two signals, to include any change that increases the ability to separate returning echoes from active jamming stimuli originating from conspecific and heterospecific organisms. Flexibility in echolocation is an important characteristic for overcoming various forms of acoustic interference and may serve a purpose in interspecific interactions as well as intraspecific ones. © 2018. Published by The Company of Biologists Ltd.

Development of echolocation calls and neural selectivity for echolocation calls in the pallid bat.

PubMed

Razak, Khaleel A; Fuzessery, Zoltan M

2015-10-01

Studies of birdsongs and neural selectivity for songs have provided important insights into principles of concurrent behavioral and auditory system development. Relatively little is known about mammalian auditory system development in terms of vocalizations or other behaviorally relevant sounds. This review suggests echolocating bats are suitable mammalian model systems to understand development of auditory behaviors. The simplicity of echolocation calls with known behavioral relevance and strong neural selectivity provides a platform to address how natural experience shapes cortical receptive field (RF) mechanisms. We summarize recent studies in the pallid bat that followed development of echolocation calls and cortical processing of such calls. We also discuss similar studies in the mustached bat for comparison. These studies suggest: (1) there are different developmental sensitive periods for different acoustic features of the same vocalization. The underlying basis is the capacity for some components of the RF to be modified independent of others. Some RF computations and maps involved in call processing are present even before the cochlea is mature and well before use of echolocation in flight. Others develop over a much longer time course. (2) Normal experience is required not just for refinement, but also for maintenance, of response properties that develop in an experience independent manner. (3) Experience utilizes millisecond range changes in timing of inhibitory and excitatory RF components as substrates to shape vocalization selectivity. We suggest that bat species and call diversity provide a unique opportunity to address developmental constraints in the evolution of neural mechanisms of vocalization processing. © 2014 Wiley Periodicals, Inc.

Ontogeny of the larynx and flight ability in Jamaican fruit bats (Phyllostomidae) with considerations for the evolution of echolocation.

PubMed

Carter, Richard T; Adams, Rick A

2014-07-01

Echolocating bats have adaptations of the larynx such as hypertrophied intrinsic musculature and calcified or ossified cartilages to support sonar emission. We examined growth and development of the larynx relative to developing flight ability in Jamaican fruit bats to assess how changes in sonar production are coordinated with the onset of flight during ontogeny as a window for understanding the evolutionary relationships between these systems. In addition, we compare the extent of laryngeal calcification in an echolocating shrew species (Sorex vagrans) and the house mouse (Mus musculus), to assess what laryngeal chiropteran adaptations are associated with flight versus echolocation. Individuals were categorized into one of five developmental flight stages (flop, flutter, flap, flight, and adult) determined by drop-tests. Larynges were cleared and stained with alcian blue and alizarin red, or sectioned and stained with hematoxylin and eosin. Our results showed calcification of the cricoid cartilage in bats, represented during the flap stage and this increased significantly in individuals at the flight stage. Thyroid and arytenoid cartilages showed no evidence of calcification and neither cricoid nor thyroid showed significant increases in rate of growth relative to the larynx as a whole. The physiological cross-sectional area of the cricothyroid muscles increased significantly at the flap stage. Shrew larynges showed signs of calcification along the margins of the cricoid and thyroid cartilages, while the mouse larynx did not. These data suggest the larynx of echolocating bats becomes stronger and sturdier in tandem with flight development, indicating possible developmental integration between flight and echolocation. © 2014 Wiley Periodicals, Inc.

Development of echolocation calls and neural selectivity for echolocation calls in the pallid bat

PubMed Central

Razak, Khaleel A.; Fuzessery, Zoltan M.

2014-01-01

Studies of birdsongs and neural selectivity for songs have provided important insights into principles of concurrent behavioral and auditory system development. Relatively little is known about mammalian auditory system development in terms of vocalizations, or other behaviorally relevant sounds. This review suggests echolocating bats are suitable mammalian model systems to understand development of auditory behaviors. The simplicity of echolocation calls with known behavioral relevance and strong neural selectivity provides a platform to address how natural experience shapes cortical receptive field (RF) mechanisms. We summarize recent studies in the pallid bat that followed development of echolocation calls and cortical processing of such calls. We also discuss similar studies in the mustached bat for comparison. These studies suggest: (1) there are different developmental sensitive periods for different acoustic features of the same vocalization. The underlying basis is the capacity for some components of the RF to be modified independent of others. Some RF computations and maps involved in call processing are present even before the cochlea is mature and well before use of echolocation in flight. Others develop over a much longer time course. (2) Normal experience is required not just for refinement, but also for maintenance, of response properties that develop in an experience independent manner. (3) Experience utilizes millisecond range changes in timing of inhibitory and excitatory RF components as substrates to shape vocalization selectivity. We suggest that bat species and call diversity provide a unique opportunity to address developmental constraints in the evolution of neural mechanisms of vocalization processing. PMID:25142131

Biosonar adjustments to target range of echolocating bottlenose dolphins (Tursiops sp.) in the wild.

PubMed

Jensen, F H; Bejder, L; Wahlberg, M; Madsen, P T

2009-04-01

Toothed whales use echolocation to locate and track prey. Most knowledge of toothed whale echolocation stems from studies on trained animals, and little is known about how toothed whales regulate and use their biosonar systems in the wild. Recent research suggests that an automatic gain control mechanism in delphinid biosonars adjusts the biosonar output to the one-way transmission loss to the target, possibly a consequence of pneumatic restrictions in how fast the sound generator can be actuated and still maintain high outputs. This study examines the relationships between target range (R), click intervals, and source levels of wild bottlenose dolphins (Tursiops sp.) by recording regular (non-buzz) echolocation clicks with a linear hydrophone array. Dolphins clicked faster with decreasing distance to the array, reflecting a decreasing delay between the outgoing echolocation click and the returning array echo. However, for interclick intervals longer than 30-40 ms, source levels were not limited by the repetition rate. Thus, pneumatic constraints in the sound-production apparatus cannot account for source level adjustments to range as a possible automatic gain control mechanism for target ranges longer than a few body lengths of the dolphin. Source level estimates drop with reducing range between the echolocating dolphins and the target as a function of 17 log(R). This may indicate either (1) an active form of time-varying gain in the biosonar independent of click intervals or (2) a bias in array recordings towards a 20 log(R) relationship for apparent source levels introduced by a threshold on received click levels included in the analysis.

Context-dependent effects of noise on echolocation pulse characteristics in free-tailed bats

PubMed Central

Smotherman, Michael S.

2010-01-01

Background noise evokes a similar suite of adaptations in the acoustic structure of communication calls across a diverse range of vertebrates. Echolocating bats may have evolved specialized vocal strategies for echolocating in noise, but also seem to exhibit generic vertebrate responses such as the ubiquitous Lombard response. We wondered how bats balance generic and echolocation-specific vocal responses to noise. To address this question, we first characterized the vocal responses of flying free-tailed bats (Tadarida brasiliensis) to broadband noises varying in amplitude. Secondly, we measured the bats’ responses to band-limited noises that varied in the extent of overlap with their echolocation pulse bandwidth. We hypothesized that the bats’ generic responses to noise would be graded proportionally with noise amplitude, total bandwidth and frequency content, and consequently that more selective responses to band-limited noise such as the jamming avoidance response could be explained by a linear decomposition of the response to broadband noise. Instead, the results showed that both the nature and the magnitude of the vocal responses varied with the acoustic structure of the outgoing pulse as well as non-linearly with noise parameters. We conclude that free-tailed bats utilize separate generic and specialized vocal responses to noise in a context-dependent fashion. PMID:19672604

The interaction of outgoing echolocation pulses and echoes in the false killer whale's auditory system: Evoked-potential study

NASA Astrophysics Data System (ADS)

Supin, Alexander Ya.; Nachtigall, Paul E.; Au, Whitlow W. L.; Breese, Marlee

2004-06-01

Brain auditory evoked potentials (AEP) associated with echolocation were recorded in a false killer whale Pseudorca crassidens trained to accept suction-cup EEG electrodes and to detect targets by echolocation. AEP collection was triggered by echolocation pulses transmitted by the animal. The target was a hollow aluminum cylinder of strength of -22 dB at a distance from 1 to 8 m. Each AEP record was obtained by averaging more than 1000 individual records. All the records contained two AEP sets: the first one of a constant latency and a second one with a delay proportional to the distance. The timing of these two AEP sets was interpreted as responses to the transmitted echolocation pulse and echo, respectively. The echo-related AEP, although slightly smaller, was comparable to the outgoing click-related AEP in amplitude, even though at a target distance as far as 8 m the echo intensity was as low as -64 dB relative to the transmitted pulse in front of the head. The amplitude of the echo-related AEP was almost independent of distance, even though variation of target distance from 1 to 8 m influenced the echo intensity by as much as 36 dB.

The interaction of outgoing echolocation pulses and echoes in the false killer whale's auditory system: evoked-potential study.

PubMed

Supin, Alexander Ya; Nachtigall, Paul E; Au, Whitlow W L; Breese, Marlee

2004-06-01

Brain auditory evoked potentials (AEP) associated with echolocation were recorded in a false killer whale Pseudorca crassidens trained to accept suction-cup EEG electrodes and to detect targets by echolocation. AEP collection was triggered by echolocation pulses transmitted by the animal. The target was a hollow aluminum cylinder of strength of -22 dB at a distance from 1 to 8 m. Each AEP record was obtained by averaging more than 1000 individual records. All the records contained two AEP sets: the first one of a constant latency and a second one with a delay proportional to the distance. The timing of these two AEP sets was interpreted as responses to the transmitted echolocation pulse and echo, respectively. The echo-related AEP, although slightly smaller, was comparable to the outgoing click-related AEP in amplitude, even though at a target distance as far as 8 m the echo intensity was as low as -64 dB relative to the transmitted pulse in front of the head. The amplitude of the echo-related AEP was almost independent of distance, even though variation of target distance from 1 to 8 m influenced the echo intensity by as much as 36 dB.

Echolocation behaviour adapted to prey in foraging Blainville's beaked whale (Mesoplodon densirostris)

PubMed Central

Johnson, M; Hickmott, L.S; Aguilar Soto, N; Madsen, P.T

2007-01-01

Toothed whales echolocating in the wild generate clicks with low repetition rates to locate prey but then produce rapid sequences of clicks, called buzzes, when attempting to capture prey. However, little is known about the factors that determine clicking rates or how prey type and behaviour influence echolocation-based foraging. Here we study Blainville's beaked whales foraging in deep water using a multi-sensor DTAG that records both outgoing echolocation clicks and echoes returning from mesopelagic prey. We demonstrate that the clicking rate at the beginning of buzzes is related to the distance between whale and prey, supporting the presumption that whales focus on a specific prey target during the buzz. One whale showed a bimodal relationship between target range and clicking rate producing abnormally slow buzz clicks while attempting to capture large echoic targets, probably schooling prey, with echo duration indicating a school diameter of up to 4.3 m. These targets were only found when the whale performed tight circling manoeuvres spending up to five times longer in water volumes with large targets than with small targets. The result indicates that toothed whales in the wild can adjust their echolocation behaviour and movement for capture of different prey on the basis of structural echo information. PMID:17986434

The influence of bat echolocation call duration and timing on auditory encoding of predator distance in noctuoid moths.

PubMed

Gordon, Shira D; Ter Hofstede, Hannah M

2018-03-22

Animals co-occur with multiple predators, making sensory systems that can encode information about diverse predators advantageous. Moths in the families Noctuidae and Erebidae have ears with two auditory receptor cells (A1 and A2) used to detect the echolocation calls of predatory bats. Bat communities contain species that vary in echolocation call duration, and the dynamic range of A1 is limited by the duration of sound, suggesting that A1 provides less information about bats with shorter echolocation calls. To test this hypothesis, we obtained intensity-response functions for both receptor cells across many moth species for sound pulse durations representing the range of echolocation call durations produced by bat species in northeastern North America. We found that the threshold and dynamic range of both cells varied with sound pulse duration. The number of A1 action potentials per sound pulse increases linearly with increasing amplitude for long-duration pulses, saturating near the A2 threshold. For short sound pulses, however, A1 saturates with only a few action potentials per pulse at amplitudes far lower than the A2 threshold for both single sound pulses and pulse sequences typical of searching or approaching bats. Neural adaptation was only evident in response to approaching bat sequences at high amplitudes, not search-phase sequences. These results show that, for short echolocation calls, a large range of sound levels cannot be coded by moth auditory receptor activity, resulting in no information about the distance of a bat, although differences in activity between ears might provide information about direction. © 2018. Published by The Company of Biologists Ltd.

Echolocation signals of free-ranging killer whales (Orcinus orca) and modeling of foraging for chinook salmon (Oncorhynchus tshawytscha)

NASA Astrophysics Data System (ADS)

Au, Whitlow W. L.; Ford, John K. B.; Horne, John K.; Allman, Kelly A. Newman

2004-02-01

Fish-eating ``resident''-type killer whales (Orcinus orca) that frequent the coastal waters off northeastern Vancouver Island, Canada have a strong preference for chinook salmon (Oncorhynchus tshawytscha). The whales in this region often forage along steep cliffs that extend into the water, echolocating their prey. Echolocation signals of resident killer whales were measured with a four-hydrophone symmetrical star array and the signals were simultaneously digitized at a sample rate of 500 kHz using a lunch-box PC. A portable VCR recorded the images from an underwater camera located adjacent to the array center. Only signals emanating from close to the beam axis (1185 total) were chosen for a detailed analysis. Killer whales project very broadband echolocation signals (Q equal 0.9 to 1.4) that tend to have bimodal frequency structure. Ninety-seven percent of the signals had center frequencies between 45 and 80 kHz with bandwidths between 35 and 50 kHz. The peak-to-peak source level of the echolocation signals decreased as a function of the one-way transmission loss to the array. Source levels varied between 195 and 224 dB re:1 μPa. Using a model of target strength for chinook salmon, the echo levels from the echolocation signals are estimated for different horizontal ranges between a whale and a salmon. At a horizontal range of 100 m, the echo level should exceed an Orcinus hearing threshold at 50 kHz by over 29 dB and should be greater than sea state 4 noise by at least 9 dB. In moderately heavy rain conditions, the detection range will be reduced substantially and the echo level at a horizontal range of 40 m would be close to the level of the rain noise.

Echolocation signals of free-ranging killer whales (Orcinus orca) and modeling of foraging for chinook salmon (Oncorhynchus tshawytscha).

PubMed

Au, Whitlow W L; Ford, John K B; Horne, John K; Allman, Kelly A Newman

2004-02-01

Fish-eating "resident"-type killer whales (Orcinus orca) that frequent the coastal waters off northeastern Vancouver Island, Canada have a strong preference for chinook salmon (Oncorhynchus tshawytscha). The whales in this region often forage along steep cliffs that extend into the water, echolocating their prey. Echolocation signals of resident killer whales were measured with a four-hydrophone symmetrical star array and the signals were simultaneously digitized at a sample rate of 500 kHz using a lunch-box PC. A portable VCR recorded the images from an underwater camera located adjacent to the array center. Only signals emanating from close to the beam axis (1185 total) were chosen for a detailed analysis. Killer whales project very broadband echolocation signals (Q equal 0.9 to 1.4) that tend to have bimodal frequency structure. Ninety-seven percent of the signals had center frequencies between 45 and 80 kHz with bandwidths between 35 and 50 kHz. The peak-to-peak source level of the echolocation signals decreased as a function of the one-way transmission loss to the array. Source levels varied between 195 and 224 dB re: 1 microPa. Using a model of target strength for chinook salmon, the echo levels from the echolocation signals are estimated for different horizontal ranges between a whale and a salmon. At a horizontal range of 100 m, the echo level should exceed an Orcinus hearing threshold at 50 kHz by over 29 dB and should be greater than sea state 4 noise by at least 9 dB. In moderately heavy rain conditions, the detection range will be reduced substantially and the echo level at a horizontal range of 40 m would be close to the level of the rain noise.

Morphology suggests noseleaf and pinnae cooperate to enhance bat echolocation.

PubMed

Kuc, Roman

2010-11-01

A protruding noseleaf and concave pinna structures suggest that some bats may use these to enhance their echolocation capabilities. This paper considers two possible mechanisms that each exploit the combination of direct and delayed acoustic paths to achieve more complex emission or sensitivity echolocation patterns. The first is an emission mechanism, in which the protruding noseleaf vibrates to emit sound in both the forward and backward directions, and pinna structures reflect the backward emission to enhance the forward beam. The second is a reception mechanism, which has a direct echo path to the ear canal and a delayed path involving pinna structures reflecting onto the noseleaf and then into the ear canal. A model using Davis' Round-eared Bat illustrates that such direct and delayed acoustic paths provide target elevation cues. The model demonstrates the delayed pinna component can increase the on-axis emission strength, narrow the beam width, and sculpt frequency-dependent beam patterns useful for echolocation.

Baird's beaked whale echolocation signals.

PubMed

Baumann-Pickering, Simone; Yack, Tina M; Barlow, Jay; Wiggins, Sean M; Hildebrand, John A

2013-06-01

Echolocation signals from Baird's beaked whales were recorded during visual and acoustic shipboard surveys of cetaceans in the California Current ecosystem and with autonomous, long-term recorders in the Southern California Bight. The preliminary measurement of the visually validated Baird's beaked whale echolocation signals from towed array data were used as a basis for identifying Baird's signals in the autonomous recorder data. Two distinct signal types were found, one being a beaked whale-like frequency modulated (FM) pulse, the other being a dolphin-like broadband click. The median FM inter-pulse interval was 230 ms. Both signal types showed a consistent multi-peak structure in their spectra with peaks at ~9, 16, 25, and 40 kHz. Depending on signal type, as well as recording aspect and distance to the hydrophone, these peaks varied in relative amplitude. The description of Baird's echolocation signals will allow for studies of their distribution and abundance using towed array data without associated visual sightings and from autonomous seafloor hydrophones.

Male sperm whale acoustic behavior observed from multipaths at a single hydrophone

NASA Astrophysics Data System (ADS)

Laplanche, Christophe; Adam, Olivier; Lopatka, Maciej; Motsch, Jean-François

2005-10-01

Sperm whales generate transient sounds (clicks) when foraging. These clicks have been described as echolocation sounds, a result of having measured the source level and the directionality of these signals and having extrapolated results from biosonar tests made on some small odontocetes. The authors propose a passive acoustic technique requiring only one hydrophone to investigate the acoustic behavior of free-ranging sperm whales. They estimate whale pitch angles from the multipath distribution of click energy. They emphasize the close bond between the sperm whale's physical and acoustic activity, leading to the hypothesis that sperm whales might, like some small odontocetes, control click level and rhythm. An echolocation model estimating the range of the sperm whale's targets from the interclick interval is computed and tested during different stages of the whale's dive. Such a hypothesis on the echolocation process would indicate that sperm whales echolocate their prey layer when initiating their dives and follow a methodic technique when foraging.

Divergent evolutionary rates in vertebrate and mammalian specific conserved non-coding elements (CNEs) in echolocating mammals.

PubMed

Davies, Kalina T J; Tsagkogeorga, Georgia; Rossiter, Stephen J

2014-12-19

The majority of DNA contained within vertebrate genomes is non-coding, with a certain proportion of this thought to play regulatory roles during development. Conserved Non-coding Elements (CNEs) are an abundant group of putative regulatory sequences that are highly conserved across divergent groups and thus assumed to be under strong selective constraint. Many CNEs may contain regulatory factor binding sites, and their frequent spatial association with key developmental genes - such as those regulating sensory system development - suggests crucial roles in regulating gene expression and cellular patterning. Yet surprisingly little is known about the molecular evolution of CNEs across diverse mammalian taxa or their role in specific phenotypic adaptations. We examined 3,110 vertebrate-specific and ~82,000 mammalian-specific CNEs across 19 and 9 mammalian orders respectively, and tested for changes in the rate of evolution of CNEs located in the proximity of genes underlying the development or functioning of auditory systems. As we focused on CNEs putatively associated with genes underlying the development/functioning of auditory systems, we incorporated echolocating taxa in our dataset because of their highly specialised and derived auditory systems. Phylogenetic reconstructions of concatenated CNEs broadly recovered accepted mammal relationships despite high levels of sequence conservation. We found that CNE substitution rates were highest in rodents and lowest in primates, consistent with previous findings. Comparisons of CNE substitution rates from several genomic regions containing genes linked to auditory system development and hearing revealed differences between echolocating and non-echolocating taxa. Wider taxonomic sampling of four CNEs associated with the homeobox genes Hmx2 and Hmx3 - which are required for inner ear development - revealed family-wise variation across diverse bat species. Specifically within one family of echolocating bats that utilise frequency-modulated echolocation calls varying widely in frequency and intensity high levels of sequence divergence were found. Levels of selective constraint acting on CNEs differed both across genomic locations and taxa, with observed variation in substitution rates of CNEs among bat species. More work is needed to determine whether this variation can be linked to echolocation, and wider taxonomic sampling is necessary to fully document levels of conservation in CNEs across diverse taxa.

Recognition of aspect-dependent three-dimensional objects by an echolocating Atlantic bottlenose dolphin.

PubMed

Helweg, D A; Roitblat, H L; Nachtigall, P E; Hautus, M J

1996-01-01

We examined the ability of a bottlenose dolphin (Tursiops truncatus) to recognize aspect-dependent objects using echolocation. An aspect-dependent object such as a cube produces acoustically different echoes at different angles relative to the echolocation signal. The dolphin recognized the objects even though the objects were free to rotate and sway. A linear discriminant analysis and nearest centroid classifier could classify the objects using average amplitude, center frequency, and bandwidth of object echoes. The results show that dolphins can use varying acoustic properties to recognize constant objects and suggest that aspect-independent representations may be formed by combining information gleaned from multiple echoes.

A blind climber: The first evidence of ultrasonic echolocation in arboreal mammals.

PubMed

Panyutina, Aleksandra A; Kuznetsov, Alexander N; Volodin, Ilya A; Abramov, Alexei V; Soldatova, Irina B

2017-03-01

The means of orientation is studied in the Vietnamese pygmy dormouse Typhlomys chapensis, a poorly known enigmatic semi-fossorial semi-arboreal rodent. Data on eye structure are presented, which prove that Typhlomys (translated as "the blind mouse") is incapable of object vision: the retina is folded and retains no more than 2500 ganglion cells in the focal plane, and the optic nerve is subject to gliosis. Hence, Typhlomys has no other means for rapid long-range orientation among tree branches other than echolocation. Ultrasonic vocalization recordings at the frequency range of 50-100 kHz support this hypothesis. The vocalizations are represented by bouts of up to 7 more or less evenly-spaced and uniform frequency-modulated sweep-like pulses in rapid succession. Structurally, these sweeps are similar to frequency-modulated ultrasonic echolocation calls of some bat species, but they are too faint to be revealed with a common bat detector. When recording video simultaneously with the ultrasonic audio, a significantly greater pulse rate during locomotion compared to that of resting animals has been demonstrated. Our findings of locomotion-associated ultrasonic vocalization in a fast-climbing but weakly-sighted small mammal ecotype add support to the "echolocation-first theory" of pre-flight origin of echolocation in bats. © 2016 International Society of Zoological Sciences, Institute of Zoology/Chinese Academy of Sciences and John Wiley & Sons Australia, Ltd.

Tight coordination of aerial flight maneuvers and sonar call production in insectivorous bats

PubMed Central

Falk, Benjamin; Kasnadi, Joseph; Moss, Cynthia F.

2015-01-01

ABSTRACT Echolocating bats face the challenge of coordinating flight kinematics with the production of echolocation signals used to guide navigation. Previous studies of bat flight have focused on kinematics of fruit and nectar-feeding bats, often in wind tunnels with limited maneuvering, and without analysis of echolocation behavior. In this study, we engaged insectivorous big brown bats in a task requiring simultaneous turning and climbing flight, and used synchronized high-speed motion-tracking cameras and audio recordings to quantify the animals' coordination of wing kinematics and echolocation. Bats varied flight speed, turn rate, climb rate and wingbeat rate as they navigated around obstacles, and they adapted their sonar signals in patterning, duration and frequency in relation to the timing of flight maneuvers. We found that bats timed the emission of sonar calls with the upstroke phase of the wingbeat cycle in straight flight, and that this relationship changed when bats turned to navigate obstacles. We also characterized the unsteadiness of climbing and turning flight, as well as the relationship between speed and kinematic parameters. Adaptations in the bats' echolocation call frequency suggest changes in beam width and sonar field of view in relation to obstacles and flight behavior. By characterizing flight and sonar behaviors in an insectivorous bat species, we find evidence of exquisitely tight coordination of sensory and motor systems for obstacle navigation and insect capture. PMID:26582935

The Use of Echolocation as a Mobility Aid for Blind Persons.

ERIC Educational Resources Information Center

Boehm, R.

1986-01-01

The value of echolocation for enhancing mobility of the blind was examined with five blind subjects and 11 sighted, blindfolded subjects. A hand held clicker provided the sounds for navigation through an unfamiliar hallway. Results indicated the blind subjects were better able to identify obstacles correctly using reflected sounds. (Author/DB)

Echolocation Reconsidered: Using Spatial Variations in the Ambient Sound Field To Guide Locomotion.

ERIC Educational Resources Information Center

Ashmead, Daniel H.; Wall, Robert S.; Eaton, Susan B.; Ebinger, Kiara A.; Snook-Hill, Mary-Maureen; And Others

1998-01-01

Presents an acoustical model and evidence from four experiments that children with visual impairments use the buildup of low-frequency sound along walls to guide locomotion. The model differs from the concept of echolocation by emphasizing sound that is ambient, rather than self-produced, and of low frequency. (Author/CR)

The energy ratio mapping algorithm: a tool to improve the energy-based detection of odontocete echolocation clicks.

PubMed

Klinck, Holger; Mellinger, David K

2011-04-01

The energy ratio mapping algorithm (ERMA) was developed to improve the performance of energy-based detection of odontocete echolocation clicks, especially for application in environments with limited computational power and energy such as acoustic gliders. ERMA systematically evaluates many frequency bands for energy ratio-based detection of echolocation clicks produced by a target species in the presence of the species mix in a given geographic area. To evaluate the performance of ERMA, a Teager-Kaiser energy operator was applied to the series of energy ratios as derived by ERMA. A noise-adaptive threshold was then applied to the Teager-Kaiser function to identify clicks in data sets. The method was tested for detecting clicks of Blainville's beaked whales while rejecting echolocation clicks of Risso's dolphins and pilot whales. Results showed that the ERMA-based detector correctly identified 81.6% of the beaked whale clicks in an extended evaluation data set. Average false-positive detection rate was 6.3% (3.4% for Risso's dolphins and 2.9% for pilot whales).

No evidence for spectral jamming avoidance in echolocation behavior of foraging pipistrelle bats

PubMed Central

Götze, Simone; Koblitz, Jens C.; Denzinger, Annette; Schnitzler, Hans-Ulrich

2016-01-01

Frequency shifts in signals of bats flying near conspecifics have been interpreted as a spectral jamming avoidance response (JAR). However, several prerequisites supporting a JAR hypothesis have not been controlled for in previous studies. We recorded flight and echolocation behavior of foraging Pipistrellus pipistrellus while flying alone and with a conspecific and tested whether frequency changes were due to a spectral JAR with an increased frequency difference, or whether changes could be explained by other reactions. P. pipistrellus reacted to conspecifics with a reduction of sound duration and often also pulse interval, accompanied by an increase in terminal frequency. This reaction is typical of behavioral situations where targets of interest have captured the bat’s attention and initiated a more detailed exploration. All observed frequency changes were predicted by the attention reaction alone, and do not support the JAR hypothesis of increased frequency separation. Reaction distances of 1–11 m suggest that the attention response may be elicited either by detection of the conspecific by short range active echolocation or by long range passive acoustic detection of echolocation calls. PMID:27502900

Spatial orientation of different frequencies within the echolocation beam of a Tursiops truncatus and Pseudorca crassidens.

PubMed

Ibsen, Stuart D; Nachtigall, Paul E; Krause-Nehring, Jacqueline; Kloepper, Laura; Breese, Marlee; Li, Songhai; Vlachos, Stephanie

2012-08-01

A two-dimensional array of 16 hydrophones was created to map the spatial distribution of different frequencies within the echolocation beam of a Tursiops truncatus and a Pseudorca crassidens. It was previously shown that both the Tursiops and Pseudorca only paid attention to frequencies between 29 and 42 kHz while echolocating. Both individuals tightly focused the 30 kHz frequency and the spatial location of the focus was consistently pointed toward the target. At 50 kHz the beam was less focused and less precisely pointed at the target. At 100 kHz the focus was often completely lost and was not pointed at the target. This indicates that these individuals actively focused the beam toward the target only in the frequency range they paid attention to. Frequencies outside this range were left unfocused and undirected. This focusing was probably achieved through sensorimotor control of the melon morphology and nasal air sacs. This indicates that both morphologically different species can control the spatial distribution of different frequency ranges within the echolocation beam to create consistent ensonation of desired targets.

Origin of the double- and multi-pulse structure of echolocation signals in Yangtze finless porpoise (Neophocaena phocaenoides asiaeorientialis)

NASA Astrophysics Data System (ADS)

Li, Songhai; Wang, Kexiong; Wang, Ding; Akamatsu, Tomonari

2005-12-01

The signals of dolphins and porpoises often exhibit a multi-pulse structure. Here, echolocation signal recordings were made from four geometrically distinct positions of seven Yangtze finless porpoises temporarily housed in a relatively small, enclosed area. Some clicks demonstrated double-pulse, and others multi-pulse, structure. The interpulse intervals between the first and second pulse of the double- and multi-pulse clicks were significantly different among data from the four different positions (p<0.01, one-way ANOVA). These results indicate that the interpulse interval and structure of the double- and multi-pulse echolocation signals depend on the hydrophone geometry of the animal, and that the double- and multi-pulse structure of echolocation signals in Yangtze finless porpoise is not caused by the phonating porpoise itself, but by the multipath propagation of the signal. Time delays in the 180° phase-shifted surface reflection pulse and the nonphase-shifted bottom reflection pulse of the multi-pulse structures, relative to the direct signal, can be used to calculate the distance to a phonating animal.

Tight coordination of aerial flight maneuvers and sonar call production in insectivorous bats.

PubMed

Falk, Benjamin; Kasnadi, Joseph; Moss, Cynthia F

2015-11-01

Echolocating bats face the challenge of coordinating flight kinematics with the production of echolocation signals used to guide navigation. Previous studies of bat flight have focused on kinematics of fruit and nectar-feeding bats, often in wind tunnels with limited maneuvering, and without analysis of echolocation behavior. In this study, we engaged insectivorous big brown bats in a task requiring simultaneous turning and climbing flight, and used synchronized high-speed motion-tracking cameras and audio recordings to quantify the animals' coordination of wing kinematics and echolocation. Bats varied flight speed, turn rate, climb rate and wingbeat rate as they navigated around obstacles, and they adapted their sonar signals in patterning, duration and frequency in relation to the timing of flight maneuvers. We found that bats timed the emission of sonar calls with the upstroke phase of the wingbeat cycle in straight flight, and that this relationship changed when bats turned to navigate obstacles. We also characterized the unsteadiness of climbing and turning flight, as well as the relationship between speed and kinematic parameters. Adaptations in the bats' echolocation call frequency suggest changes in beam width and sonar field of view in relation to obstacles and flight behavior. By characterizing flight and sonar behaviors in an insectivorous bat species, we find evidence of exquisitely tight coordination of sensory and motor systems for obstacle navigation and insect capture. © 2015. Published by The Company of Biologists Ltd.

Comparisons of MRI images, and auditory-related and vocal-related protein expressions in the brain of echolocation bats and rodents.

PubMed

Hsiao, Chun-Jen; Hsu, Chih-Hsiang; Lin, Ching-Lung; Wu, Chung-Hsin; Jen, Philip Hung-Sun

2016-08-17

Although echolocating bats and other mammals share the basic design of laryngeal apparatus for sound production and auditory system for sound reception, they have a specialized laryngeal mechanism for ultrasonic sound emissions as well as a highly developed auditory system for processing species-specific sounds. Because the sounds used by bats for echolocation and rodents for communication are quite different, there must be differences in the central nervous system devoted to producing and processing species-specific sounds between them. The present study examines the difference in the relative size of several brain structures and expression of auditory-related and vocal-related proteins in the central nervous system of echolocation bats and rodents. Here, we report that bats using constant frequency-frequency-modulated sounds (CF-FM bats) and FM bats for echolocation have a larger volume of midbrain nuclei (inferior and superior colliculi) and cerebellum relative to the size of the brain than rodents (mice and rats). However, the former have a smaller volume of the cerebrum and olfactory bulb, but greater expression of otoferlin and forkhead box protein P2 than the latter. Although the size of both midbrain colliculi is comparable in both CF-FM and FM bats, CF-FM bats have a larger cerebrum and greater expression of otoferlin and forkhead box protein P2 than FM bats. These differences in brain structure and protein expression are discussed in relation to their biologically relevant sounds and foraging behavior.

Development of echolocation and communication vocalizations in the big brown bat, Eptesicus fuscus.

PubMed

Monroy, Jenna A; Carter, Matthew E; Miller, Kimberly E; Covey, Ellen

2011-05-01

Big brown bats form large maternity colonies of up to 200 mothers and their pups. If pups are separated from their mothers, they can locate each other using vocalizations. The goal of this study was to systematically characterize the development of echolocation and communication calls from birth through adulthood to determine whether they develop from a common precursor at the same or different rates, or whether both types are present initially. Three females and their six pups were isolated from our captive breeding colony. We recorded vocal activity from postnatal day 1 to 35, both when the pups were isolated and when they were reunited with their mothers. At birth, pups exclusively emitted isolation calls, with a fundamental frequency range <20 kHz, and duration >30 ms. By the middle of week 1, different types of vocalizations began to emerge. Starting in week 2, pups in the presence of their mothers emitted sounds that resembled adult communication vocalizations, with a lower frequency range and longer durations than isolation calls or echolocation signals. During weeks 2 and 3, these vocalizations were extremely heterogeneous, suggesting that the pups went through a babbling stage before establishing a repertoire of stereotyped adult vocalizations around week 4. By week 4, vocalizations emitted when pups were alone were identical to adult echolocation signals. Echolocation and communication signals both appear to develop from the isolation call, diverging during week 2 and continuing to develop at different rates for several weeks until the adult vocal repertoire is established.

Dynamics of the echolocation beam during prey pursuit in aerial hawking bats.

PubMed

Jakobsen, Lasse; Olsen, Mads Nedergaard; Surlykke, Annemarie

2015-06-30

In the evolutionary arms race between prey and predator, measures and countermeasures continuously evolve to increase survival on both sides. Bats and moths are prime examples. When exposed to intense ultrasound, eared moths perform dramatic escape behaviors. Vespertilionid and rhinolophid bats broaden their echolocation beam in the final stage of pursuit, presumably as a countermeasure to keep evading moths within their "acoustic field of view." In this study, we investigated if dynamic beam broadening is a general property of echolocation when catching moving prey. We recorded three species of emballonurid bats, Saccopteryx bilineata, Saccopteryx leptura, and Rhynchonycteris naso, catching airborne insects in the field. The study shows that S. bilineata and S. leptura maintain a constant beam shape during the entire prey pursuit, whereas R. naso broadens the beam by lowering the peak call frequency from 100 kHz during search and approach to 67 kHz in the buzz. Surprisingly, both Saccopteryx bats emit calls with very high energy throughout the pursuit, up to 60 times more than R. naso and Myotis daubentonii (a similar sized vespertilionid), providing them with as much, or more, peripheral "vision" than the vespertilionids, but ensonifying objects far ahead suggesting more clutter. Thus, beam broadening is not a fundamental property of the echolocation system. However, based on the results, we hypothesize that increased peripheral detection is crucial to all aerial hawking bats in the final stages of prey pursuit and speculate that beam broadening is a feature characterizing more advanced echolocation.

Fast sensory–motor reactions in echolocating bats to sudden changes during the final buzz and prey intercept

PubMed Central

Geberl, Cornelia; Brinkløv, Signe; Wiegrebe, Lutz; Surlykke, Annemarie

2015-01-01

Echolocation is an active sense enabling bats and toothed whales to orient in darkness through echo returns from their ultrasonic signals. Immediately before prey capture, both bats and whales emit a buzz with such high emission rates (≥180 Hz) and overall duration so short that its functional significance remains an enigma. To investigate sensory–motor control during the buzz of the insectivorous bat Myotis daubentonii, we removed prey, suspended in air or on water, before expected capture. The bats responded by shortening their echolocation buzz gradually; the earlier prey was removed down to approximately 100 ms (30 cm) before expected capture, after which the full buzz sequence was emitted both in air and over water. Bats trawling over water also performed the full capture behavior, but in-air capture motions were aborted, even at very late prey removals (<20 ms = 6 cm before expected contact). Thus, neither the buzz nor capture movements are stereotypical, but dynamically adapted based on sensory feedback. The results indicate that echolocation is controlled mainly by acoustic feedback, whereas capture movements are adjusted according to both acoustic and somatosensory feedback, suggesting separate (but coordinated) central motor control of the two behaviors based on multimodal input. Bat echolocation, especially the terminal buzz, provides a unique window to extremely fast decision processes in response to sensory feedback and modulation through attention in a naturally behaving animal. PMID:25775538

Target detection, shape discrimination, and signal characteristics of an echolocating false killer whale (Pseudorca crassidens).

PubMed

Brill, R L; Pawloski, J L; Helweg, D A; Au, W W; Moore, P W

1992-09-01

This study demonstrated the ability of a false killer whale (Pseudorca crassidens) to discriminate between two targets and investigated the parameters of the whale's emitted signals for changes related to test conditions. Target detection performance comparable to the bottlenose dolphin's (Tursiops truncatus) has previously been reported for echolocating false killer whales. No other echolocation capabilities have been reported. A false killer whale, naive to conditioned echolocation tasks, was initially trained to detect a cylinder in a "go/no-go" procedure over ranges of 3 to 8 m. The transition from a detection task to a discrimination task was readily achieved by introducing a spherical comparison target. Finally, the cylinder was successfully compared to spheres of two different sizes and target strengths. Multivariate analyses were used to evaluate the parameters of emitted signals. Duncan's multiple range tests showed significant decreases (df = 185, p less than 0.05) in both source level and bandwidth in the transition from detection to discrimination. Analysis of variance revealed a significant decrease in the number of clicks over test conditions [F(5.26) = 5.23, p less than 0.0001]. These data suggest that the whale relied on cues relevant to target shape as well as target strength, that changes in source level and bandwidth were task-related, that the decrease in clicks was associated with learning experience, and that Pseudorca's ability to discriminate shapes using echolocation may be comparable to that of Tursiops truncatus.

Adaptations for Substrate Gleaning in Bats: The Pallid Bat as a Case Study.

PubMed

Razak, Khaleel A

2018-06-06

Substrate gleaning is a foraging strategy in which bats use a mixture of echolocation, prey-generated sounds, and vision to localize and hunt surface-dwelling prey. Many substrate-gleaning species depend primarily on prey-generated noise to hunt. Use of echolocation is limited to general orientation and obstacle avoidance. This foraging strategy involves a different set of selective pressures on morphology, behavior, and auditory system organization of bats compared to the use of echolocation for both hunting and navigation. Gleaning likely evolved to hunt in cluttered environments and/or as a counterstrategy to reduce detection by eared prey. Gleaning bats simultaneously receive streams of echoes from obstacles and prey-generated noise, and have to segregate these acoustic streams to attend to one or both. Not only do these bats have to be exquisitely sensitive to the soft, low frequency sounds produced by walking/rustling prey, they also have to precisely localize these sounds. Gleaners typically use low intensity echolocation calls. Such stealth echolocation requires a nervous system that is attuned to low intensity sound processing. In addition, landing on the ground to hunt may bring gleaners in close proximity to venomous prey. In fact, at least 2 gleaning bat species are known to hunt highly venomous scorpions. While a number of studies have addressed adaptations for echolocation in bats that hunt in the air, very little is known about the morphological, behavioral, and neural specializations for gleaning in bats. This review highlights the novel insights gleaning bats provide into bat evolution, particularly auditory pathway organization and ion channel structure/function relationships. Gleaning bats are found in multiple families, suggesting convergent evolution of specializations for gleaning as a foraging strategy. However, most of this review is based on recent work on a single species - the pallid bat (Antrozous palli dus) - symptomatic of the fact that more comparative work is needed to identify the mechanisms that facilitate gleaning behavior. © 2018 S. Karger AG, Basel.

The adaptive function of tiger moth clicks against echolocating bats: an experimental and synthetic approach.

PubMed

Ratcliffe, John M; Fullard, James H

2005-12-01

We studied the efficiency and effects of the multiple sensory cues of tiger moths on echolocating bats. We used the northern long-eared bat, Myotis septentrionalis, a purported moth specialist that takes surface-bound prey (gleaning) and airborne prey (aerial hawking), and the dogbane tiger moth, Cycnia tenera, an eared species unpalatable to bats that possesses conspicuous colouration and sound-producing organs (tymbals). This is the first study to investigate the interaction of tiger moths and wild-caught bats under conditions mimicking those found in nature and to demand the use of both aerial hawking and gleaning strategies by bats. Further, it is the first to report spectrograms of the sounds produced by tiger moths while under aerial attack by echolocating bats. During both aerial hawking and gleaning trials, all muted C. tenera and perched intact C. tenera were attacked by M. septentrionalis, indicating that M. septentrionalis did not discriminate C. tenera from palatable moths based on potential echoic and/or non-auditory cues. Intact C. tenera were attacked significantly less often than muted C. tenera during aerial hawking attacks: tymbal clicks were therefore an effective deterrent in an aerial hawking context. During gleaning attacks, intact and muted C. tenera were always attacked and suffered similar mortality rates, suggesting that while handling prey this bat uses primarily chemical signals. Our results also show that C. tenera temporally matches the onset of click production to the ;approach phase' echolocation calls produced by aerial hawking attacking bats and that clicks themselves influence the echolocation behaviour of attacking bats. In the context of past research, these findings support the hypotheses that the clicks of arctiid moths are both an active defence (through echolocation disruption) and a reliable indicator of chemical defence against aerial-hawking bats. We suggest these signals are specialized for an aerial context.

Evolution of the heteroharmonic strategy for target-range computation in the echolocation of Mormoopidae

PubMed Central

Mora, Emanuel C.; Macías, Silvio; Hechavarría, Julio; Vater, Marianne; Kössl, Manfred

2013-01-01

Echolocating bats use the time elapsed from biosonar pulse emission to the arrival of echo (defined as echo-delay) to assess target-distance. Target-distance is represented in the brain by delay-tuned neurons that are classified as either “heteroharmonic” or “homoharmormic.” Heteroharmonic neurons respond more strongly to pulse-echo pairs in which the timing of the pulse is given by the fundamental biosonar harmonic while the timing of echoes is provided by one (or several) of the higher order harmonics. On the other hand, homoharmonic neurons are tuned to the echo delay between similar harmonics in the emitted pulse and echo. It is generally accepted that heteroharmonic computations are advantageous over homoharmonic computations; i.e., heteroharmonic neurons receive information from call and echo in different frequency-bands which helps to avoid jamming between pulse and echo signals. Heteroharmonic neurons have been found in two species of the family Mormoopidae (Pteronotus parnellii and Pteronotus quadridens) and in Rhinolophus rouxi. Recently, it was proposed that heteroharmonic target-range computations are a primitive feature of the genus Pteronotus that was preserved in the evolution of the genus. Here, we review recent findings on the evolution of echolocation in Mormoopidae, and try to link those findings to the evolution of the heteroharmonic computation strategy (HtHCS). We stress the hypothesis that the ability to perform heteroharmonic computations evolved separately from the ability of using long constant-frequency echolocation calls, high duty cycle echolocation, and Doppler Shift Compensation. Also, we present the idea that heteroharmonic computations might have been of advantage for categorizing prey size, hunting eared insects, and living in large conspecific colonies. We make five testable predictions that might help future investigations to clarify the evolution of the heteroharmonic echolocation in Mormoopidae and other families. PMID:23781209

Comments on "Killer whale (Orcinus orca) behavioral audiograms" [J. Acoust. Soc. Am. 141, 2387-2398 (2017)].

PubMed

Heffner, Henry E; Heffner, Rickye S

2018-01-01

Branstetter and his colleagues present the audiograms of eight killer whales and provide a comprehensive review of previous killer whale audiograms. In their paper, they say that the present authors have reported a relationship between size and high-frequency hearing but that echolocating cetaceans might be a special case. The purpose of these comments is to clarify that the relationship of a species' high-frequency hearing is not to its size (mass) but to its "functional interaural distance" (a measure of the availability of sound-localization cues). Moreover, it has previously been noted that echolocating animals, cetaceans as well as bats, have extended their high-frequency hearing somewhat beyond the frequencies used by comparable non-echolocators for passive localization.

The beluga whale produces two pulses to form its sonar signal

PubMed Central

Lammers, Marc O.; Castellote, Manuel

2009-01-01

Odontocete cetaceans use biosonar clicks to acoustically probe their aquatic environment with an aptitude unmatched by man-made sonar. A cornerstone of this ability is their use of short, broadband pulses produced in the region of the upper nasal passages. Here we provide empirical evidence that a beluga whale (Delphinapterus leucas) uses two signal generators simultaneously when echolocating. We show that the pulses of the two generators are combined as they are transmitted through the melon to produce a single echolocation click emitted from the front of the animal. Generating two pulses probably offers the beluga the ability to control the energy and frequency distribution of the emitted click and may allow it to acoustically steer its echolocation beam. PMID:19324643

Dynamics of the echolocation beam during prey pursuit in aerial hawking bats

PubMed Central

Jakobsen, Lasse; Olsen, Mads Nedergaard; Surlykke, Annemarie

2015-01-01

In the evolutionary arms race between prey and predator, measures and countermeasures continuously evolve to increase survival on both sides. Bats and moths are prime examples. When exposed to intense ultrasound, eared moths perform dramatic escape behaviors. Vespertilionid and rhinolophid bats broaden their echolocation beam in the final stage of pursuit, presumably as a countermeasure to keep evading moths within their “acoustic field of view.” In this study, we investigated if dynamic beam broadening is a general property of echolocation when catching moving prey. We recorded three species of emballonurid bats, Saccopteryx bilineata, Saccopteryx leptura, and Rhynchonycteris naso, catching airborne insects in the field. The study shows that S. bilineata and S. leptura maintain a constant beam shape during the entire prey pursuit, whereas R. naso broadens the beam by lowering the peak call frequency from 100 kHz during search and approach to 67 kHz in the buzz. Surprisingly, both Saccopteryx bats emit calls with very high energy throughout the pursuit, up to 60 times more than R. naso and Myotis daubentonii (a similar sized vespertilionid), providing them with as much, or more, peripheral “vision” than the vespertilionids, but ensonifying objects far ahead suggesting more clutter. Thus, beam broadening is not a fundamental property of the echolocation system. However, based on the results, we hypothesize that increased peripheral detection is crucial to all aerial hawking bats in the final stages of prey pursuit and speculate that beam broadening is a feature characterizing more advanced echolocation. PMID:26080398

Aural localization of silent objects by active human biosonar: neural representations of virtual echo-acoustic space.

PubMed

Wallmeier, Ludwig; Kish, Daniel; Wiegrebe, Lutz; Flanagin, Virginia L

2015-03-01

Some blind humans have developed the remarkable ability to detect and localize objects through the auditory analysis of self-generated tongue clicks. These echolocation experts show a corresponding increase in 'visual' cortex activity when listening to echo-acoustic sounds. Echolocation in real-life settings involves multiple reflections as well as active sound production, neither of which has been systematically addressed. We developed a virtualization technique that allows participants to actively perform such biosonar tasks in virtual echo-acoustic space during magnetic resonance imaging (MRI). Tongue clicks, emitted in the MRI scanner, are picked up by a microphone, convolved in real time with the binaural impulse responses of a virtual space, and presented via headphones as virtual echoes. In this manner, we investigated the brain activity during active echo-acoustic localization tasks. Our data show that, in blind echolocation experts, activations in the calcarine cortex are dramatically enhanced when a single reflector is introduced into otherwise anechoic virtual space. A pattern-classification analysis revealed that, in the blind, calcarine cortex activation patterns could discriminate left-side from right-side reflectors. This was found in both blind experts, but the effect was significant for only one of them. In sighted controls, 'visual' cortex activations were insignificant, but activation patterns in the planum temporale were sufficient to discriminate left-side from right-side reflectors. Our data suggest that blind and echolocation-trained, sighted subjects may recruit different neural substrates for the same active-echolocation task. © 2015 Federation of European Neuroscience Societies and John Wiley & Sons Ltd.

Dynamic Echo Information Guides Flight in the Big Brown Bat

PubMed Central

Warnecke, Michaela; Lee, Wu-Jung; Krishnan, Anand; Moss, Cynthia F.

2016-01-01

Animals rely on sensory feedback from their environment to guide locomotion. For instance, visually guided animals use patterns of optic flow to control their velocity and to estimate their distance to objects (e.g., Srinivasan et al., 1991, 1996). In this study, we investigated how acoustic information guides locomotion of animals that use hearing as a primary sensory modality to orient and navigate in the dark, where visual information is unavailable. We studied flight and echolocation behaviors of big brown bats as they flew under infrared illumination through a corridor with walls constructed from a series of individual vertical wooden poles. The spacing between poles on opposite walls of the corridor was experimentally manipulated to create dense/sparse and balanced/imbalanced spatial structure. The bats’ flight trajectories and echolocation signals were recorded with high-speed infrared motion-capture cameras and ultrasound microphones, respectively. As bats flew through the corridor, successive biosonar emissions returned cascades of echoes from the walls of the corridor. The bats flew through the center of the corridor when the pole spacing on opposite walls was balanced and closer to the side with wider pole spacing when opposite walls had an imbalanced density. Moreover, bats produced shorter duration echolocation calls when they flew through corridors with smaller spacing between poles, suggesting that clutter density influences features of the bat’s sonar signals. Flight speed and echolocation call rate did not, however, vary with dense and sparse spacing between the poles forming the corridor walls. Overall, these data demonstrate that bats adapt their flight and echolocation behavior dynamically when flying through acoustically complex environments. PMID:27199690

Morphology of the Nasal Apparatus in Pygmy (Kogia Breviceps) and Dwarf (K. Sima) Sperm Whales.

PubMed

Thornton, Steven W; Mclellan, William A; Rommel, Sentiel A; Dillaman, Richard M; Nowacek, Douglas P; Koopman, Heather N; Pabst, D Ann

2015-07-01

Odontocete echolocation clicks are generated by pneumatically driven phonic lips within the nasal passage, and propagated through specialized structures within the forehead. This study investigated the highly derived echolocation structures of the pygmy (Kogia breviceps) and dwarf (K. sima) sperm whales through careful dissections (N = 18 K. breviceps, 6 K. sima) and histological examinations (N = 5 K. breviceps). This study is the first to show that the entire kogiid sound production and transmission pathway is acted upon by complex facial muscles (likely derivations of the m. maxillonasolabialis). Muscles appear capable of tensing and separating the solitary pair of phonic lips, which would control echolocation click frequencies. The phonic lips are enveloped by the "vocal cap," a morphologically complex, connective tissue structure unique to kogiids. Extensive facial muscles appear to control the position of this structure and its spatial relationship to the phonic lips. The vocal cap's numerous air crypts suggest that it may reflect sounds. Muscles encircling the connective tissue case that surrounds the spermaceti organ may change its shape and/or internal pressure. These actions may influence the acoustic energy transmitted from the phonic lips, through this lipid body, to the melon. Facial and rostral muscles act upon the length of the melon, suggesting that the sound "beam" can be focused as it travels through the melon and into the environment. This study suggests that the kogiid echolocation system is highly tunable. Future acoustic studies are required to test these hypotheses and gain further insight into the kogiid echolocation system. © 2015 Wiley Periodicals, Inc.

The Acuity of Echolocation: Spatial Resolution in Sighted Persons Compared to the Performance of an Expert Who Is Blind

ERIC Educational Resources Information Center

Teng, Santani; Whitney, David

2011-01-01

Echolocation is a specialized application of spatial hearing that uses reflected auditory information to localize objects and represent the external environment. Although it has been documented extensively in nonhuman species, such as bats and dolphins, its use by some persons who are blind as a navigation and object-identification aid has…

Action Enhances Acoustic Cues for 3-D Target Localization by Echolocating Bats

PubMed Central

Wohlgemuth, Melville J.

2016-01-01

Under natural conditions, animals encounter a barrage of sensory information from which they must select and interpret biologically relevant signals. Active sensing can facilitate this process by engaging motor systems in the sampling of sensory information. The echolocating bat serves as an excellent model to investigate the coupling between action and sensing because it adaptively controls both the acoustic signals used to probe the environment and movements to receive echoes at the auditory periphery. We report here that the echolocating bat controls the features of its sonar vocalizations in tandem with the positioning of the outer ears to maximize acoustic cues for target detection and localization. The bat’s adaptive control of sonar vocalizations and ear positioning occurs on a millisecond timescale to capture spatial information from arriving echoes, as well as on a longer timescale to track target movement. Our results demonstrate that purposeful control over sonar sound production and reception can serve to improve acoustic cues for localization tasks. This finding also highlights the general importance of movement to sensory processing across animal species. Finally, our discoveries point to important parallels between spatial perception by echolocation and vision. PMID:27608186

Classification of echolocation clicks from odontocetes in the Southern California Bight.

PubMed

Roch, Marie A; Klinck, Holger; Baumann-Pickering, Simone; Mellinger, David K; Qui, Simon; Soldevilla, Melissa S; Hildebrand, John A

2011-01-01

This study presents a system for classifying echolocation clicks of six species of odontocetes in the Southern California Bight: Visually confirmed bottlenose dolphins, short- and long-beaked common dolphins, Pacific white-sided dolphins, Risso's dolphins, and presumed Cuvier's beaked whales. Echolocation clicks are represented by cepstral feature vectors that are classified by Gaussian mixture models. A randomized cross-validation experiment is designed to provide conditions similar to those found in a field-deployed system. To prevent matched conditions from inappropriately lowering the error rate, echolocation clicks associated with a single sighting are never split across the training and test data. Sightings are randomly permuted before assignment to folds in the experiment. This allows different combinations of the training and test data to be used while keeping data from each sighting entirely in the training or test set. The system achieves a mean error rate of 22% across 100 randomized three-fold cross-validation experiments. Four of the six species had mean error rates lower than the overall mean, with the presumed Cuvier's beaked whale clicks showing the best performance (<2% error rate). Long-beaked common and bottlenose dolphins proved the most difficult to classify, with mean error rates of 53% and 68%, respectively.

Psychophysics of human echolocation.

PubMed

Schörnich, Sven; Wallmeier, Ludwig; Gessele, Nikodemus; Nagy, Andreas; Schranner, Michael; Kish, Daniel; Wiegrebe, Lutz

2013-01-01

The skills of some blind humans orienting in their environment through the auditory analysis of reflections from self-generated sounds have received only little scientific attention to date. Here we present data from a series of formal psychophysical experiments with sighted subjects trained to evaluate features of a virtual echo-acoustic space, allowing for rigid and fine-grain control of the stimulus parameters. The data show how subjects shape both their vocalisations and auditory analysis of the echoes to serve specific echo-acoustic tasks. First, we show that humans can echo-acoustically discriminate target distances with a resolution of less than 1 m for reference distances above 3.4 m. For a reference distance of 1.7 m, corresponding to an echo delay of only 10 ms, distance JNDs were typically around 0.5 m. Second, we explore the interplay between the precedence effect and echolocation. We show that the strong perceptual asymmetry between lead and lag is weakened during echolocation. Finally, we show that through the auditory analysis of self-generated sounds, subjects discriminate room-size changes as small as 10%.In summary, the current data confirm the practical efficacy of human echolocation, and they provide a rigid psychophysical basis for addressing its neural foundations.

Gleaning bat echolocation calls do not elicit antipredator behaviour in the Pacific field cricket, Teleogryllus oceanicus (Orthoptera: Gryllidae).

PubMed

ter Hofstede, Hannah M; Killow, Joanne; Fullard, James H

2009-08-01

Bats that glean prey (capture them from surfaces) produce relatively inconspicuous echolocation calls compared to aerially foraging bats and could therefore be difficult predators to detect, even for insects with ultrasound sensitive ears. In the cricket Teleogryllus oceanicus, an auditory interneuron (AN2) responsive to ultrasound is known to elicit turning behaviour, but only when the cricket is in flight. Turning would not save a cricket from a gleaning bat so we tested the hypothesis that AN2 elicits more appropriate antipredator behaviours when crickets are on the ground. The echolocation calls of Nyctophilus geoffroyi, a sympatric gleaning bat, were broadcast to singing male and walking female T. oceanicus. Males did not cease singing and females did not pause walking more than usual in response to the bat calls up to intensities of 82 dB peSPL. Extracellular recordings from the cervical connective revealed that the echolocation calls elicited AN2 action potentials at high firing rates, indicating that the crickets could hear these stimuli. AN2 appears to elicit antipredator behaviour only in flight, and we discuss possible reasons for this context-dependent function.

Target distance-dependent variation of hearing sensitivity during echolocation in a false killer whale.

PubMed

Supin, Alexander Ya; Nachtigall, Paul E; Breese, Marlee

2010-06-01

Evidence of varying hearing sensitivity according to the target distance was obtained in a false killer whale Pseudorca crassidens during echolocation. Auditory evoked potentials (AEPs) triggered by echolocation clicks were recorded. The target distance varied from 1 to 6 m. The records contained AEPs to the self-heard emitted click and AEPs to the echoes. Mean level of echolocation clicks depended on distance (the longer the distance, the higher the click level), however, the effect of click level on AEP amplitude was eliminated by extracting AEPs to clicks of certain particular levels. The amplitude of the echo-provoked AEP was almost independent of distance, however, the amplitude of the AEP to the emitted click, did depend on distance within a range from 1 to 4 m: the longer the distance, the higher the amplitude. The latter result is interpreted as confirmational evidence that the animal is capable of varying hearing sensitivity according to target distance. The variation of hearing sensitivity may help to compensate for the echo attenuation with distance; as a secondary effect, this variation manifested itself in a variation of the amplitude of the AEP to emitted clicks.

Cortical neurons of bats respond best to echoes from nearest targets when listening to natural biosonar multi-echo streams.

PubMed

Beetz, M Jerome; Hechavarría, Julio C; Kössl, Manfred

2016-10-27

Bats orientate in darkness by listening to echoes from their biosonar calls, a behaviour known as echolocation. Recent studies showed that cortical neurons respond in a highly selective manner when stimulated with natural echolocation sequences that contain echoes from single targets. However, it remains unknown how cortical neurons process echolocation sequences containing echo information from multiple objects. In the present study, we used echolocation sequences containing echoes from three, two or one object separated in the space depth as stimuli to study neuronal activity in the bat auditory cortex. Neuronal activity was recorded with multi-electrode arrays placed in the dorsal auditory cortex, where neurons tuned to target-distance are found. Our results show that target-distance encoding neurons are mostly selective to echoes coming from the closest object, and that the representation of echo information from distant objects is selectively suppressed. This suppression extends over a large part of the dorsal auditory cortex and may override possible parallel processing of multiple objects. The presented data suggest that global cortical suppression might establish a cortical "default mode" that allows selectively focusing on close obstacle even without active attention from the animals.

Delayed response and biosonar perception explain movement coordination in trawling bats.

PubMed

Giuggioli, Luca; McKetterick, Thomas J; Holderied, Marc

2015-03-01

Animal coordinated movement interactions are commonly explained by assuming unspecified social forces of attraction, repulsion and alignment with parameters drawn from observed movement data. Here we propose and test a biologically realistic and quantifiable biosonar movement interaction mechanism for echolocating bats based on spatial perceptual bias, i.e. actual sound field, a reaction delay, and observed motor constraints in speed and acceleration. We found that foraging pairs of bats flying over a water surface swapped leader-follower roles and performed chases or coordinated manoeuvres by copying the heading a nearby individual has had up to 500 ms earlier. Our proposed mechanism based on the interplay between sensory-motor constraints and delayed alignment was able to recreate the observed spatial actor-reactor patterns. Remarkably, when we varied model parameters (response delay, hearing threshold and echolocation directionality) beyond those observed in nature, the spatio-temporal interaction patterns created by the model only recreated the observed interactions, i.e. chases, and best matched the observed spatial patterns for just those response delays, hearing thresholds and echolocation directionalities found to be used by bats. This supports the validity of our sensory ecology approach of movement coordination, where interacting bats localise each other by active echolocation rather than eavesdropping.

Cortical neurons of bats respond best to echoes from nearest targets when listening to natural biosonar multi-echo streams

PubMed Central

Beetz, M. Jerome; Hechavarría, Julio C.; Kössl, Manfred

2016-01-01

Bats orientate in darkness by listening to echoes from their biosonar calls, a behaviour known as echolocation. Recent studies showed that cortical neurons respond in a highly selective manner when stimulated with natural echolocation sequences that contain echoes from single targets. However, it remains unknown how cortical neurons process echolocation sequences containing echo information from multiple objects. In the present study, we used echolocation sequences containing echoes from three, two or one object separated in the space depth as stimuli to study neuronal activity in the bat auditory cortex. Neuronal activity was recorded with multi-electrode arrays placed in the dorsal auditory cortex, where neurons tuned to target-distance are found. Our results show that target-distance encoding neurons are mostly selective to echoes coming from the closest object, and that the representation of echo information from distant objects is selectively suppressed. This suppression extends over a large part of the dorsal auditory cortex and may override possible parallel processing of multiple objects. The presented data suggest that global cortical suppression might establish a cortical “default mode” that allows selectively focusing on close obstacle even without active attention from the animals. PMID:27786252

Recording and quantification of ultrasonic echolocation clicks from free-ranging toothed whales

NASA Astrophysics Data System (ADS)

Madsen, P. T.; Wahlberg, M.

2007-08-01

Toothed whales produce short, ultrasonic clicks of high directionality and source level to probe their environment acoustically. This process, termed echolocation, is to a large part governed by the properties of the emitted clicks. Therefore derivation of click source parameters from free-ranging animals is of increasing importance to understand both how toothed whales use echolocation in the wild and how they may be monitored acoustically. This paper addresses how source parameters can be derived from free-ranging toothed whales in the wild using calibrated multi-hydrophone arrays and digital recorders. We outline the properties required of hydrophones, amplifiers and analog to digital converters, and discuss the problems of recording echolocation clicks on the axis of a directional sound beam. For accurate localization the hydrophone array apertures must be adapted and scaled to the behavior of, and the range to, the clicking animal, and precise information on hydrophone locations is critical. We provide examples of localization routines and outline sources of error that lead to uncertainties in localizing clicking animals in time and space. Furthermore we explore approaches to time series analysis of discrete versions of toothed whale clicks that are meaningful in a biosonar context.

Echolocation click rates and behavior of foraging Hawaiian spinner dolphins

NASA Astrophysics Data System (ADS)

Benoit-Bird, Kelly J.; Au, Whitlow W. L.

2004-05-01

Groups of spinner dolphins work together to actively aggregate small animals in the deep-scattering layer that serve as their prey. Detailed information on dolphin foraging behavior, obtained with a 200-kHz multibeam sonar (Simrad MS2000), made it possible to correlate echolocation and foraging. Fifty-six groups of spinner dolphins foraging at night within a midwater micronekton sound-scattering layer were observed with the sonar. During sonar surveys, the rates of whistles and echolocation clicks were measured using four hydrophones at 6-m depth intervals. Significant differences in click rates were found between depths and between the different stages of foraging. Groups of foraging dolphins ranged in size from 16 to 28 dolphins. Click rates were not significantly affected by the number of dolphins in a foraging group. Contrary to initial predictions, click rates were relatively low when sonar data indicated that pairs of dolphins were actively feeding. Highest echolocation rates occurred within the scattering layer, during transitions between foraging states. Whistles were only detected when dolphins were not in a foraging formation and when animals were surfacing. This suggests clicks may be used directly or indirectly to cue group movement during foraging.

Acoustic behavior of echolocating bats in complex environments

NASA Astrophysics Data System (ADS)

Moss, Cynthia; Ghose, Kaushik; Jensen, Marianne; Surlykke, Annemarie

2004-05-01

The echolocating bat controls the direction of its sonar beam, just as visually dominant animals control the movement of their eyes to foveate targets of interest. The sonar beam aim of the echolocating bat can therefore serve as an index of the animal's attention to objects in the environment. Until recently, spatial attention has not been studied in the context of echolocation, perhaps due to the difficulty in obtaining an objective measure. Here, we describe measurements of the bat's sonar beam aim, serving as an index of acoustic gaze and attention to objects, in tasks that require localization of obstacles and insect prey. Measurements of the bat's sonar beam aim are taken from microphone array recordings of vocal signals produced by a free-flying bat under experimentally controlled conditions. In some situations, the animal relies on spatial memory over reflected sounds, perhaps because its perceptual system cannot easily organize cascades of echoes from obstacles and prey. This highlights the complexity of the bat's orientation behavior, which can alternate between active sensing and spatial memory systems. The bat's use of spatial memory for orientation also will be addressed in this talk. [Work supported by NSF-IBN-0111973 and the Danish Research Council.

Detection and classification of underwater targets by echolocating dolphins

NASA Astrophysics Data System (ADS)

Au, Whitlow

2003-10-01

Many experiments have been performed with echolocating dolphins to determine their target detection and discrimination capabilities. Target detection experiments have been performed in a naturally noisy environment, with masking noise and with both phantom echoes and masking noise, and in reverberation. The echo energy to rms noise spectral density for the Atlantic bottlenose dolphin (Tursiops truncatus) at the 75% correct response threshold is approximately 7.5 dB whereas for the beluga whale (Delphinapterus leucas) the threshold is approximately 1 dB. The dolphin's detection threshold in reverberation is approximately 2.5 dB vs 2 dB for the beluga. The difference in performance between species can probably be ascribed to differences in how both species perceived the task. The bottlenose dolphin may be performing a combination detection/discrimination task whereas the beluga may be performing a simple detection task. Echolocating dolphins also have the capability to make fine discriminate of target properties such as wall thickness difference of water-filled cylinders and material differences in metallic plates. The high resolution property of the animal's echolocation signals and the high dynamic range of its auditory system are important factors in their outstanding discrimination capabilities.

Object representation in the bottlenose dolphin (Tursiops truncatus): integration of visual and echoic information.

PubMed

Harley, H E; Roitblat, H L; Nachtigall, P E

1996-04-01

A dolphin performed a 3-alternative matching-to-sample task in different modality conditions (visual/echoic, both vision and echolocation: visual, vision only; echoic, echolocation only). In Experiment 1, training occurred in the dual-modality (visual/echoic) condition. Choice accuracy in tests of all conditions was above chance without further training. In Experiment 2, unfamiliar objects with complementary similarity relations in vision and echolocation were presented in single-modality conditions until accuracy was about 70%. When tested in the visual/echoic condition, accuracy immediately rose (95%), suggesting integration across modalities. In Experiment 3, conditions varied between presentation of sample and alternatives. The dolphin successfully matched familiar objects in the cross-modal conditions. These data suggest that the dolphin has an object-based representational system.

Regulation of bat echolocation pulse acoustics by striatal dopamine.

PubMed

Tressler, Jedediah; Schwartz, Christine; Wellman, Paul; Hughes, Samuel; Smotherman, Michael

2011-10-01

The ability to control the bandwidth, amplitude and duration of echolocation pulses is a crucial aspect of echolocation performance but few details are known about the neural mechanisms underlying the control of these voice parameters in any mammal. The basal ganglia (BG) are a suite of forebrain nuclei centrally involved in sensory-motor control and are characterized by their dependence on dopamine. We hypothesized that pharmacological manipulation of brain dopamine levels could reveal how BG circuits might influence the acoustic structure of bat echolocation pulses. A single intraperitoneal injection of a low dose (5 mg kg(-1)) of the neurotoxin 1-methyl-4-phenylpyridine (MPTP), which selectively targets dopamine-producing cells of the substantia nigra, produced a rapid degradation in pulse acoustic structure and eliminated the bat's ability to make compensatory changes in pulse amplitude in response to background noise, i.e. the Lombard response. However, high-performance liquid chromatography (HPLC) measurements of striatal dopamine concentrations revealed that the main effect of MPTP was a fourfold increase rather than the predicted decrease in striatal dopamine levels. After first using autoradiographic methods to confirm the presence and location of D(1)- and D(2)-type dopamine receptors in the bat striatum, systemic injections of receptor subtype-specific agonists showed that MPTP's effects on pulse acoustics were mimicked by a D(2)-type dopamine receptor agonist (Quinpirole) but not by a D(1)-type dopamine receptor agonist (SKF82958). The results suggest that BG circuits have the capacity to influence echolocation pulse acoustics, particularly via D(2)-type dopamine receptor-mediated pathways, and may therefore represent an important mechanism for vocal control in bats.

Bat guilds, a concept to classify the highly diverse foraging and echolocation behaviors of microchiropteran bats

PubMed Central

Denzinger, Annette; Schnitzler, Hans-Ulrich

2013-01-01

Throughout evolution the foraging and echolocation behaviors as well as the motor systems of bats have been adapted to the tasks they have to perform while searching and acquiring food. When bats exploit the same class of environmental resources in a similar way, they perform comparable tasks and thus share similar adaptations independent of their phylogeny. Species with similar adaptations are assigned to guilds or functional groups. Habitat type and foraging mode mainly determine the foraging tasks and thus the adaptations of bats. Therefore, we use habitat type and foraging mode to define seven guilds. The habitat types open, edge and narrow space are defined according to the bats' echolocation behavior in relation to the distance between bat and background or food item and background. Bats foraging in the aerial, trawling, flutter detecting, or active gleaning mode use only echolocation to acquire their food. When foraging in the passive gleaning mode bats do not use echolocation but rely on sensory cues from the food item to find it. Bat communities often comprise large numbers of species with a high diversity in foraging areas, foraging modes, and diets. The assignment of species living under similar constraints into guilds identifies patterns of community structure and helps to understand the factors that underlie the organization of highly diverse bat communities. Bat species from different guilds do not compete for food as they differ in their foraging behavior and in the environmental resources they use. However, sympatric living species belonging to the same guild often exploit the same class of resources. To avoid competition they should differ in their niche dimensions. The fine grain structure of bat communities below the rather coarse classification into guilds is determined by mechanisms that result in niche partitioning. PMID:23840190

Regulation of bat echolocation pulse acoustics by striatal dopamine

PubMed Central

Tressler, Jedediah; Schwartz, Christine; Wellman, Paul; Hughes, Samuel; Smotherman, Michael

2011-01-01

SUMMARY The ability to control the bandwidth, amplitude and duration of echolocation pulses is a crucial aspect of echolocation performance but few details are known about the neural mechanisms underlying the control of these voice parameters in any mammal. The basal ganglia (BG) are a suite of forebrain nuclei centrally involved in sensory-motor control and are characterized by their dependence on dopamine. We hypothesized that pharmacological manipulation of brain dopamine levels could reveal how BG circuits might influence the acoustic structure of bat echolocation pulses. A single intraperitoneal injection of a low dose (5 mg kg–1) of the neurotoxin 1-methyl-4-phenylpyridine (MPTP), which selectively targets dopamine-producing cells of the substantia nigra, produced a rapid degradation in pulse acoustic structure and eliminated the bat's ability to make compensatory changes in pulse amplitude in response to background noise, i.e. the Lombard response. However, high-performance liquid chromatography (HPLC) measurements of striatal dopamine concentrations revealed that the main effect of MPTP was a fourfold increase rather than the predicted decrease in striatal dopamine levels. After first using autoradiographic methods to confirm the presence and location of D1- and D2-type dopamine receptors in the bat striatum, systemic injections of receptor subtype-specific agonists showed that MPTP's effects on pulse acoustics were mimicked by a D2-type dopamine receptor agonist (Quinpirole) but not by a D1-type dopamine receptor agonist (SKF82958). The results suggest that BG circuits have the capacity to influence echolocation pulse acoustics, particularly via D2-type dopamine receptor-mediated pathways, and may therefore represent an important mechanism for vocal control in bats. PMID:21900471

Flexible echolocation behavior of trawling bats during approach of continuous or transient prey cues

PubMed Central

Übernickel, Kirstin; Tschapka, Marco; Kalko, Elisabeth K. V.

2013-01-01

Trawling bats use echolocation not only to detect and classify acoustically continuous cues originated from insects at and above water surfaces, but also to detect small water-dwelling prey items breaking the water surface for a very short time, producing only transient cues to be perceived acoustically. Generally, bats need to adjust their echolocation behavior to the specific task on hand, and because of the diversity of prey cues they use in hunting, trawling bats should be highly flexible in their echolocation behavior. We studied the adaptations in the behavior of Noctilio leporinus when approaching either a continuous cue or a transient cue that disappeared during the approach of the bat. Normally the bats reacted by dipping their feet in the water at the cue location. We found that the bats typically started to adapt their calling behavior at approximately 410 ms before prey contact in continuous cue trials, but were also able to adapt their approach behavior to stimuli onsets as short as 177 ms before contact, within a minimum reaction time of 50.9 ms in response to transient cues. In both tasks the approach phase ended between 32 and 53 ms before prey contact. Call emission always continued after the end of the approach phase until around prey contact. In some failed capture attempts, call emission did not cease at all after prey contact. Probably bats used spatial memory to dip at the original location of the transient cue after its disappearance. The duration of the pointed dips was significantly longer in transient cue trials than in continuous cue trials. Our results suggest that trawling bats possess the ability to modify their generally rather stereotyped echolocation behavior during approaches within very short reaction times depending on the sensory information available. PMID:23675352

Echolocating Big Brown Bats, Eptesicus fuscus, Modulate Pulse Intervals to Overcome Range Ambiguity in Cluttered Surroundings

PubMed Central

Wheeler, Alyssa R.; Fulton, Kara A.; Gaudette, Jason E.; Simmons, Ryan A.; Matsuo, Ikuo; Simmons, James A.

2016-01-01

Big brown bats (Eptesicus fuscus) emit trains of brief, wideband frequency-modulated (FM) echolocation sounds and use echoes of these sounds to orient, find insects, and guide flight through vegetation. They are observed to emit sounds that alternate between short and long inter-pulse intervals (IPIs), forming sonar sound groups. The occurrence of these strobe groups has been linked to flight in cluttered acoustic environments, but how exactly bats use sonar sound groups to orient and navigate is still a mystery. Here, the production of sound groups during clutter navigation was examined. Controlled flight experiments were conducted where the proximity of the nearest obstacles was systematically decreased while the extended scene was kept constant. Four bats flew along a corridor of varying widths (100, 70, and 40 cm) bounded by rows of vertically hanging plastic chains while in-flight echolocation calls were recorded. Bats shortened their IPIs for more rapid spatial sampling and also grouped their sounds more tightly when flying in narrower corridors. Bats emitted echolocation calls with progressively shorter IPIs over the course of a flight, and began their flights by emitting shorter starting IPI calls when clutter was denser. The percentage of sound groups containing 3 or more calls increased with increasing clutter proximity. Moreover, IPI sequences having internal structure become more pronounced when corridor width narrows. A novel metric for analyzing the temporal organization of sound sequences was developed, and the results indicate that the time interval between echolocation calls depends heavily on the preceding time interval. The occurrence of specific IPI patterns were dependent upon clutter, which suggests that sonar sound grouping may be an adaptive strategy for coping with pulse-echo ambiguity in cluttered surroundings. PMID:27445723

[Dietary composition, echolocation pulses and morphological measurements of the long-fingered bat Miniopterus fuliginosus (Chiroptera: Vespertilioninae)].

PubMed

Hu, Kai-Liang; Wei, Li; Zhu, Teng-Teng; Wang, Xu-Zhong; Zhang, Li-Biao

2011-04-01

We investigated food (insect) availability in foraging areas utilized by the long-fingered bat Miniopterus fuliginosus using light traps, fish netting and fecal analysis. The dominant preys of M. fuliginosus were Lepidoptera (55%, by volume percent) and Coleoptera (38%) of a relatively large body size. M. fuliginosus has relatively long, narrow wings and a wing span of 6.58+/-0.12 and high wing loading of 9.85+/-0.83 N/m2. The echolocation calls of free flying M. fuliginosus were FM signals, with a pulse duration of 1.45+/-0.06 ms, interpulse interval of 63.08+/-21.55 ms, and low dominant frequency of 44.50+/-2.26 kHz. This study shows that the morphological characteristics and echolocation calls of long-fingered bats are closely linked to their predatory behavior.

The Effects of Behavioral Change in Response to Acoustic Disturbance on the Health of the Population of Blainville’s Beaked Whales (Mesoplodon densirostris) in the Tongue of the Ocean

DTIC Science & Technology

2014-09-30

acoustic data from detection of Md echolocation clicks and sonar, observational data from expert surface observers for group size, surface behavior...on-axis echolocation click source level is in excess of 200 dB), it is therefore highly likely that a GVP will be detected on more than one

Possible age-related hearing loss (presbycusis) and corresponding change in echolocation parameters in a stranded Indo-Pacific humpback dolphin.

PubMed

Li, Songhai; Wang, Ding; Wang, Kexiong; Hoffmann-Kuhnt, Matthias; Fernando, Nimal; Taylor, Elizabeth A; Lin, Wenzhi; Chen, Jialin; Ng, Timothy

2013-11-15

The hearing and echolocation clicks of a stranded Indo-Pacific humpback dolphin (Sousa chinensis) in Zhuhai, China, were studied. This animal had been repeatedly observed in the wild before it was stranded and its age was estimated to be ~40 years. The animal's hearing was measured using a non-invasive auditory evoked potential (AEP) method. Echolocation clicks produced by the dolphin were recorded when the animal was freely swimming in a 7.5 m (width)×22 m (length)×4.8 m (structural depth) pool with a water depth of ~2.5 m. The hearing and echolocation clicks of the studied dolphin were compared with those of a conspecific younger individual, ~13 years of age. The results suggested that the cut-off frequency of the high-frequency hearing of the studied dolphin was ~30-40 kHz lower than that of the younger individual. The peak and centre frequencies of the clicks produced by the older dolphin were ~16 kHz lower than those of the clicks produced by the younger animal. Considering that the older dolphin was ~40 years old, its lower high-frequency hearing range with lower click peak and centre frequencies could probably be explained by age-related hearing loss (presbycusis).

Hearing diversity in moths confronting a neotropical bat assemblage.

PubMed

Cobo-Cuan, Ariadna; Kössl, Manfred; Mora, Emanuel C

2017-09-01

The tympanal ear is an evolutionary acquisition which helps moths survive predation from bats. The greater diversity of bats and echolocation strategies in the Neotropics compared with temperate zones would be expected to impose different sensory requirements on the neotropical moths. However, even given some variability among moth assemblages, the frequencies of best hearing of moths from different climate zones studied to date have been roughly the same: between 20 and 60 kHz. We have analyzed the auditory characteristics of tympanate moths from Cuba, a neotropical island with high levels of bat diversity and a high incidence of echolocation frequencies above those commonly at the upper limit of moths' hearing sensitivity. Moths of the superfamilies Noctuoidea, Geometroidea and Pyraloidea were examined. Audiograms were determined by non-invasively measuring distortion-product otoacoustic emissions. We also quantified the frequency spectrum of the echolocation sounds to which this moth community is exposed. The hearing ranges of moths in our study showed best frequencies between 36 and 94 kHz. High sensitivity to frequencies above 50 kHz suggests that the auditory sensitivity of moths is suited to the sounds used by sympatric echolocating bat fauna. Biodiversity characterizes predators and prey in the Neotropics, but the bat-moth acoustic interaction keeps spectrally matched.

Global warming alters sound transmission: differential impact on the prey detection ability of echolocating bats

PubMed Central

Luo, Jinhong; Koselj, Klemen; Zsebők, Sándor; Siemers, Björn M.; Goerlitz, Holger R.

2014-01-01

Climate change impacts the biogeography and phenology of plants and animals, yet the underlying mechanisms are little known. Here, we present a functional link between rising temperature and the prey detection ability of echolocating bats. The maximum distance for echo-based prey detection is physically determined by sound attenuation. Attenuation is more pronounced for high-frequency sound, such as echolocation, and is a nonlinear function of both call frequency and ambient temperature. Hence, the prey detection ability, and thus possibly the foraging efficiency, of echolocating bats and susceptible to rising temperatures through climate change. Using present-day climate data and projected temperature rises, we modelled this effect for the entire range of bat call frequencies and climate zones around the globe. We show that depending on call frequency, the prey detection volume of bats will either decrease or increase: species calling above a crossover frequency will lose and species emitting lower frequencies will gain prey detection volume, with crossover frequency and magnitude depending on the local climatic conditions. Within local species assemblages, this may cause a change in community composition. Global warming can thus directly affect the prey detection ability of individual bats and indirectly their interspecific interactions with competitors and prey. PMID:24335559

Global warming alters sound transmission: differential impact on the prey detection ability of echolocating bats.

PubMed

Luo, Jinhong; Koselj, Klemen; Zsebok, Sándor; Siemers, Björn M; Goerlitz, Holger R

2014-02-06

Climate change impacts the biogeography and phenology of plants and animals, yet the underlying mechanisms are little known. Here, we present a functional link between rising temperature and the prey detection ability of echolocating bats. The maximum distance for echo-based prey detection is physically determined by sound attenuation. Attenuation is more pronounced for high-frequency sound, such as echolocation, and is a nonlinear function of both call frequency and ambient temperature. Hence, the prey detection ability, and thus possibly the foraging efficiency, of echolocating bats and susceptible to rising temperatures through climate change. Using present-day climate data and projected temperature rises, we modelled this effect for the entire range of bat call frequencies and climate zones around the globe. We show that depending on call frequency, the prey detection volume of bats will either decrease or increase: species calling above a crossover frequency will lose and species emitting lower frequencies will gain prey detection volume, with crossover frequency and magnitude depending on the local climatic conditions. Within local species assemblages, this may cause a change in community composition. Global warming can thus directly affect the prey detection ability of individual bats and indirectly their interspecific interactions with competitors and prey.

Delayed Response and Biosonar Perception Explain Movement Coordination in Trawling Bats

PubMed Central

Giuggioli, Luca; McKetterick, Thomas J.; Holderied, Marc

2015-01-01

Animal coordinated movement interactions are commonly explained by assuming unspecified social forces of attraction, repulsion and alignment with parameters drawn from observed movement data. Here we propose and test a biologically realistic and quantifiable biosonar movement interaction mechanism for echolocating bats based on spatial perceptual bias, i.e. actual sound field, a reaction delay, and observed motor constraints in speed and acceleration. We found that foraging pairs of bats flying over a water surface swapped leader-follower roles and performed chases or coordinated manoeuvres by copying the heading a nearby individual has had up to 500 ms earlier. Our proposed mechanism based on the interplay between sensory-motor constraints and delayed alignment was able to recreate the observed spatial actor-reactor patterns. Remarkably, when we varied model parameters (response delay, hearing threshold and echolocation directionality) beyond those observed in nature, the spatio-temporal interaction patterns created by the model only recreated the observed interactions, i.e. chases, and best matched the observed spatial patterns for just those response delays, hearing thresholds and echolocation directionalities found to be used by bats. This supports the validity of our sensory ecology approach of movement coordination, where interacting bats localise each other by active echolocation rather than eavesdropping. PMID:25811627

Adaptive beam-width control of echolocation sounds by CF-FM bats, Rhinolophus ferrumequinum nippon, during prey-capture flight.

PubMed

Matsuta, Naohiro; Hiryu, Shizuko; Fujioka, Emyo; Yamada, Yasufumi; Riquimaroux, Hiroshi; Watanabe, Yoshiaki

2013-04-01

The echolocation sounds of Japanese CF-FM bats (Rhinolophus ferrumequinum nippon) were measured while the bats pursued a moth (Goniocraspidum pryeri) in a flight chamber. Using a 31-channel microphone array system, we investigated how CF-FM bats adjust pulse direction and beam width according to prey position. During the search and approach phases, the horizontal and vertical beam widths were ±22±5 and ±13±5 deg, respectively. When bats entered the terminal phase approximately 1 m from a moth, distinctive evasive flight by G. pryeri was sometimes observed. Simultaneously, the bats broadened the beam widths of some emissions in both the horizontal (44% of emitted echolocation pulses) and vertical planes (71%). The expanded beam widths were ±36±7 deg (horizontal) and ±30±9 deg (vertical). When moths began evasive flight, the tracking accuracy decreased compared with that during the approach phase. However, in 97% of emissions during the terminal phase, the beam width was wider than the misalignment (the angular difference between the pulse and target directions). These findings indicate that bats actively adjust their beam width to retain the moving target within a spatial echolocation window during the final capture stages.

Transmission beam characteristics of a Risso's dolphin (Grampus griseus).

PubMed

Smith, Adam B; Kloepper, Laura N; Yang, Wei-Cheng; Huang, Wan-Hsiu; Jen, I-Fan; Rideout, Brendan P; Nachtigall, Paul E

2016-01-01

The echolocation system of the Risso's dolphin (Grampus griseus) remains poorly studied compared to other odontocete species. In this study, echolocation signals were recorded from a stationary Risso's dolphin with an array of 16 hydrophones and the two-dimensional beam shape was explored using frequency-dependent amplitude plots. Click source parameters were similar to those already described for this species. Centroid frequency of click signals increased with increasing sound pressure level, while the beamwidth decreased with increasing center frequency. Analysis revealed primarily single-lobed, and occasionally vertically dual-lobed, beam shapes. Overall beam directivity was found to be greater than that of the harbor porpoise, bottlenose dolphin, and a false killer whale. The relationship between frequency content, beam directivity, and head size for this Risso's dolphin deviated from the trend described for other species. These are the first reported measurements of echolocation beam shape and directivity in G. griseus.

Discriminative echolocation in a porpoise, 12

PubMed Central

Turner, Ronald N.; Norris, Kenneth S.

1966-01-01

Operant conditioning techniques were used to establish a discriminative echolocation performance in a porpoise. Pairs of spheres of disparate diameters were presented in an under-water display, and the positions of the spheres were switched according to a scrambled sequence while the blindfolded porpoise responded on a pair of submerged response levers. Responses which identified the momentary state of the display were food-reinforced, while those which did not (errors) produced time out. Errors were then studied in relation to decreased disparity between the spheres. As disparity was decreased, errors which terminated runs of correct responses occurred more frequently and were followed by longer strings of consecutive errors. Increased errors and disruption of a stable pattern of collateral behavior were associated. Since some sources of error other than decreased disparity were present, the porpoise's final performance did not fully reflect the acuity of its echolocation channel. PMID:5964509

Innate recognition of water bodies in echolocating bats.

PubMed

Greif, Stefan; Siemers, Björn M

2010-11-02

In the course of their lives, most animals must find different specific habitat and microhabitat types for survival and reproduction. Yet, in vertebrates, little is known about the sensory cues that mediate habitat recognition. In free flying bats the echolocation of insect-sized point targets is well understood, whereas how they recognize and classify spatially extended echo targets is currently unknown. In this study, we show how echolocating bats recognize ponds or other water bodies that are crucial for foraging, drinking and orientation. With wild bats of 15 different species (seven genera from three phylogenetically distant, large bat families), we found that bats perceived any extended, echo-acoustically smooth surface to be water, even in the presence of conflicting information from other sensory modalities. In addition, naive juvenile bats that had never before encountered a water body showed spontaneous drinking responses from smooth plates. This provides the first evidence for innate recognition of a habitat cue in a mammal.

Does similarity in call structure or foraging ecology explain interspecific information transfer in wild Myotis bats?

PubMed

Hügel, Theresa; van Meir, Vincent; Muñoz-Meneses, Amanda; Clarin, B-Markus; Siemers, Björn M; Goerlitz, Holger R

2017-01-01

Animals can gain important information by attending to the signals and cues of other animals in their environment, with acoustic information playing a major role in many taxa. Echolocation call sequences of bats contain information about the identity and behaviour of the sender which is perceptible to close-by receivers. Increasing evidence supports the communicative function of echolocation within species, yet data about its role for interspecific information transfer is scarce. Here, we asked which information bats extract from heterospecific echolocation calls during foraging. In three linked playback experiments, we tested in the flight room and field if foraging Myotis bats approached the foraging call sequences of conspecifics and four heterospecifics that were similar in acoustic call structure only (acoustic similarity hypothesis), in foraging ecology only (foraging similarity hypothesis), both, or none. Compared to the natural prey capture rate of 1.3 buzzes per minute of bat activity, our playbacks of foraging sequences with 23-40 buzzes/min simulated foraging patches with significantly higher profitability. In the flight room, M. capaccinii only approached call sequences of conspecifics and of the heterospecific M. daubentonii with similar acoustics and foraging ecology. In the field, M. capaccinii and M. daubentonii only showed a weak positive response to those two species. Our results confirm information transfer across species boundaries and highlight the importance of context on the studied behaviour, but cannot resolve whether information transfer in trawling Myotis is based on acoustic similarity only or on a combination of similarity in acoustics and foraging ecology. Animals transfer information, both voluntarily and inadvertently, and within and across species boundaries. In echolocating bats, acoustic call structure and foraging ecology are linked, making echolocation calls a rich source of information about species identity, ecology and activity of the sender, which receivers might exploit to find profitable foraging grounds. We tested in three lab and field experiments if information transfer occurs between bat species and if bats obtain information about ecology from echolocation calls. Myotis capaccinii/daubentonii bats approached call playbacks, but only those from con- and heterospecifics with similar call structure and foraging ecology, confirming interspecific information transfer. Reactions differed between lab and field, emphasising situation-dependent differences in animal behaviour, the importance of field research, and the need for further studies on the underlying mechanism of information transfer and the relative contributions of acoustic and ecological similarity.

Energy spectrum analysis - A model of echolocation processing. [in animals

NASA Technical Reports Server (NTRS)

Johnson, R. A.; Titlebaum, E. L.

1976-01-01

The paper proposes a frequency domain approach based on energy spectrum analysis of the combination of a signal and its echoes as the processing mechanism for the echolocation process used by bats and other animals. The mechanism is a generalized wide-band one and can account for the large diversity of wide-band signals used for orientation. The coherency in the spectrum of the signal-echo combination is shown to be equivalent to correlation.

Acoustic Scattering of Broadband Echolocation Signals from Prey of Blainville’s Beaked Whales: Modeling and Analysis

DTIC Science & Technology

2006-09-01

biosonar , summarized in the following paragraphs, provides context for this study. 1.1.1 Echolocation in bats Researchers have debated for over two...centuries the capabilities of certain species of animals to use biosonar in orientation, communication, and prey capture. As early as 1793 Italian...marine organisms In complement to the research on the biosonar systems of these capable predators, a concurrent body of research has been conducted on

Clicking in a Killer Whale Habitat: Narrow-Band, High-Frequency Biosonar Clicks of Harbour Porpoise (Phocoena phocoena) and Dall’s Porpoise (Phocoenoides dalli)

PubMed Central

Kyhn, Line A.; Tougaard, Jakob; Beedholm, Kristian; Jensen, Frants H.; Ashe, Erin; Williams, Rob; Madsen, Peter T.

2013-01-01

Odontocetes produce a range of different echolocation clicks but four groups in different families have converged on producing the same stereotyped narrow band high frequency (NBHF) click. In microchiropteran bats, sympatric species have evolved the use of different acoustic niches and subtly different echolocation signals to avoid competition among species. In this study, we examined whether similar adaptations are at play among sympatric porpoise species that use NBHF echolocation clicks. We used a six-element hydrophone array to record harbour and Dall’s porpoises in British Columbia (BC), Canada, and harbour porpoises in Denmark. The click source properties of all porpoise groups were remarkably similar and had an average directivity index of 25 dB. Yet there was a small, but consistent and significant 4 kHz difference in centroid frequency between sympatric Dall’s (137±3 kHz) and Canadian harbour porpoises (141±2 kHz). Danish harbour porpoise clicks (136±3 kHz) were more similar to Dall’s porpoise than to their conspecifics in Canada. We suggest that the spectral differences in echolocation clicks between the sympatric porpoises are consistent with evolution of a prezygotic isolating barrier (i.e., character displacement) to avoid hybridization of sympatric species. In practical terms, these spectral differences have immediate application to passive acoustic monitoring. PMID:23723996

Clicking in a killer whale habitat: narrow-band, high-frequency biosonar clicks of harbour porpoise (Phocoena phocoena) and Dall's porpoise (Phocoenoides dalli).

PubMed

Kyhn, Line A; Tougaard, Jakob; Beedholm, Kristian; Jensen, Frants H; Ashe, Erin; Williams, Rob; Madsen, Peter T

2013-01-01

Odontocetes produce a range of different echolocation clicks but four groups in different families have converged on producing the same stereotyped narrow band high frequency (NBHF) click. In microchiropteran bats, sympatric species have evolved the use of different acoustic niches and subtly different echolocation signals to avoid competition among species. In this study, we examined whether similar adaptations are at play among sympatric porpoise species that use NBHF echolocation clicks. We used a six-element hydrophone array to record harbour and Dall's porpoises in British Columbia (BC), Canada, and harbour porpoises in Denmark. The click source properties of all porpoise groups were remarkably similar and had an average directivity index of 25 dB. Yet there was a small, but consistent and significant 4 kHz difference in centroid frequency between sympatric Dall's (137±3 kHz) and Canadian harbour porpoises (141±2 kHz). Danish harbour porpoise clicks (136±3 kHz) were more similar to Dall's porpoise than to their conspecifics in Canada. We suggest that the spectral differences in echolocation clicks between the sympatric porpoises are consistent with evolution of a prezygotic isolating barrier (i.e., character displacement) to avoid hybridization of sympatric species. In practical terms, these spectral differences have immediate application to passive acoustic monitoring.

Three-dimensional tracking of Cuvier's beaked whales' echolocation sounds using nested hydrophone arrays.

PubMed

Gassmann, Martin; Wiggins, Sean M; Hildebrand, John A

2015-10-01

Cuvier's beaked whales (Ziphius cavirostris) were tracked using two volumetric small-aperture (∼1 m element spacing) hydrophone arrays, embedded into a large-aperture (∼1 km element spacing) seafloor hydrophone array of five nodes. This array design can reduce the minimum number of nodes that are needed to record the arrival of a strongly directional echolocation sound from 5 to 2, while providing enough time-differences of arrivals for a three-dimensional localization without depending on any additional information such as multipath arrivals. To illustrate the capabilities of this technique, six encounters of up to three Cuvier's beaked whales were tracked over a two-month recording period within an area of 20 km(2) in the Southern California Bight. Encounter periods ranged from 11 min to 33 min. Cuvier's beaked whales were found to reduce the time interval between echolocation clicks while alternating between two inter-click-interval regimes during their descent towards the seafloor. Maximum peak-to-peak source levels of 179 and 224 dB re 1 μPa @ 1 m were estimated for buzz sounds and on-axis echolocation clicks (directivity index = 30 dB), respectively. Source energy spectra of the on-axis clicks show significant frequency components between 70 and 90 kHz, in addition to their typically noted FM upsweep at 40-60 kHz.

Adaptive evolution of tight junction protein claudin-14 in echolocating whales.

PubMed

Xu, Huihui; Liu, Yang; He, Guimei; Rossiter, Stephen J; Zhang, Shuyi

2013-11-10

Toothed whales and bats have independently evolved specialized ultrasonic hearing for echolocation. Recent findings have suggested that several genes including Prestin, Tmc1, Pjvk and KCNQ4 appear to have undergone molecular adaptations associated with the evolution of this ultrasonic hearing in mammals. Here we studied the hearing gene Cldn14, which encodes the claudin-14 protein and is a member of tight junction proteins that functions in the organ of Corti in the inner ear to maintain a cationic gradient between endolymph and perilymph. Particular mutations in human claudin-14 give rise to non-syndromic deafness, suggesting an essential role in hearing. Our results uncovered two bursts of positive selection, one in the ancestral branch of all toothed whales and a second in the branch leading to the delphinid, phocoenid and ziphiid whales. These two branches are the same as those previously reported to show positive selection in the Prestin gene. Furthermore, as with Prestin, the estimated hearing frequencies of whales significantly correlate with numbers of branch-wise non-synonymous substitutions in Cldn14, but not with synonymous changes. However, in contrast to Prestin, we found no evidence of positive selection in bats. Our findings from Cldn14, and comparisons with Prestin, strongly implicate multiple loci in the acquisition of echolocation in cetaceans, but also highlight possible differences in the evolutionary route to echolocation taken by whales and bats. © 2013.

Sensorimotor Model of Obstacle Avoidance in Echolocating Bats

PubMed Central

Vanderelst, Dieter; Holderied, Marc W.; Peremans, Herbert

2015-01-01

Bat echolocation is an ability consisting of many subtasks such as navigation, prey detection and object recognition. Understanding the echolocation capabilities of bats comes down to isolating the minimal set of acoustic cues needed to complete each task. For some tasks, the minimal cues have already been identified. However, while a number of possible cues have been suggested, little is known about the minimal cues supporting obstacle avoidance in echolocating bats. In this paper, we propose that the Interaural Intensity Difference (IID) and travel time of the first millisecond of the echo train are sufficient cues for obstacle avoidance. We describe a simple control algorithm based on the use of these cues in combination with alternating ear positions modeled after the constant frequency bat Rhinolophus rouxii. Using spatial simulations (2D and 3D), we show that simple phonotaxis can steer a bat clear from obstacles without performing a reconstruction of the 3D layout of the scene. As such, this paper presents the first computationally explicit explanation for obstacle avoidance validated in complex simulated environments. Based on additional simulations modelling the FM bat Phyllostomus discolor, we conjecture that the proposed cues can be exploited by constant frequency (CF) bats and frequency modulated (FM) bats alike. We hypothesize that using a low level yet robust cue for obstacle avoidance allows bats to comply with the hard real-time constraints of this basic behaviour. PMID:26502063

Echolocation in the Risso's dolphin, Grampus griseus

NASA Astrophysics Data System (ADS)

Philips, Jennifer D.; Nachtigall, Paul E.; Au, Whitlow W. L.; Pawloski, Jeffrey L.; Roitblat, Herbert L.

2003-01-01

The Risso's dolphin (Grampus griseus) is an exclusively cephalopod-consuming delphinid with a distinctive vertical indentation along its forehead. To investigate whether or not the species echolocates, a female Risso's dolphin was trained to discriminate an aluminum cylinder from a nylon sphere (experiment 1) or an aluminum sphere (experiment 2) while wearing eyecups and free swimming in an open-water pen in Kaneohe Bay, Hawaii. The dolphin completed the task with little difficulty despite being blindfolded. Clicks emitted by the dolphin were acquired at average amplitudes of 192.6 dB re 1 μPa, with estimated sources levels up to 216 dB re 1 μPa-1 m. Clicks were acquired with peak frequencies as high as 104.7 kHz (Mfp=47.9 kHz), center frequencies as high as 85.7 kHz (Mf0=56.5 kHz), 3-dB bandwidths up to 94.1 kHz (MBW=39.7 kHz), and root-mean-square bandwidths up to 32.8 kHz (MRMS=23.3 kHz). Click durations were between 40 and 70 μs. The data establish that the Risso's dolphin echolocates, and that, aside from slightly lower amplitudes and frequencies, the clicks emitted by the dolphin were similar to those emitted by other echolocating odontocetes. The particular acoustic and behavioral findings in the study are discussed with respect to the possible direction of the sonar transmission beam of the species.

Variability in echolocation call intensity in a community of horseshoe bats: a role for resource partitioning or communication?

PubMed

Schuchmann, Maike; Siemers, Björn M

2010-09-17

Only recently data on bat echolocation call intensities is starting to accumulate. Yet, intensity is an ecologically crucial parameter, as it determines the extent of the bats' perceptual space and, specifically, prey detection distance. Interspecifically, we thus asked whether sympatric, congeneric bat species differ in call intensities and whether differences play a role for niche differentiation. Specifically, we investigated whether R. mehelyi that calls at a frequency clearly above what is predicted by allometry, compensates for frequency-dependent loss in detection distance by using elevated call intensity. Maximum echolocation call intensities might depend on body size or condition and thus be used as an honest signal of quality for intraspecific communication. We for the first time investigated whether a size-intensity relation is present in echolocating bats. We measured maximum call intensities and frequencies for all five European horseshoe bat species. Maximum intensity differed among species largely due to R. euryale. Furthermore, we found no compensation for frequency-dependent loss in detection distance in R. mehelyi. Intraspecifically, there is a negative correlation between forearm lengths and intensity in R. euryale and a trend for a negative correlation between body condition index and intensity in R. ferrumequinum. In R. hipposideros, females had 8 dB higher intensities than males. There were no correlations with body size or sex differences and intensity for the other species. Based on call intensity and frequency measurements, we estimated echolocation ranges for our study community. These suggest that intensity differences result in different prey detection distances and thus likely play some role for resource access. It is interesting and at first glance counter-intuitive that, where a correlation was found, smaller bats called louder than large individuals. Such negative relationship between size or condition and vocal amplitude may indicate an as yet unknown physiological or sexual selection pressure.

Variability in Echolocation Call Intensity in a Community of Horseshoe Bats: A Role for Resource Partitioning or Communication?

PubMed Central

Schuchmann, Maike; Siemers, Björn M.

2010-01-01

Background Only recently data on bat echolocation call intensities is starting to accumulate. Yet, intensity is an ecologically crucial parameter, as it determines the extent of the bats' perceptual space and, specifically, prey detection distance. Interspecifically, we thus asked whether sympatric, congeneric bat species differ in call intensities and whether differences play a role for niche differentiation. Specifically, we investigated whether R. mehelyi that calls at a frequency clearly above what is predicted by allometry, compensates for frequency-dependent loss in detection distance by using elevated call intensity. Maximum echolocation call intensities might depend on body size or condition and thus be used as an honest signal of quality for intraspecific communication. We for the first time investigated whether a size-intensity relation is present in echolocating bats. Methodology/Principal Findings We measured maximum call intensities and frequencies for all five European horseshoe bat species. Maximum intensity differed among species largely due to R. euryale. Furthermore, we found no compensation for frequency-dependent loss in detection distance in R. mehelyi. Intraspecifically, there is a negative correlation between forearm lengths and intensity in R. euryale and a trend for a negative correlation between body condition index and intensity in R. ferrumequinum. In R. hipposideros, females had 8 dB higher intensities than males. There were no correlations with body size or sex differences and intensity for the other species. Conclusions/Significance Based on call intensity and frequency measurements, we estimated echolocation ranges for our study community. These suggest that intensity differences result in different prey detection distances and thus likely play some role for resource access. It is interesting and at first glance counter-intuitive that, where a correlation was found, smaller bats called louder than large individuals. Such negative relationship between size or condition and vocal amplitude may indicate an as yet unknown physiological or sexual selection pressure. PMID:20862252

Modeling of Habitat and Foraging Behavior of Beaked Whales in the Southern California Bight

DTIC Science & Technology

2012-09-30

patterns of beaked whale echolocation signals in the North Pacific over 26 sites (Figure 3) (Baumann-Pickering et al., 2012a) revealed that Wake Atoll ... Atoll are not considered typical territory for this species and would be a surprising finding. BW43 signal encounters were restricted to the...Roch, M. A., Schnitzler, H. U., and Hildebrand, J. A. (2010). "Echolocation signals of a beaked whale at Palmyra Atoll ," J. Acoust. Soc. Am. 127

Analysis and Modeling of Echolocation Signals Emitted by Mediterranean Bottlenose Dolphins

NASA Astrophysics Data System (ADS)

Greco, Maria; Gini, Fulvio

2006-12-01

We analyzed the echolocation sounds emitted by Mediterranean bottlenose dolphins. We extracted the click trains by visual inspection of the data files recorded along the coast of the Tuscany with the collaboration of the CETUS Research Center. We modeled the extracted sonar clicks as Gaussian or exponential multicomponent signals, we estimated the characteristic parameters and compared the data with the reconstructed signals based on the estimates. Results about the estimation and the data fitting are largely shown in the paper.

Automatic detection of echolocation clicks based on a Gabor model of their waveform.

PubMed

Madhusudhana, Shyam; Gavrilov, Alexander; Erbe, Christine

2015-06-01

Prior research has shown that echolocation clicks of several species of terrestrial and marine fauna can be modelled as Gabor-like functions. Here, a system is proposed for the automatic detection of a variety of such signals. By means of mathematical formulation, it is shown that the output of the Teager-Kaiser Energy Operator (TKEO) applied to Gabor-like signals can be approximated by a Gaussian function. Based on the inferences, a detection algorithm involving the post-processing of the TKEO outputs is presented. The ratio of the outputs of two moving-average filters, a Gaussian and a rectangular filter, is shown to be an effective detection parameter. Detector performance is assessed using synthetic and real (taken from MobySound database) recordings. The detection method is shown to work readily with a variety of echolocation clicks and in various recording scenarios. The system exhibits low computational complexity and operates several times faster than real-time. Performance comparisons are made to other publicly available detectors including pamguard.

Recording animal vocalizations from a UAV: bat echolocation during roost re-entry.

PubMed

Kloepper, Laura N; Kinniry, Morgan

2018-05-17

Unmanned aerial vehicles (UAVs) are rising in popularity for wildlife monitoring, but direct recordings of animal vocalizations have not yet been accomplished, likely due to the noise generated by the UAV. Echolocating bats, especially Tadarida brasiliensis, are good candidates for UAV recording due to their high-speed, high-altitude flight. Here, we use a UAV to record the signals of bats during morning roost re-entry. We designed a UAV to block the noise of the propellers from the receiving microphone, and report on the characteristics of bioacoustic recordings from a UAV. We report the first published characteristics of echolocation signals from bats during group flight and cave re-entry. We found changes in inter-individual time-frequency shape, suggesting that bats may use differences in call design when sensing in complex groups. Furthermore, our first documented successful recordings of animals in their natural habitat demonstrate that UAVs can be important tools for bioacoustic monitoring, and we discuss the ethical considerations for such monitoring.

Buzzing during biosonar-based interception of prey in the delphinids Tursiops truncatus and Pseudorca crassidens.

PubMed

Wisniewska, Danuta M; Johnson, Mark; Nachtigall, Paul E; Madsen, Peter T

2014-12-15

Echolocating bats and toothed whales probe their environment with ultrasonic sound pulses, using returning echoes to navigate and find prey in a process that appears to have resulted from a remarkable convergence of the two taxa. Here, we report the first detailed quantification of echolocation behaviour during prey capture in the most studied delphinid species, a false killer whale and a bottlenose dolphin. Using acoustic DTAGs, we demonstrate that just prior to prey interception these delphinids change their acoustic gaze dramatically by reducing inter-click intervals and output >10-fold in a high repetition rate, low output buzz. Buzz click rates of 250-500 Hz for large but agile animals suggest that sampling rates during capture are scaled with the whale's manoeuvrability. These observations support the growing notion that fast sonar sampling accompanied by a low output level is critical for high rate feedback to inform motor patterns during prey interception in all echolocating toothed whales. © 2014. Published by The Company of Biologists Ltd.

Fractal scaling in bottlenose dolphin (Tursiops truncatus) echolocation: A case study

NASA Astrophysics Data System (ADS)

Perisho, Shaun T.; Kelty-Stephen, Damian G.; Hajnal, Alen; Houser, Dorian; Kuczaj, Stan A., II

2016-02-01

Fractal scaling patterns, which entail a power-law relationship between magnitude of fluctuations in a variable and the scale at which the variable is measured, have been found in many aspects of human behavior. These findings have led to advances in behavioral models (e.g. providing empirical support for cascade-driven theories of cognition) and have had practical medical applications (e.g. providing new methods for early diagnosis of medical conditions). In the present paper, fractal analysis is used to investigate whether similar fractal scaling patterns exist in inter-click interval and peak-peak amplitude measurements of bottlenose dolphin click trains. Several echolocation recordings taken from two male bottlenose dolphins were analyzed using Detrended Fluctuation Analysis and Higuchi's (1988) method for determination of fractal dimension. Both animals were found to exhibit fractal scaling patterns near what is consistent with persistent long range correlations. These findings suggest that recent advances in human cognition and medicine may have important parallel applications to echolocation as well.

How Nectar-Feeding Bats Localize their Food: Echolocation Behavior of Leptonycteris yerbabuenae Approaching Cactus Flowers

PubMed Central

Koblitz, Jens C.; Fleming, Theodore H.; Medellín, Rodrigo A.; Kalko, Elisabeth K. V.; Schnitzler, Hans-Ulrich; Tschapka, Marco

2016-01-01

Nectar-feeding bats show morphological, physiological, and behavioral adaptations for feeding on nectar. How they find and localize flowers is still poorly understood. While scent cues alone allow no precise localization of a floral target, the spatial properties of flower echoes are very precise and could play a major role, particularly at close range. The aim of this study is to understand the role of echolocation for classification and localization of flowers. We compared the approach behavior of Leptonycteris yerbabuenae to flowers of a columnar cactus, Pachycereus pringlei, to that to an acrylic hollow hemisphere that is acoustically conspicuous to bats, but has different acoustic properties and, contrary to the cactus flower, present no scent. For recording the flight and echolocation behaviour we used two infrared video cameras under stroboscopic illumination synchronized with ultrasound recordings. During search flights all individuals identified both targets as a possible food source and initiated an approach flight; however, they visited only the cactus flower. In experiments with the acrylic hemisphere bats aborted the approach at ca. 40–50 cm. In the last instant before the flower visit the bats emitted a long terminal group of 10–20 calls. This is the first report of this behaviour for a nectar-feeding bat. Our findings suggest that L. yerbabuenae use echolocation for classification and localization of cactus flowers and that the echo-acoustic characteristics of the flower guide the bats directly to the flower opening. PMID:27684373

Adaptive changes in echolocation sounds by Pipistrellus abramus in response to artificial jamming sounds.

PubMed

Takahashi, Eri; Hyomoto, Kiri; Riquimaroux, Hiroshi; Watanabe, Yoshiaki; Ohta, Tetsuo; Hiryu, Shizuko

2014-08-15

The echolocation behavior of Pipistrellus abramus during exposure to artificial jamming sounds during flight was investigated. Echolocation pulses emitted by the bats were recorded using a telemetry microphone mounted on the bats' backs, and their adaptation based on acoustic characteristics of emitted pulses was assessed in terms of jamming-avoidance responses (JARs). In experiment 1, frequency-modulated jamming sounds (3 ms duration) mimicking echolocation pulses of P. abramus were prepared. All bats showed significant increases in the terminal frequency of the frequency-modulated pulse by an average of 2.1-4.5 kHz when the terminal frequency of the jamming sounds was lower than the bats' own pulses. This frequency shift was not observed using jamming frequencies that overlapped with or were higher than the bats' own pulses. These findings suggest that JARs in P. abramus are sensitive to the terminal frequency of jamming pulses and that the bats' response pattern was dependent on the slight difference in stimulus frequency. In experiment 2, when bats were repeatedly exposed to a band-limited noise of 70 ms duration, the bats in flight more frequently emitted pulses during silent periods between jamming sounds, suggesting that the bats could actively change the timing of pulse emissions, even during flight, to avoid temporal overlap with jamming sounds. Our findings demonstrate that bats could adjust their vocalized frequency and emission timing during flight in response to acoustic jamming stimuli. © 2014. Published by The Company of Biologists Ltd.

Hearing Sensation Levels of Emitted Biosonar Clicks in an Echolocating Atlantic Bottlenose Dolphin

PubMed Central

Li, Songhai; Nachtigall, Paul E.; Breese, Marlee; Supin, Alexander Ya.

2012-01-01

Emitted biosonar clicks and auditory evoked potential (AEP) responses triggered by the clicks were synchronously recorded during echolocation in an Atlantic bottlenose dolphin (Tursiops truncatus) trained to wear suction-cup EEG electrodes and to detect targets by echolocation. Three targets with target strengths of −34, −28, and −22 dB were used at distances of 2 to 6.5 m for each target. The AEP responses were sorted according to the corresponding emitted click source levels in 5-dB bins and averaged within each bin to extract biosonar click-related AEPs from noise. The AEP amplitudes were measured peak-to-peak and plotted as a function of click source levels for each target type, distance, and target-present or target-absent condition. Hearing sensation levels of the biosonar clicks were evaluated by comparing the functions of the biosonar click-related AEP amplitude-versus-click source level to a function of external (in free field) click-related AEP amplitude-versus-click sound pressure level. The results indicated that the dolphin's hearing sensation levels to her own biosonar clicks were equal to that of external clicks with sound pressure levels 16 to 36 dB lower than the biosonar click source levels, varying with target type, distance, and condition. These data may be assumed to indicate that the bottlenose dolphin possesses effective protection mechanisms to isolate the self-produced intense biosonar beam from the animal's ears during echolocation. PMID:22238654

Hearing sensation levels of emitted biosonar clicks in an echolocating Atlantic bottlenose dolphin.

PubMed

Li, Songhai; Nachtigall, Paul E; Breese, Marlee; Supin, Alexander Ya

2012-01-01

Emitted biosonar clicks and auditory evoked potential (AEP) responses triggered by the clicks were synchronously recorded during echolocation in an Atlantic bottlenose dolphin (Tursiops truncatus) trained to wear suction-cup EEG electrodes and to detect targets by echolocation. Three targets with target strengths of -34, -28, and -22 dB were used at distances of 2 to 6.5 m for each target. The AEP responses were sorted according to the corresponding emitted click source levels in 5-dB bins and averaged within each bin to extract biosonar click-related AEPs from noise. The AEP amplitudes were measured peak-to-peak and plotted as a function of click source levels for each target type, distance, and target-present or target-absent condition. Hearing sensation levels of the biosonar clicks were evaluated by comparing the functions of the biosonar click-related AEP amplitude-versus-click source level to a function of external (in free field) click-related AEP amplitude-versus-click sound pressure level. The results indicated that the dolphin's hearing sensation levels to her own biosonar clicks were equal to that of external clicks with sound pressure levels 16 to 36 dB lower than the biosonar click source levels, varying with target type, distance, and condition. These data may be assumed to indicate that the bottlenose dolphin possesses effective protection mechanisms to isolate the self-produced intense biosonar beam from the animal's ears during echolocation.

Echolocation behavior in big brown bats is not impaired after intense broadband noise exposures.

PubMed

Hom, Kelsey N; Linnenschmidt, Meike; Simmons, James A; Simmons, Andrea Megela

2016-10-15

Echolocating bats emit trains of intense ultrasonic biosonar pulses and listen to weaker echoes returning from objects in their environment. Identification and categorization of echoes are crucial for orientation and prey capture. Bats are social animals and often fly in groups in which they are exposed to their own emissions and to those from other bats, as well as to echoes from multiple surrounding objects. Sound pressure levels in these noisy conditions can exceed 110 dB, with no obvious deleterious effects on echolocation performance. Psychophysical experiments show that big brown bats (Eptesicus fuscus) do not experience temporary threshold shifts after exposure to intense broadband ultrasonic noise, but it is not known if they make fine-scale adjustments in their pulse emissions to compensate for any effects of the noise. We investigated whether big brown bats adapt the number, temporal patterning or relative amplitude of their emitted pulses while flying through an acoustically cluttered corridor after exposure to intense broadband noise (frequency range 10-100 kHz; sound exposure level 152 dB). Under these conditions, four bats made no significant changes in navigation errors or in pulse number, timing and amplitude 20 min, 24 h or 48 h after noise exposure. These data suggest that big brown bats remain able to perform difficult echolocation tasks after exposure to ecologically realistic levels of broadband noise. © 2016. Published by The Company of Biologists Ltd.

Broadband noise exposure does not affect hearing sensitivity in big brown bats (Eptesicus fuscus).

PubMed

Simmons, Andrea Megela; Hom, Kelsey N; Warnecke, Michaela; Simmons, James A

2016-04-01

In many vertebrates, exposure to intense sounds under certain stimulus conditions can induce temporary threshold shifts that reduce hearing sensitivity. Susceptibility to these hearing losses may reflect the relatively quiet environments in which most of these species have evolved. Echolocating big brown bats (Eptesicus fuscus) live in extremely intense acoustic environments in which they navigate and forage successfully, both alone and in company with other bats. We hypothesized that bats may have evolved a mechanism to minimize noise-induced hearing losses that otherwise could impair natural echolocation behaviors. The hearing sensitivity of seven big brown bats was measured in active echolocation and passive hearing tasks, before and after exposure to broadband noise spanning their audiometric range (10-100 kHz, 116 dB SPL re. 20 µPa rms, 1 h duration; sound exposure level 152 dB). Detection thresholds measured 20 min, 2 h or 24 h after exposure did not vary significantly from pre-exposure thresholds or from thresholds in control (sham exposure) conditions. These results suggest that big brown bats may be less susceptible to temporary threshold shifts than are other terrestrial mammals after exposure to similarly intense broadband sounds. These experiments provide fertile ground for future research on possible mechanisms employed by echolocating bats to minimize hearing losses while orienting effectively in noisy biological soundscapes. © 2016. Published by The Company of Biologists Ltd.

Echolocation in the Risso's dolphin, Grampus griseus.

PubMed

Philips, Jennifer D; Nachtigall, Paul E; Au, Whitlow W L; Pawloski, Jeffrey L; Roitblat, Herbert L

2003-01-01

The Risso's dolphin (Grampus griseus) is an exclusively cephalopod-consuming delphinid with a distinctive vertical indentation along its forehead. To investigate whether or not the species echolocates, a female Risso's dolphin was trained to discriminate an aluminum cylinder from a nylon sphere (experiment 1) or an aluminum sphere (experiment 2) while wearing eyecups and free swimming in an open-water pen in Kaneohe Bay, Hawaii. The dolphin completed the task with little difficulty despite being blindfolded. Clicks emitted by the dolphin were acquired at average amplitudes of 192.6 dB re 1 microPa, with estimated sources levels up to 216 dB re 1 microPa-1 m. Clicks were acquired with peak frequencies as high as 104.7 kHz (Mf(p) = 47.9 kHz), center frequencies as high as 85.7 kHz (Mf(0) = 56.5 kHz), 3-dB bandwidths up to 94.1 kHz (M(BW) = 39.7 kHz), and root-mean-square bandwidths up to 32.8 kHz (M(RMS) = 23.3 kHz). Click durations were between 40 and 70 micros. The data establish that the Risso's dolphin echolocates, and that, aside from slightly lower amplitudes and frequencies, the clicks emitted by the dolphin were similar to those emitted by other echolocating odontocetes. The particular acoustic and behavioral findings in the study are discussed with respect to the possible direction of the sonar transmission beam of the species.

Characteristics of hearing and echolocation in under-studied odontocete species

NASA Astrophysics Data System (ADS)

Smith, Adam B.

All odontoctes (toothed whales and dolphins) studied to date have been shown to echolocate. They use sound as their primary means for foraging, navigation, and communication with conspecifics and are thus considered acoustic specialists. However, the vast majority of what is known about odontocete acoustic systems comes from only a handful of the 76 recognized extant species. The research presented in this dissertation investigated basic characteristics of odontocete hearing and echolocation, including auditory temporal resolution, auditory pathways, directional hearing, and transmission beam characteristics, in individuals of five different odontocete species that are understudied. Modulation rate transfer functions were measured from formerly stranded individuals of four different species (Stenella longirostris, Feresa attenuata, Globicephala melas, Mesoplodon densirostris) using non-invasive auditory evoked potential methods. All individuals showed acute auditory temporal resolution that was comparable to other studied odontocete species. Using the same electrophysiological methods, auditory pathways and directional hearing were investigated in a Risso's dolphin (Grampus griseus) using both localized and far-field acoustic stimuli. The dolphin's hearing showed significant, frequency dependent asymmetry to localized sound presented on the right and left sides of its head. The dolphin also showed acute, but mostly symmetrical, directional auditory sensitivity to sounds presented in the far-field. Furthermore, characteristics of the echolocation transmission beam of this same individual Risso's dolphin were measured using a 16 element hydrophone array. The dolphin exhibited both single and dual lobed beam shapes that were more directional than similar measurements from a bottlenose dolphin, harbor porpoise, and false killer whale.

Sensory trait variation in an echolocating bat suggests roles for both selection and plasticity

PubMed Central

2014-01-01

Background Across heterogeneous environments selection and gene flow interact to influence the rate and extent of adaptive trait evolution. This complex relationship is further influenced by the rarely considered role of phenotypic plasticity in the evolution of adaptive population variation. Plasticity can be adaptive if it promotes colonization and survival in novel environments and in doing so may increase the potential for future population differentiation via selection. Gene flow between selectively divergent environments may favour the evolution of phenotypic plasticity or conversely, plasticity itself may promote gene flow, leading to a pattern of trait differentiation in the presence of gene flow. Variation in sensory traits is particularly informative in testing the role of environment in trait and population differentiation. Here we test the hypothesis of ‘adaptive differentiation with minimal gene flow’ in resting echolocation frequencies (RF) of Cape horseshoe bats (Rhinolophus capensis) across a gradient of increasingly cluttered habitats. Results Our analysis reveals a geographically structured pattern of increasing RF from open to highly cluttered habitats in R. capensis; however genetic drift appears to be a minor player in the processes influencing this pattern. Although Bayesian analysis of population structure uncovered a number of spatially defined mitochondrial groups and coalescent methods revealed regional-scale gene flow, phylogenetic analysis of mitochondrial sequences did not correlate with RF differentiation. Instead, habitat discontinuities between biomes, and not genetic and geographic distances, best explained echolocation variation in this species. We argue that both selection for increased detection distance in relatively less cluttered habitats and adaptive phenotypic plasticity may have influenced the evolution of matched echolocation frequencies and habitats across different populations. Conclusions Our study reveals significant sensory trait differentiation in the presence of historical gene flow and suggests roles for both selection and plasticity in the evolution of echolocation variation in R. capensis. These results highlight the importance of population level analyses to i) illuminate the subtle interplay between selection, plasticity and gene flow in the evolution of adaptive traits and ii) demonstrate that evolutionary processes may act simultaneously and that their relative influence may vary across different environments. PMID:24674227

By the Light of the Moon: North Pacific Dolphins Optimize Foraging with the Lunar Cycle

NASA Astrophysics Data System (ADS)

Simonis, Anne Elizabeth

The influence of the lunar cycle on dolphin foraging behavior was investigated in the productive, southern California Current Ecosystem and the oligotrophic Hawaiian Archipelago. Passive acoustic recordings from 2009 to 2015 were analyzed to document the presence of echolocation from four dolphin species that demonstrate distinct foraging preferences and diving abilities. Visual observations of dolphins, cloud coverage, commercial landings of market squid (Doryteuthis opalescens) and acoustic backscatter of fish were also considered in the Southern California Bight. The temporal variability of echolocation is described from daily to annual timescales, with emphasis on the lunar cycle as an established behavioral driver for potential dolphin prey. For dolphins that foraged at night, the presence of echolocation was reduced during nights of the full moon and during times of night that the moon was present in the night sky. In the Southern California Bight, echolocation activity was reduced for both shallow- diving common dolphins (Delphinus delphis) and deeper-diving Risso's dolphins (Grampus griseus) during times of increased illumination. Seasonal differences in acoustic behavior for both species suggest a geographic shift in dolphin populations, shoaling scattering layers or prey switching behavior during warm months, whereby dolphins target prey that do not vertically migrate. In the Hawaiian Archipelago, deep-diving short-finned pilot whales (Globicephala macrorhynchus) and shallow-diving false killer whales (Pseudorca crassidens) also showed reduced echolocation behavior during periods of increased lunar illumination. In contrast to nocturnal foraging in the northwestern Hawaiian Islands, false killer whales in the main Hawaiian Islands mainly foraged during the day and the lunar cycle showed little influence on their nocturnal acoustic behavior. Different temporal patterns in false killer whale acoustic behavior between the main and northwestern Hawaiian Islands can likely be attributed to the presence of distinct populations or social clusters with dissimilar foraging strategies. Consistent observations of reduced acoustic activity during times of increased lunar illumination show that the lunar cycle is an important predictor for nocturnal dolphin foraging behavior. The result of this research advances the scientific understanding of how dolphins optimize their foraging behavior in response to the changing distribution and abundance of their prey.

Sensory trait variation in an echolocating bat suggests roles for both selection and plasticity.

PubMed

Odendaal, Lizelle J; Jacobs, David S; Bishop, Jacqueline M

2014-03-27

Across heterogeneous environments selection and gene flow interact to influence the rate and extent of adaptive trait evolution. This complex relationship is further influenced by the rarely considered role of phenotypic plasticity in the evolution of adaptive population variation. Plasticity can be adaptive if it promotes colonization and survival in novel environments and in doing so may increase the potential for future population differentiation via selection. Gene flow between selectively divergent environments may favour the evolution of phenotypic plasticity or conversely, plasticity itself may promote gene flow, leading to a pattern of trait differentiation in the presence of gene flow. Variation in sensory traits is particularly informative in testing the role of environment in trait and population differentiation. Here we test the hypothesis of 'adaptive differentiation with minimal gene flow' in resting echolocation frequencies (RF) of Cape horseshoe bats (Rhinolophus capensis) across a gradient of increasingly cluttered habitats. Our analysis reveals a geographically structured pattern of increasing RF from open to highly cluttered habitats in R. capensis; however genetic drift appears to be a minor player in the processes influencing this pattern. Although Bayesian analysis of population structure uncovered a number of spatially defined mitochondrial groups and coalescent methods revealed regional-scale gene flow, phylogenetic analysis of mitochondrial sequences did not correlate with RF differentiation. Instead, habitat discontinuities between biomes, and not genetic and geographic distances, best explained echolocation variation in this species. We argue that both selection for increased detection distance in relatively less cluttered habitats and adaptive phenotypic plasticity may have influenced the evolution of matched echolocation frequencies and habitats across different populations. Our study reveals significant sensory trait differentiation in the presence of historical gene flow and suggests roles for both selection and plasticity in the evolution of echolocation variation in R. capensis. These results highlight the importance of population level analyses to i) illuminate the subtle interplay between selection, plasticity and gene flow in the evolution of adaptive traits and ii) demonstrate that evolutionary processes may act simultaneously and that their relative influence may vary across different environments.

Adaptive echolocation behavior in bats for the analysis of auditory scenes

PubMed Central

Chiu, Chen; Xian, Wei; Moss, Cynthia F.

2009-01-01

Summary Echolocating bats emit sonar pulses and listen to returning echoes to probe their surroundings. Bats adapt their echolocation call design to cope with dynamic changes in the acoustic environment, including habitat change or the presence of nearby conspecifics/heterospecifics. Seven pairs of big brown bats, Eptesicus fuscus, were tested in this study to examine how they adjusted their echolocation calls when flying and competing with a conspecific for food. Results showed that differences in five call parameters, start/end frequencies, duration, bandwidth and sweep rate, significantly increased in the two-bat condition compared with the baseline data. In addition, the magnitude of spectral separation of calls was negatively correlated with the baseline call design differences in individual bats. Bats with small baseline call frequency differences showed larger increases in call frequency separation when paired than those with large baseline call frequency differences, suggesting that bats actively change their sonar call structure if pre-existing differences in call design are small. Call design adjustments were also influenced by physical spacing between two bats. Calls of paired bats exhibited the largest design separations when inter-bat distance was shorter than 0.5 m, and the separation decreased as the spacing increased. All individuals modified at least one baseline call parameter in response to the presence of another conspecific. We propose that dissimilarity between the time–frequency features of sonar calls produced by different bats aids each individual in segregating echoes of its own sonar vocalizations from the acoustic signals of neighboring bats. PMID:19376960

Automated classification of dolphin echolocation click types from the Gulf of Mexico.

PubMed

Frasier, Kaitlin E; Roch, Marie A; Soldevilla, Melissa S; Wiggins, Sean M; Garrison, Lance P; Hildebrand, John A

2017-12-01

Delphinids produce large numbers of short duration, broadband echolocation clicks which may be useful for species classification in passive acoustic monitoring efforts. A challenge in echolocation click classification is to overcome the many sources of variability to recognize underlying patterns across many detections. An automated unsupervised network-based classification method was developed to simulate the approach a human analyst uses when categorizing click types: Clusters of similar clicks were identified by incorporating multiple click characteristics (spectral shape and inter-click interval distributions) to distinguish within-type from between-type variation, and identify distinct, persistent click types. Once click types were established, an algorithm for classifying novel detections using existing clusters was tested. The automated classification method was applied to a dataset of 52 million clicks detected across five monitoring sites over two years in the Gulf of Mexico (GOM). Seven distinct click types were identified, one of which is known to be associated with an acoustically identifiable delphinid (Risso's dolphin) and six of which are not yet identified. All types occurred at multiple monitoring locations, but the relative occurrence of types varied, particularly between continental shelf and slope locations. Automatically-identified click types from autonomous seafloor recorders without verifiable species identification were compared with clicks detected on sea-surface towed hydrophone arrays in the presence of visually identified delphinid species. These comparisons suggest potential species identities for the animals producing some echolocation click types. The network-based classification method presented here is effective for rapid, unsupervised delphinid click classification across large datasets in which the click types may not be known a priori.

Automated classification of dolphin echolocation click types from the Gulf of Mexico

PubMed Central

Roch, Marie A.; Soldevilla, Melissa S.; Wiggins, Sean M.; Garrison, Lance P.; Hildebrand, John A.

2017-01-01

Delphinids produce large numbers of short duration, broadband echolocation clicks which may be useful for species classification in passive acoustic monitoring efforts. A challenge in echolocation click classification is to overcome the many sources of variability to recognize underlying patterns across many detections. An automated unsupervised network-based classification method was developed to simulate the approach a human analyst uses when categorizing click types: Clusters of similar clicks were identified by incorporating multiple click characteristics (spectral shape and inter-click interval distributions) to distinguish within-type from between-type variation, and identify distinct, persistent click types. Once click types were established, an algorithm for classifying novel detections using existing clusters was tested. The automated classification method was applied to a dataset of 52 million clicks detected across five monitoring sites over two years in the Gulf of Mexico (GOM). Seven distinct click types were identified, one of which is known to be associated with an acoustically identifiable delphinid (Risso’s dolphin) and six of which are not yet identified. All types occurred at multiple monitoring locations, but the relative occurrence of types varied, particularly between continental shelf and slope locations. Automatically-identified click types from autonomous seafloor recorders without verifiable species identification were compared with clicks detected on sea-surface towed hydrophone arrays in the presence of visually identified delphinid species. These comparisons suggest potential species identities for the animals producing some echolocation click types. The network-based classification method presented here is effective for rapid, unsupervised delphinid click classification across large datasets in which the click types may not be known a priori. PMID:29216184

Clicking in shallow rivers: short-range echolocation of Irrawaddy and Ganges River dolphins in a shallow, acoustically complex habitat.

PubMed

Jensen, Frants H; Rocco, Alice; Mansur, Rubaiyat M; Smith, Brian D; Janik, Vincent M; Madsen, Peter T

2013-01-01

Toothed whales (Cetacea, odontoceti) use biosonar to navigate their environment and to find and catch prey. All studied toothed whale species have evolved highly directional, high-amplitude ultrasonic clicks suited for long-range echolocation of prey in open water. Little is known about the biosonar signals of toothed whale species inhabiting freshwater habitats such as endangered river dolphins. To address the evolutionary pressures shaping the echolocation signal parameters of non-marine toothed whales, we investigated the biosonar source parameters of Ganges river dolphins (Platanista gangetica gangetica) and Irrawaddy dolphins (Orcaella brevirostris) within the river systems of the Sundarban mangrove forest. Both Ganges and Irrawaddy dolphins produced echolocation clicks with a high repetition rate and low source level compared to marine species. Irrawaddy dolphins, inhabiting coastal and riverine habitats, produced a mean source level of 195 dB (max 203 dB) re 1 µPapp whereas Ganges river dolphins, living exclusively upriver, produced a mean source level of 184 dB (max 191) re 1 µPapp. These source levels are 1-2 orders of magnitude lower than those of similar sized marine delphinids and may reflect an adaptation to a shallow, acoustically complex freshwater habitat with high reverberation and acoustic clutter. The centroid frequency of Ganges river dolphin clicks are an octave lower than predicted from scaling, but with an estimated beamwidth comparable to that of porpoises. The unique bony maxillary crests found in the Platanista forehead may help achieve a higher directionality than expected using clicks nearly an octave lower than similar sized odontocetes.

Clicking in Shallow Rivers: Short-Range Echolocation of Irrawaddy and Ganges River Dolphins in a Shallow, Acoustically Complex Habitat

PubMed Central

Jensen, Frants H.; Rocco, Alice; Mansur, Rubaiyat M.; Smith, Brian D.; Janik, Vincent M.; Madsen, Peter T.

2013-01-01

Toothed whales (Cetacea, odontoceti) use biosonar to navigate their environment and to find and catch prey. All studied toothed whale species have evolved highly directional, high-amplitude ultrasonic clicks suited for long-range echolocation of prey in open water. Little is known about the biosonar signals of toothed whale species inhabiting freshwater habitats such as endangered river dolphins. To address the evolutionary pressures shaping the echolocation signal parameters of non-marine toothed whales, we investigated the biosonar source parameters of Ganges river dolphins (Platanista gangetica gangetica) and Irrawaddy dolphins (Orcaella brevirostris) within the river systems of the Sundarban mangrove forest. Both Ganges and Irrawaddy dolphins produced echolocation clicks with a high repetition rate and low source level compared to marine species. Irrawaddy dolphins, inhabiting coastal and riverine habitats, produced a mean source level of 195 dB (max 203 dB) re 1 µPapp whereas Ganges river dolphins, living exclusively upriver, produced a mean source level of 184 dB (max 191) re 1 µPapp. These source levels are 1–2 orders of magnitude lower than those of similar sized marine delphinids and may reflect an adaptation to a shallow, acoustically complex freshwater habitat with high reverberation and acoustic clutter. The centroid frequency of Ganges river dolphin clicks are an octave lower than predicted from scaling, but with an estimated beamwidth comparable to that of porpoises. The unique bony maxillary crests found in the Platanista forehead may help achieve a higher directionality than expected using clicks nearly an octave lower than similar sized odontocetes. PMID:23573197

Ambient noise causes independent changes in distinct spectro-temporal features of echolocation calls in horseshoe bats.

PubMed

Hage, Steffen R; Jiang, Tinglei; Berquist, Sean W; Feng, Jiang; Metzner, Walter

2014-07-15

One of the most efficient mechanisms to optimize signal-to-noise ratios is the Lombard effect - an involuntary rise in call amplitude due to ambient noise. It is often accompanied by changes in the spectro-temporal composition of calls. We examined the effects of broadband-filtered noise on the spectro-temporal composition of horseshoe bat echolocation calls, which consist of a constant-frequency component and initial and terminal frequency-modulated components. We found that the frequency-modulated components became larger for almost all noise conditions, whereas the bandwidth of the constant-frequency component increased only when broadband-filtered noise was centered on or above the calls' dominant or fundamental frequency. This indicates that ambient noise independently modifies the associated acoustic parameters of the Lombard effect, such as spectro-temporal features, and could significantly affect the bat's ability to detect and locate targets. Our findings may be of significance in evaluating the impact of environmental noise on echolocation behavior in bats. © 2014. Published by The Company of Biologists Ltd.

Dynamic adjustment of echolocation pulse structure of big-footed myotis (Myotis macrodactylus) in response to different habitats.

PubMed

Wang, Lei; Luo, Jinhong; Wang, Hongna; Ou, Wei; Jiang, Tinglei; Liu, Ying; Lyle, Dennis; Feng, Jiang

2014-02-01

Studying relationships between characteristics of sonar pulses and habitat clutter level is important for the understanding of signal design in bat echolocation. However, most studies have focused on overall spectral and temporal parameters of such vocalizations, with focus less on potential variation in frequency modulation rates (MRs) occurring within each pulse. In the current study, frequency modulation (FM) characteristics were examined in echolocation pulses recorded from big-footed myotis (Myotis macrodactylus) bats as these animals searched for prey in five habitats differing in relative clutter level. Pulses were analyzed using ten parameters, including four structure-related characters which were derived by dividing each pulse into three elements based on two knees in the FM sweep. Results showed that overall frequency, pulse duration, and MR all varied across habitat. The strongest effects were found for MR in the body of the pulse, implying that this particular component plays a major role as M. macrodactylus, and potentially other bat species, adjust to varying clutter levels in their foraging habitats.

Tiger moths and the threat of bats: decision-making based on the activity of a single sensory neuron.

PubMed

Ratcliffe, John M; Fullard, James H; Arthur, Benjamin J; Hoy, Ronald R

2009-06-23

Echolocating bats and eared moths are a model system of predator-prey interaction within an almost exclusively auditory world. Through selective pressures from aerial-hawking bats, noctuoid moths have evolved simple ears that contain one to two auditory neurons and function to detect bat echolocation calls and initiate defensive flight behaviours. Among these moths, some chemically defended and mimetic tiger moths also produce ultrasonic clicks in response to bat echolocation calls; these defensive signals are effective warning signals and may interfere with bats' ability to process echoic information. Here, we demonstrate that the activity of a single auditory neuron (the A1 cell) provides sufficient information for the toxic dogbane tiger moth, Cycnia tenera, to decide when to initiate defensive sound production in the face of bats. Thus, despite previous suggestions to the contrary, these moths' only other auditory neuron, the less sensitive A2 cell, is not necessary for initiating sound production. However, we found a positive linear relationship between combined A1 and A2 activity and the number of clicks the dogbane tiger moth produces.

Assessing bat detectability and occupancy with multiple automated echolocation detectors

USGS Publications Warehouse

Gorresen, P.M.; Miles, A.C.; Todd, C.M.; Bonaccorso, F.J.; Weller, T.J.

2008-01-01

Occupancy analysis and its ability to account for differential detection probabilities is important for studies in which detecting echolocation calls is used as a measure of bat occurrence and activity. We examined the feasibility of remotely acquiring bat encounter histories to estimate detection probability and occupancy. We used echolocation detectors coupled to digital recorders operating at a series of proximate sites on consecutive nights in 2 trial surveys for the Hawaiian hoary bat (Lasiurus cinereus semotus). Our results confirmed that the technique is readily amenable for use in occupancy analysis. We also conducted a simulation exercise to assess the effects of sampling effort on parameter estimation. The results indicated that the precision and bias of parameter estimation were often more influenced by the number of sites sampled than number of visits. Acceptable accuracy often was not attained until at least 15 sites or 15 visits were used to estimate detection probability and occupancy. The method has significant potential for use in monitoring trends in bat activity and in comparative studies of habitat use. ?? 2008 American Society of Mammalogists.

The cochlear size of bats and rodents derived from MRI images and histology.

PubMed

Hsiao, Chun Jen; Jen, Philip Hung-Sun; Wu, Chung Hsin

2015-05-27

From the evolutionary perspective, the ear of each animal species is built for effective processing of the biologically relevant signals used for communication and acoustically guided orientation. Because the sound pulses used by echolocating bats for orientation and rodents for communication are quite different, the basic design of the mammalian auditory system commonly shared by echolocating bats must be specialized in some manner to effectively process their species-specific sounds. The present study examines the difference in the cochlea of these animal species using MRI images and histological techniques. We report here that, although all these animal species share a similar cochlear structure, they vary in their cochlear size and turns. Bats using constant frequency-frequency-modulated pulses (CF-FM bats) and frequency-modulated pulses (FM bats) for echolocation have a larger cochlear size and more cochlear turns than rodents (mice and rats). However, CF-FM bats have the largest cochlear size and most cochlear turns. This difference in cochlear size and turns of these animal species is discussed in relation to their biologically relevant sounds and acoustic behavior.

Echolocation signals of wild Atlantic spotted dolphin (Stenella frontalis)

NASA Astrophysics Data System (ADS)

Au, Whitlow W. L.; Herzing, Denise L.

2003-01-01

An array of four hydrophones arranged in a symmetrical star configuration was used to measure the echolocation signals of the Atlantic spotted dolphin (Stenella frontalis) in the Bahamas. The spacing between the center hydrophone and the other hydrophones was 45.7 cm. A video camera was attached to the array and a video tape recorder was time synchronized with the computer used to digitize the acoustic signals. The echolocation signals had bi-modal frequency spectra with a low-frequency peak between 40 and 50 kHz and a high-frequency peak between 110 and 130 kHz. The low-frequency peak was dominant when the signal the source level was low and the high-frequency peak dominated when the source level was high. Peak-to-peak source levels as high as 210 dB re 1 μPa were measured. The source level varied in amplitude approximately as a function of the one-way transmission loss for signals traveling from the animals to the array. The characteristics of the signals were similar to those of captive Tursiops truncatus, Delphinapterus leucas and Pseudorca crassidens measured in open waters under controlled conditions.

Listening for bats: the hearing range of the bushcricket Phaneroptera falcata for bat echolocation calls measured in the field.

PubMed Central

Schul, J; Matt, F; von Helversen, O

2000-01-01

The hearing range of the tettigoniid Phaneropterafalcata for the echolocation calls of freely flying mouseeared bats (Myotis myotis) was determined in the field. The hearing of the insect was monitored using hook electrode recordings from an auditory interneuron, which is as sensitive as the hearing organ for frequencies above 16 kHz. The flight path of the bat relative to the insect's position was tracked by recording the echolocation calls with two microphone arrays, and calculating the bat's position from the arrival time differences of the calls at each microphone. The hearing distances ranged from 13 to 30 m. The large variability appeared both between different insects and between different bat approaches to an individual insect. The escape time of the bushcricket, calculated from the detection distance of the insect and the instantaneous flight speed of the bat, ranged from 1.5 to more than 4s. The hearing ranges of bushcrickets suggest that the insect hears the approaching bat long before the bat can detect an echo from the flying insect. PMID:12233766

Echolocation of insects using intermittent frequency-modulated sounds.

PubMed

Matsuo, Ikuo; Takanashi, Takuma

2015-09-01

Using echolocation influenced by Doppler shift, bats can capture flying insects in real three-dimensional space. On the basis of this principle, a model that estimates object locations using frequency modulated (FM) sound was proposed. However, no investigation was conducted to verify whether the model can localize flying insects from their echoes. This study applied the model to estimate the range and direction of flying insects by extracting temporal changes from the time-frequency pattern and interaural range difference, respectively. The results obtained confirm that a living insect's position can be estimated using this model with echoes measured while emitting intermittent FM sounds.

Light-emitting diode street lights reduce last-ditch evasive manoeuvres by moths to bat echolocation calls

PubMed Central

Wakefield, Andrew; Stone, Emma L.; Jones, Gareth; Harris, Stephen

2015-01-01

The light-emitting diode (LED) street light market is expanding globally, and it is important to understand how LED lights affect wildlife populations. We compared evasive flight responses of moths to bat echolocation calls experimentally under LED-lit and -unlit conditions. Significantly, fewer moths performed ‘powerdive’ flight manoeuvres in response to bat calls (feeding buzz sequences from Nyctalus spp.) under an LED street light than in the dark. LED street lights reduce the anti-predator behaviour of moths, shifting the balance in favour of their predators, aerial hawking bats. PMID:26361558

From the ultrasonic to the infrared: molecular evolution and the sensory biology of bats

PubMed Central

Jones, Gareth; Teeling, Emma C.; Rossiter, Stephen J.

2013-01-01

Great advances have been made recently in understanding the genetic basis of the sensory biology of bats. Research has focused on the molecular evolution of candidate sensory genes, genes with known functions [e.g., olfactory receptor (OR) genes] and genes identified from mutations associated with sensory deficits (e.g., blindness and deafness). For example, the FoxP2 gene, underpinning vocal behavior and sensorimotor coordination, has undergone diversification in bats, while several genes associated with audition show parallel amino acid substitutions in unrelated lineages of echolocating bats and, in some cases, in echolocating dolphins, representing a classic case of convergent molecular evolution. Vision genes encoding the photopigments rhodopsin and the long-wave sensitive opsin are functional in bats, while that encoding the short-wave sensitive opsin has lost functionality in rhinolophoid bats using high-duty cycle laryngeal echolocation, suggesting a sensory trade-off between investment in vision and echolocation. In terms of olfaction, bats appear to have a distinctive OR repertoire compared with other mammals, and a gene involved in signal transduction in the vomeronasal system has become non-functional in most bat species. Bitter taste receptors appear to have undergone a “birth-and death” evolution involving extensive gene duplication and loss, unlike genes coding for sweet and umami tastes that show conservation across most lineages but loss in vampire bats. Common vampire bats have also undergone adaptations for thermoperception, via alternative splicing resulting in the evolution of a novel heat-sensitive channel. The future for understanding the molecular basis of sensory biology is promising, with great potential for comparative genomic analyses, studies on gene regulation and expression, exploration of the role of alternative splicing in the generation of proteomic diversity, and linking genetic mechanisms to behavioral consequences. PMID:23755015

Bat detective-Deep learning tools for bat acoustic signal detection.

PubMed

Mac Aodha, Oisin; Gibb, Rory; Barlow, Kate E; Browning, Ella; Firman, Michael; Freeman, Robin; Harder, Briana; Kinsey, Libby; Mead, Gary R; Newson, Stuart E; Pandourski, Ivan; Parsons, Stuart; Russ, Jon; Szodoray-Paradi, Abigel; Szodoray-Paradi, Farkas; Tilova, Elena; Girolami, Mark; Brostow, Gabriel; Jones, Kate E

2018-03-01

Passive acoustic sensing has emerged as a powerful tool for quantifying anthropogenic impacts on biodiversity, especially for echolocating bat species. To better assess bat population trends there is a critical need for accurate, reliable, and open source tools that allow the detection and classification of bat calls in large collections of audio recordings. The majority of existing tools are commercial or have focused on the species classification task, neglecting the important problem of first localizing echolocation calls in audio which is particularly problematic in noisy recordings. We developed a convolutional neural network based open-source pipeline for detecting ultrasonic, full-spectrum, search-phase calls produced by echolocating bats. Our deep learning algorithms were trained on full-spectrum ultrasonic audio collected along road-transects across Europe and labelled by citizen scientists from www.batdetective.org. When compared to other existing algorithms and commercial systems, we show significantly higher detection performance of search-phase echolocation calls with our test sets. As an example application, we ran our detection pipeline on bat monitoring data collected over five years from Jersey (UK), and compared results to a widely-used commercial system. Our detection pipeline can be used for the automatic detection and monitoring of bat populations, and further facilitates their use as indicator species on a large scale. Our proposed pipeline makes only a small number of bat specific design decisions, and with appropriate training data it could be applied to detecting other species in audio. A crucial novelty of our work is showing that with careful, non-trivial, design and implementation considerations, state-of-the-art deep learning methods can be used for accurate and efficient monitoring in audio.

Bat detective—Deep learning tools for bat acoustic signal detection

PubMed Central

Barlow, Kate E.; Firman, Michael; Freeman, Robin; Harder, Briana; Kinsey, Libby; Mead, Gary R.; Newson, Stuart E.; Pandourski, Ivan; Russ, Jon; Szodoray-Paradi, Abigel; Tilova, Elena; Girolami, Mark; Jones, Kate E.

2018-01-01

Passive acoustic sensing has emerged as a powerful tool for quantifying anthropogenic impacts on biodiversity, especially for echolocating bat species. To better assess bat population trends there is a critical need for accurate, reliable, and open source tools that allow the detection and classification of bat calls in large collections of audio recordings. The majority of existing tools are commercial or have focused on the species classification task, neglecting the important problem of first localizing echolocation calls in audio which is particularly problematic in noisy recordings. We developed a convolutional neural network based open-source pipeline for detecting ultrasonic, full-spectrum, search-phase calls produced by echolocating bats. Our deep learning algorithms were trained on full-spectrum ultrasonic audio collected along road-transects across Europe and labelled by citizen scientists from www.batdetective.org. When compared to other existing algorithms and commercial systems, we show significantly higher detection performance of search-phase echolocation calls with our test sets. As an example application, we ran our detection pipeline on bat monitoring data collected over five years from Jersey (UK), and compared results to a widely-used commercial system. Our detection pipeline can be used for the automatic detection and monitoring of bat populations, and further facilitates their use as indicator species on a large scale. Our proposed pipeline makes only a small number of bat specific design decisions, and with appropriate training data it could be applied to detecting other species in audio. A crucial novelty of our work is showing that with careful, non-trivial, design and implementation considerations, state-of-the-art deep learning methods can be used for accurate and efficient monitoring in audio. PMID:29518076

Discrimination of fast click-series produced by tagged Risso's dolphins (Grampus griseus) for echolocation or communication.

PubMed

Arranz, P; DeRuiter, S L; Stimpert, A K; Neves, S; Friedlaender, A S; Goldbogen, J A; Visser, F; Calambokidis, J; Southall, B L; Tyack, P L

2016-09-15

Early studies that categorized odontocete pulsed sounds had few means of discriminating signals used for biosonar-based foraging from those used for communication. This capability to identify the function of sounds is important for understanding and interpreting behavior; it is also essential for monitoring and mitigating potential disturbance from human activities. Archival tags were placed on free-ranging Grampus griseus to quantify and discriminate between pulsed sounds used for echolocation-based foraging and those used for communication. Two types of rapid click-series pulsed sounds, buzzes and burst pulses, were identified as produced by the tagged dolphins and classified using a Gaussian mixture model based on their duration, association with jerk (i.e. rapid change of acceleration) and temporal association with click trains. Buzzes followed regular echolocation clicks and coincided with a strong jerk signal from accelerometers on the tag. They consisted of series averaging 359±210 clicks (mean±s.d.) with an increasing repetition rate and relatively low amplitude. Burst pulses consisted of relatively short click series averaging 45±54 clicks with decreasing repetition rate and longer inter-click interval that were less likely to be associated with regular echolocation and the jerk signal. These results suggest that the longer, relatively lower amplitude, jerk-associated buzzes are used in this species to capture prey, mostly during the bottom phase of foraging dives, as seen in other odontocetes. In contrast, the shorter, isolated burst pulses that are generally emitted by the dolphins while at or near the surface are used outside of a direct, known foraging context. © 2016. Published by The Company of Biologists Ltd.

Biomimetic Signal Processing Using the Biosonar Measurement Tool (BMT)

NASA Astrophysics Data System (ADS)

Abawi, Ahmad T.; Hursky, Paul; Porter, Michael B.; Tiemann, Chris; Martin, Stephen

2004-11-01

In this paper data recorded on the Biosonar Measurement Tool (BMT) during a target echolocation experiment are used to 1) find ways to separate target echoes from clutter echoes, 2) analyze target returns and 3) find features in target returns that distinguish them from clutter returns. The BMT is an instrumentation package used in dolphin echolocation experiments developed at SPAWARSYSCEN. It can be held by the dolphin using a bite-plate during echolocation experiments and records the movement and echolocation strategy of a target-hunting dolphin without interfering with its motion through the search field. The BMT was developed to record a variety of data from a free-swimming dolphin engaged in a bottom target detection task. These data include the three dimensional location of the dolphin, including its heading, pitch roll and velocity as well as passive acoustic data recorded on three channels. The outgoing dolphin click is recorded on one channel and the resulting echoes are recorded on the two remaining channels. For each outgoing click the BMT records a large number of echoes that come from the entire ensonified field. Given the large number of transmitted clicks and the returned echoes, it is almost impossible to find a target return from the recorded data on the BMT. As a means of separating target echoes from those of clutter, an echo-mapping tool was developed. This tool produces an echomap on which echoes from targets (and other regular objects such as surface buoys, the side of a boat and so on) stack together as tracks, while echoes from clutter are scattered. Once these tracks are identified, the retuned echoes can easily be extracted for further analysis.

Drinking and Flying: Does Alcohol Consumption Affect the Flight and Echolocation Performance of Phyllostomid Bats?

PubMed Central

Orbach, Dara N.; Veselka, Nina; Dzal, Yvonne; Lazure, Louis; Fenton, M. Brock

2010-01-01

Background In the wild, frugivorous and nectarivorous bats often eat fermenting fruits and nectar, and thus may consume levels of ethanol that could induce inebriation. To understand if consumption of ethanol by bats alters their access to food and general survival requires examination of behavioural responses to its ingestion, as well as assessment of interspecific variation in those responses. We predicted that bats fed ethanol would show impaired flight and echolocation behaviour compared to bats fed control sugar water, and that there would be behavioural differences among species. Methodology/Principal Findings We fed wild caught Artibeus jamaicensis, A. lituratus, A. phaeotis, Carollia sowelli, Glossophaga soricina, and Sturnira lilium (Chiroptera, Phyllostomidae) sugar water (44 g of table sugar in 500 ml of water) or sugar water with ethanol before challenging them to fly through an obstacle course while we simultaneously recorded their echolocation calls. We used bat saliva, a non-invasive proxy, to measure blood ethanol concentrations ranging from 0 to >0.3% immediately before flight trials. Flight performance and echolocation behaviour were not significantly affected by consumption of ethanol, but species differed in their blood alcohol concentrations after consuming it. Conclusions/Significance The bats we studied display a tolerance for ethanol that could have ramifications for the adaptive radiation of frugivorous and nectarivorous bats by allowing them to use ephemeral food resources over a wide span of time. By sampling across phyllostomid genera, we show that patterns of apparent ethanol tolerance in New World bats are broad, and thus may have been an important early step in the evolution of frugivory and nectarivory in these animals. PMID:20126552

Congruent representation of visual and acoustic space in the superior colliculus of the echolocating bat Phyllostomus discolor.

PubMed

Hoffmann, Susanne; Vega-Zuniga, Tomas; Greiter, Wolfgang; Krabichler, Quirin; Bley, Alexandra; Matthes, Mariana; Zimmer, Christiane; Firzlaff, Uwe; Luksch, Harald

2016-11-01

The midbrain superior colliculus (SC) commonly features a retinotopic representation of visual space in its superficial layers, which is congruent with maps formed by multisensory neurons and motor neurons in its deep layers. Information flow between layers is suggested to enable the SC to mediate goal-directed orienting movements. While most mammals strongly rely on vision for orienting, some species such as echolocating bats have developed alternative strategies, which raises the question how sensory maps are organized in these animals. We probed the visual system of the echolocating bat Phyllostomus discolor and found that binocular high acuity vision is frontally oriented and thus aligned with the biosonar system, whereas monocular visual fields cover a large area of peripheral space. For the first time in echolocating bats, we could show that in contrast with other mammals, visual processing is restricted to the superficial layers of the SC. The topographic representation of visual space, however, followed the general mammalian pattern. In addition, we found a clear topographic representation of sound azimuth in the deeper collicular layers, which was congruent with the superficial visual space map and with a previously documented map of orienting movements. Especially for bats navigating at high speed in densely structured environments, it is vitally important to transfer and coordinate spatial information between sensors and motor systems. Here, we demonstrate first evidence for the existence of congruent maps of sensory space in the bat SC that might serve to generate a unified representation of the environment to guide motor actions. © 2016 Federation of European Neuroscience Societies and John Wiley & Sons Ltd.

Operating range of a differential-absorption lidar based on a CO{sub 2} laser

DOE Office of Scientific and Technical Information (OSTI.GOV)

Ivashchenko, M V; Sherstov, I V

2000-08-31

The echolocation range and the remote sensing of ethylene in the atmosphere are simulated for a differential-absorption lidar based on TEA CO{sub 2} lasers. The dependence of the lidar echolocation range on the energy and the peak power of probe pulses is shown to be close to logarithmic. It is demonstrated that the use of narrow-band spectral filters is justified only for low-noise detectors and viewing angles of the receiver exceeding 5 mrad. The relative measurement error of the ethylene concentration in the atmosphere is estimated for various detection modes. (laser applications and other topics in quantum electronics)

Some results of studying the acoustics of dolphins

NASA Astrophysics Data System (ADS)

Romanenko, E. V.

2004-05-01

An experimental study of the echolocation ability of dolphins ( Tursiops truncatus) is performed in the presence of correlated and uncorrelated broadband noise acting on their organs of hearing. It is shown that, under such conditions, the echolocating pulses of a dolphin become noticeably modified: in the absence of noise, standard broadband pulses are produced, while in the presence of noise, the pulses acquire an oscillatory character (become narrowband). Sounds and air pressure that occur inside the respiratory tract of a dolphin when the animal produces whistles and pulsed signals are studied. Data testifying in favor of the pneumatic origin of sounds generated by dolphins are obtained.

High-frequency modulated signals of killer whales (Orcinus orca) in the North Pacific.

PubMed

Simonis, Anne E; Baumann-Pickering, Simone; Oleson, Erin; Melcón, Mariana L; Gassmann, Martin; Wiggins, Sean M; Hildebrand, John A

2012-04-01

Killer whales in the North Pacific, similar to Atlantic populations, produce high-frequency modulated signals, based on acoustic recordings from ship-based hydrophone arrays and autonomous recorders at multiple locations. The median peak frequency of these signals ranged from 19.6-36.1 kHz and median duration ranged from 50-163 ms. Source levels were 185-193 dB peak-to-peak re: 1 μPa at 1 m. These uniform, repetitive, down-swept signals are similar to bat echolocation signals and possibly could have echolocation functionality. A large geographic range of occurrence suggests that different killer whale ecotypes may utilize these signals.

Echolocation clicks from killer whales (Orcinus orca) feeding on herring (Clupea harengus).

PubMed

Simon, Malene; Wahlberg, Magnus; Miller, Lee A

2007-02-01

Echolocation clicks from Norwegian killer whales feeding on herring schools were recorded using a four-hydrophone array. The clicks had broadband bimodal frequency spectra with low and high frequency peaks at 24 and 108 kHz, respectively. The -10 dB bandwidth was 35 kHz. The average source level varied from 173 to 202 dB re 1 microPa (peak-to-peak) at 1 m. This is considerably lower than source levels described for Canadian killer whales foraging on salmon. It is suggested that biosonar clicks of Norwegian killer whales are adapted for localization of prey with high target strength and acute hearing abilities.

The function of male sperm whale slow clicks in a high latitude habitat: communication, echolocation, or prey debilitation?

PubMed

Oliveira, Cláudia; Wahlberg, Magnus; Johnson, Mark; Miller, Patrick J O; Madsen, Peter T

2013-05-01

Sperm whales produce different click types for echolocation and communication. Usual clicks and buzzes appear to be used primarily in foraging while codas are thought to function in social communication. The function of slow clicks is less clear, but they appear to be produced by males at higher latitudes, where they primarily forage solitarily, and on the breeding grounds, where they roam between groups of females. Here the behavioral context in which these vocalizations are produced and the function they may serve was investigated. Ninety-nine hours of acoustic and diving data were analyzed from sound recording tags on six male sperm whales in Northern Norway. The 755 slow clicks detected were produced by tagged animals at the surface (52%), ascending from a dive (37%), and during the bottom phase (11%), but never during the descent. Slow clicks were not associated with the production of buzzes, other echolocation clicks, or fast maneuvering that would indicate foraging. Some slow clicks were emitted in seemingly repetitive temporal patterns supporting the hypothesis that the function for slow clicks on the feeding grounds is long range communication between males, possibly relaying information about individual identity or behavioral states.

Species-specific beaked whale echolocation signals.

PubMed

Baumann-Pickering, Simone; McDonald, Mark A; Simonis, Anne E; Solsona Berga, Alba; Merkens, Karlina P B; Oleson, Erin M; Roch, Marie A; Wiggins, Sean M; Rankin, Shannon; Yack, Tina M; Hildebrand, John A

2013-09-01

Beaked whale echolocation signals are mostly frequency-modulated (FM) upsweep pulses and appear to be species specific. Evolutionary processes of niche separation may have driven differentiation of beaked whale signals used for spatial orientation and foraging. FM pulses of eight species of beaked whales were identified, as well as five distinct pulse types of unknown species, but presumed to be from beaked whales. Current evidence suggests these five distinct but unidentified FM pulse types are also species-specific and are each produced by a separate species. There may be a relationship between adult body length and center frequency with smaller whales producing higher frequency signals. This could be due to anatomical and physiological restraints or it could be an evolutionary adaption for detection of smaller prey for smaller whales with higher resolution using higher frequencies. The disadvantage of higher frequencies is a shorter detection range. Whales echolocating with the highest frequencies, or broadband, likely lower source level signals also use a higher repetition rate, which might compensate for the shorter detection range. Habitat modeling with acoustic detections should give further insights into how niches and prey may have shaped species-specific FM pulse types.

Invariance of evoked-potential echo-responses to target strength and distance in an echolocating false killer whale

NASA Astrophysics Data System (ADS)

Supin, Alexander Ya.; Nachtigall, Paul E.; Au, Whitlow W. L.; Breese, Marlee

2005-06-01

Brain auditory evoked potentials (AEPs) were recorded in a false killer whale Pseudorca crassidens trained to accept suction-cup EEG electrodes and to detect targets by echolocation. AEP collection was triggered by echolocation pulses transmitted by the animal. The target strength varied from -22 to -40 dB the distance varied from 1.5 to 6 m. All the records contained two AEP sets: the first one of a constant latency (transmission-related AEP) and a second one with a delay proportional to the distance (echo-related AEP). The amplitude of echo-related AEPs was almost independent of both target strength and distance, though combined variation of these two parameters resulted in echo intensity variation within a range of 42 dB. The amplitude of transmission-related AEPs was independent of distance but dependent on target strength: the less the target strength, the higher the amplitude. Recording of transmitted pulses has not shown their intensity dependence on target strength. It is supposed that the constancy of echo-related AEP results from variation of hearing sensitivity depending on the target strength and release of echo-related responses from masking by transmitted pulses depending on the distance. .

Basis of acoustic discrimination of Chinook salmon from other salmons by echolocating Orcinus orca.

PubMed

Au, Whitlow W L; Horne, John K; Jones, Christopher

2010-10-01

The "resident" ecotype of killer whales (Orcinus orca) in the waters of British Columbia and Washington State have a strong preference for Chinook salmon even in months when Chinook comprise less than about 10% of the salmon population. The foraging behavior of killer whales suggests that they depend on echolocation to detect and recognize their prey. In order to determine possible cues in echoes from salmon species, a series of backscatter measurements were made at the Applied Physics Laboratory (Univ. of Wash.) Facility on Lake Union, on three different salmon species using simulated killer whale echolocation signals. The fish were attached to a monofilament net panel and rotated while echoes were collected, digitized and stored on a laptop computer. Three transducer depths were used; same depth, 22° and 45° above the horizontal plane of the fish. Echoes were collected from five Chinook, three coho and one sockeye salmon. Radiograph images of all specimens were obtained to examine the swimbladder shape and orientation. The results show that echo structure from similar length but different species of salmon were different and probably recognizable by foraging killer whales.

Invariance of evoked-potential echo-responses to target strength and distance in an echolocating false killer whale.

PubMed

Supin, Alexander Ya; Nachtigall, Paul E; Au, Whitlow W L; Breese, Marlee

2005-06-01

Brain auditory evoked potentials (AEPs) were recorded in a false killer whale Pseudorca crassidens trained to accept suction-cup EEG electrodes and to detect targets by echolocation. AEP collection was triggered by echolocation pulses transmitted by the animal. The target strength varied from -22 to -40 dB; the distance varied from 1.5 to 6 m. All the records contained two AEP sets: the first one of a constant latency (transmission-related AEP) and a second one with a delay proportional to the distance (echo-related AEP). The amplitude of echo-related AEPs was almost independent of both target strength and distance, though combined variation of these two parameters resulted in echo intensity variation within a range of 42 dB. The amplitude of transmission-related AEPs was independent of distance but dependent on target strength: the less the target strength, the higher the amplitude. Recording of transmitted pulses has not shown their intensity dependence on target strength. It is supposed that the constancy of echo-related AEP results from variation of hearing sensitivity depending on the target strength and release of echo-related responses from masking by transmitted pulses depending on the distance.

Rapid jamming avoidance in biosonar.

PubMed

Gillam, Erin H; Ulanovsky, Nachum; McCracken, Gary F

2007-03-07

The sonar systems of bats and dolphins are in many ways superior to man-made sonar and radar systems, and considerable effort has been devoted to understanding the signal-processing strategies underlying these capabilities. A major feature determining the efficiency of sonar systems is the sensitivity to noise and jamming signals. Previous studies indicated that echolocating bats may adjust their signal structure to avoid jamming ('jamming avoidance response'; JAR). However, these studies relied on behavioural correlations and not controlled experiments. Here, we provide the first experimental evidence for JAR in bats. We presented bats (Tadarida brasiliensis) with 'playback stimuli' consisting of recorded echolocation calls at one of six frequencies. The bats exhibited a JAR by shifting their call frequency away from the presented playback frequency. When the approaching bats were challenged by an abrupt change in the playback stimulus, they responded by shifting their call frequencies upwards, away from the playback. Interestingly, even bats initially calling below the playback's frequency shifted their frequencies upwards, 'jumping' over the playback frequency. These spectral shifts in the bats' calls occurred often within less than 200 ms, in the first echolocation call emitted after the stimulus switch-suggesting that rapid jamming avoidance is important for the bat.

A neural mechanism for detecting the distance of a selected target by modulating the FM sweep rate of biosonar in echolocation of bat.

PubMed

Kamata, Eigo; Inoue, Satoru; Zheng, MeiHong; Kashimori, Yoshiki; Kambara, Takeshi

2004-01-01

Most species of bats making echolocation use frequency modulated (FM) ultrasonic pulses to measure the distance to targets. These bats detect with a high accuracy the arrival time differences between emitted pulses and their echoes generated by targets. In order to clarify the neural mechanism for echolocation, we present neural model of inferior colliculus (IC), medial geniculate body (MGB) and auditory cortex (AC) along which information of echo delay times is processed. The bats increase the downward frequency sweep rate of emitted FM pulse as they approach the target. The functional role of this modulation of sweep rate is not yet clear. In order to investigate the role, we calculated the response properties of our models of IC, MGB, and AC changing the target distance and the sweep rate. We found based on the simulations that the distance of a target in various ranges may be encoded the most clearly into the activity pattern of delay time map network in AC, when the sweep rate of FM pulse used is coincided with the observed value which the bats adopt for each range of target distance.

Classification of communication signals of the little brown bat

NASA Astrophysics Data System (ADS)

Melendez, Karla V.; Jones, Douglas L.; Feng, Albert S.

2005-09-01

Little brown bats, Myotis lucifugus, are known for their ability to echolocate and utilize their echolocation system to navigate, locate, and identify prey. Their echolocation signals have been characterized in detail, but their communication signals are poorly understood despite their widespread use during the social interactions. The goal of this study was to characterize the communication signals of little brown bats. Sound recordings were made overnight on five individual bats (housed separately from a large group of captive bats) for 7 nights, using a Pettersson ultrasound detector D240x bat detector and Nagra ARES-BB digital recorder. The spectral and temporal characteristics of recorded sounds were first analyzed using BATSOUND software from Pettersson. Sounds were first classified by visual observation of calls' temporal pattern and spectral composition, and later using an automatic classification scheme based on multivariate statistical parameters in MATLAB. Human- and machine-based analysis revealed five discrete classes of bat's communication signals: downward frequency-modulated calls, constant frequency calls, broadband noise bursts, broadband chirps, and broadband click trains. Future studies will focus on analysis of calls' spectrotemporal modulations to discriminate any subclasses that may exist. [Research supported by Grant R01-DC-04998 from the National Institute for Deafness and Communication Disorders.

Discrimination of complex synthetic echoes by an echolocating bottlenose dolphin

NASA Astrophysics Data System (ADS)

Helweg, David A.; Moore, Patrick W.; Dankiewicz, Lois A.; Zafran, Justine M.; Brill, Randall L.

2003-02-01

Bottlenose dolphins (Tursiops truncatus) detect and discriminate underwater objects by interrogating the environment with their native echolocation capabilities. Study of dolphins' ability to detect complex (multihighlight) signals in noise suggest echolocation object detection using an approximate 265-μs energy integration time window sensitive to the echo region of highest energy or containing the highlight with highest energy. Backscatter from many real objects contains multiple highlights, distributed over multiple integration windows and with varying amplitude relationships. This study used synthetic echoes with complex highlight structures to test whether high-amplitude initial highlights would interfere with discrimination of low-amplitude trailing highlights. A dolphin was trained to discriminate two-highlight synthetic echoes using differences in the center frequencies of the second highlights. The energy ratio (ΔdB) and the timing relationship (ΔT) between the first and second highlights were manipulated. An iso-sensitivity function was derived using a factorial design testing ΔdB at -10, -15, -20, and -25 dB and ΔT at 10, 20, 40, and 80 μs. The results suggest that the animal processed multiple echo highlights as separable analyzable features in the discrimination task, perhaps perceived through differences in spectral rippling across the duration of the echoes.

Beaked whales echolocate on prey.

PubMed Central

Johnson, Mark; Madsen, Peter T; Zimmer, Walter M X; de Soto, Natacha Aguilar; Tyack, Peter L

2004-01-01

Beaked whales (Cetacea: Ziphiidea) of the genera Ziphius and Mesoplodon are so difficult to study that they are mostly known from strandings. How these elusive toothed whales use and react to sound is of concern because they mass strand during naval sonar exercises. A new non-invasive acoustic ording tag was attached to four beaked whales(two Mesoplodon densirostris and two Ziphius cavirostris) and recorded high-frequency clicks during deep dives. The tagged whales only clicked at depths below 200 m, down to a maximum depth of 1267 m. Both species produced a large number of short, directional, ultrasonic clicks with significant energy below 20 kHz. The tags recorded echoes from prey items; to our knowledge, a first for any animal echolocating in the wild. As far as we are aware, these echoes provide the first direct evidence on how free-ranging toothed whales use echolocation in foraging. The strength of these echoes suggests that the source level of Mesoplodon clicks is in the range of 200-220 dB re 1 microPa at 1 m.This paper presents conclusive data on the normal vocalizations of these beaked whale species, which may enable acoustic monitoring to mitigate exposure to sounds intense enough to harm them. PMID:15801582

Acoustic features of objects matched by an echolocating bottlenose dolphin.

PubMed

Delong, Caroline M; Au, Whitlow W L; Lemonds, David W; Harley, Heidi E; Roitblat, Herbert L

2006-03-01

The focus of this study was to investigate how dolphins use acoustic features in returning echolocation signals to discriminate among objects. An echolocating dolphin performed a match-to-sample task with objects that varied in size, shape, material, and texture. After the task was completed, the features of the object echoes were measured (e.g., target strength, peak frequency). The dolphin's error patterns were examined in conjunction with the between-object variation in acoustic features to identify the acoustic features that the dolphin used to discriminate among the objects. The present study explored two hypotheses regarding the way dolphins use acoustic information in echoes: (1) use of a single feature, or (2) use of a linear combination of multiple features. The results suggested that dolphins do not use a single feature across all object sets or a linear combination of six echo features. Five features appeared to be important to the dolphin on four or more sets: the echo spectrum shape, the pattern of changes in target strength and number of highlights as a function of object orientation, and peak and center frequency. These data suggest that dolphins use multiple features and integrate information across echoes from a range of object orientations.

Human listening studies reveal insights into object features extracted by echolocating dolphins

NASA Astrophysics Data System (ADS)

Delong, Caroline M.; Au, Whitlow W. L.; Roitblat, Herbert L.

2004-05-01

Echolocating dolphins extract object feature information from the acoustic parameters of object echoes. However, little is known about which object features are salient to dolphins or how they extract those features. To gain insight into how dolphins might be extracting feature information, human listeners were presented with echoes from objects used in a dolphin echoic-visual cross-modal matching task. Human participants performed a task similar to the one the dolphin had performed; however, echoic samples consisting of 23-echo trains were presented via headphones. The participants listened to the echoic sample and then visually selected the correct object from among three alternatives. The participants performed as well as or better than the dolphin (M=88.0% correct), and reported using a combination of acoustic cues to extract object features (e.g., loudness, pitch, timbre). Participants frequently reported using the pattern of aural changes in the echoes across the echo train to identify the shape and structure of the objects (e.g., peaks in loudness or pitch). It is likely that dolphins also attend to the pattern of changes across echoes as objects are echolocated from different angles.

Functional Organization and Dynamic Activity in the Superior Colliculus of the Echolocating Bat, Eptesicus fuscus.

PubMed

Wohlgemuth, Melville J; Kothari, Ninad B; Moss, Cynthia F

2018-01-03

Sensory-guided behaviors require the transformation of sensory information into task-specific motor commands. Prior research on sensorimotor integration has emphasized visuomotor processes in the context of simplified orienting movements in controlled laboratory tasks rather than an animal's more complete, natural behavioral repertoire. Here, we conducted a series of neural recording experiments in the midbrain superior colliculus (SC) of echolocating bats engaged in a sonar target-tracking task that invoked dynamic active sensing behaviors. We hypothesized that SC activity in freely behaving animals would reveal dynamic shifts in neural firing patterns within and across sensory, sensorimotor, and premotor layers. We recorded neural activity in the SC of freely echolocating bats (three females and one male) and replicated the general trends reported in other species with sensory responses in the dorsal divisions and premotor activity in ventral divisions of the SC. However, within this coarse functional organization, we discovered that sensory and motor neurons are comingled within layers throughout the volume of the bat SC. In addition, as the bat increased pulse rate adaptively to increase resolution of the target location with closing distance, the activity of sensory and vocal premotor neurons changed such that auditory response times decreased, and vocal premotor lead times shortened. This finding demonstrates that SC activity can be modified dynamically in concert with adaptive behaviors and suggests that an integrated functional organization within SC laminae supports rapid and local integration of sensory and motor signals for natural, adaptive behaviors. SIGNIFICANCE STATEMENT Natural sensory-guided behaviors involve the rapid integration of information from the environment to direct flexible motor actions. The vast majority of research on sensorimotor integration has used artificial stimuli and simplified behaviors, leaving open questions about nervous system function in the context of natural tasks. Our work investigated mechanisms of dynamic sensorimotor feedback control by analyzing patterns of neural activity in the midbrain superior colliculus (SC) of an echolocating bat tracking and intercepting moving prey. Recordings revealed that sensory and motor neurons comingle within laminae of the SC to support rapid sensorimotor integration. Further, we discovered that neural activity in the bat SC changes with dynamic adaptations in the animal's echolocation behavior. Copyright © 2018 the authors 0270-6474/18/380245-12$15.00/0.

Extremely high frequency sensitivity in a 'simple' ear.

PubMed

Moir, Hannah M; Jackson, Joseph C; Windmill, James F C

2013-08-23

An evolutionary war is being played out between the bat, which uses ultrasonic calls to locate insect prey, and the moth, which uses microscale ears to listen for the approaching bat. While the highest known frequency of bat echolocation calls is 212 kHz, the upper limit of moth hearing is considered much lower. Here, we show that the greater wax moth, Galleria mellonella, is capable of hearing ultrasonic frequencies approaching 300 kHz; the highest frequency sensitivity of any animal. With auditory frequency sensitivity that is unprecedented in the animal kingdom, the greater wax moth is ready and armed for any echolocation call adaptations made by the bat in the on-going bat-moth evolutionary war.

Cloud Model Bat Algorithm

PubMed Central

Zhou, Yongquan; Xie, Jian; Li, Liangliang; Ma, Mingzhi

2014-01-01

Bat algorithm (BA) is a novel stochastic global optimization algorithm. Cloud model is an effective tool in transforming between qualitative concepts and their quantitative representation. Based on the bat echolocation mechanism and excellent characteristics of cloud model on uncertainty knowledge representation, a new cloud model bat algorithm (CBA) is proposed. This paper focuses on remodeling echolocation model based on living and preying characteristics of bats, utilizing the transformation theory of cloud model to depict the qualitative concept: “bats approach their prey.” Furthermore, Lévy flight mode and population information communication mechanism of bats are introduced to balance the advantage between exploration and exploitation. The simulation results show that the cloud model bat algorithm has good performance on functions optimization. PMID:24967425

Does nasal echolocation influence the modularity of the mammal skull?

PubMed

Santana, S E; Lofgren, S E

2013-11-01

In vertebrates, changes in cranial modularity can evolve rapidly in response to selection. However, mammals have apparently maintained their pattern of cranial integration throughout their evolutionary history and across tremendous morphological and ecological diversity. Here, we use phylogenetic, geometric morphometric and comparative analyses to test the hypothesis that the modularity of the mammalian skull has been remodelled in rhinolophid bats due to the novel and critical function of the nasal cavity in echolocation. We predicted that nasal echolocation has resulted in the evolution of a third cranial module, the 'nasal dome', in addition to the braincase and rostrum modules, which are conserved across mammals. We also test for similarities in the evolution of skull shape in relation to habitat across rhinolophids. We find that, despite broad variation in the shape of the nasal dome, the integration of the rhinolophid skull is highly consistent with conserved patterns of modularity found in other mammals. Across their broad geographical distribution, cranial shape in rhinolophids follows two major divisions that could reflect adaptations to dietary and environmental differences in African versus South Asian distributions. Our results highlight the potential of a relatively simple modular template to generate broad morphological and functional variation in mammals. © 2013 The Authors. Journal of Evolutionary Biology © 2013 European Society For Evolutionary Biology.

Calling louder and longer: how bats use biosonar under severe acoustic interference from other bats

PubMed Central

Amichai, Eran; Blumrosen, Gaddi; Yovel, Yossi

2015-01-01

Active-sensing systems such as echolocation provide animals with distinct advantages in dark environments. For social animals, however, like many bat species, active sensing can present problems as well: when many individuals emit bio-sonar calls simultaneously, detecting and recognizing the faint echoes generated by one's own calls amid the general cacophony of the group becomes challenging. This problem is often termed ‘jamming’ and bats have been hypothesized to solve it by shifting the spectral content of their calls to decrease the overlap with the jamming signals. We tested bats’ response in situations of extreme interference, mimicking a high density of bats. We played-back bat echolocation calls from multiple speakers, to jam flying Pipistrellus kuhlii bats, simulating a naturally occurring situation of many bats flying in proximity. We examined behavioural and echolocation parameters during search phase and target approach. Under severe interference, bats emitted calls of higher intensity and longer duration, and called more often. Slight spectral shifts were observed but they did not decrease the spectral overlap with jamming signals. We also found that pre-existing inter-individual spectral differences could allow self-call recognition. Results suggest that the bats’ response aimed to increase the signal-to-noise ratio and not to avoid spectral overlap. PMID:26702045

Compensating for the effects of site and equipment variation on delphinid species identification from their echolocation clicks.

PubMed

Roch, Marie A; Stinner-Sloan, Johanna; Baumann-Pickering, Simone; Wiggins, Sean M

2015-01-01

A concern for applications of machine learning techniques to bioacoustics is whether or not classifiers learn the categories for which they were trained. Unfortunately, information such as characteristics of specific recording equipment or noise environments can also be learned. This question is examined in the context of identifying delphinid species by their echolocation clicks. To reduce the ambiguity between species classification performance and other confounding factors, species whose clicks can be readily distinguished were used in this study: Pacific white-sided and Risso's dolphins. A subset of data from autonomous acoustic recorders located at seven sites in the Southern California Bight collected between 2006 and 2012 was selected. Cepstral-based features were extracted for each echolocation click and Gaussian mixture models were used to classify groups of 100 clicks. One hundred Monte-Carlo three-fold experiments were conducted to examine classification performance where fold composition was determined by acoustic encounter, recorder characteristics, or recording site. The error rate increased from 6.1% when grouped by acoustic encounter to 18.1%, 46.2%, and 33.2% for grouping by equipment, equipment category, and site, respectively. A noise compensation technique reduced error for these grouping schemes to 2.7%, 4.4%, 6.7%, and 11.4%, respectively, a reduction in error rate of 56%-86%.

Multi-component separation and analysis of bat echolocation calls.

PubMed

DiCecco, John; Gaudette, Jason E; Simmons, James A

2013-01-01

The vast majority of animal vocalizations contain multiple frequency modulated (FM) components with varying amounts of non-linear modulation and harmonic instability. This is especially true of biosonar sounds where precise time-frequency templates are essential for neural information processing of echoes. Understanding the dynamic waveform design by bats and other echolocating animals may help to improve the efficacy of man-made sonar through biomimetic design. Bats are known to adapt their call structure based on the echolocation task, proximity to nearby objects, and density of acoustic clutter. To interpret the significance of these changes, a method was developed for component separation and analysis of biosonar waveforms. Techniques for imaging in the time-frequency plane are typically limited due to the uncertainty principle and interference cross terms. This problem is addressed by extending the use of the fractional Fourier transform to isolate each non-linear component for separate analysis. Once separated, empirical mode decomposition can be used to further examine each component. The Hilbert transform may then successfully extract detailed time-frequency information from each isolated component. This multi-component analysis method is applied to the sonar signals of four species of bats recorded in-flight by radiotelemetry along with a comparison of other common time-frequency representations.

Hearing sensitivity during target presence and absence while a whale echolocates.

PubMed

Supin, Alexander Ya; Nachtigall, Paul E; Breese, Marlee

2008-01-01

Hearing sensitivity was measured in a false killer whale during echolocation. Sensitivity was measured using probe stimuli as sinusoidally amplitude modulated signals with a 22.5-kHz carrier frequency and recording auditory evoked potentials as envelope-following responses. The probes were presented and responses were recorded during short 2-s periods when the animal echolocated to detect the presence or absence of a target in a go/no-go paradigm. In the target-absent trials, a hearing threshold of 90.4 dB re 1 muPa was found; in the target-present trials, the threshold was 109.8 dB. Thus, a 19.4-dB difference was found between thresholds in the target-present and target-absent trials. To check the possibility that this difference was the result of different masking degree of the probe by the emitted sonar clicks, click statistics were investigated in similar trials. No indication was found that the energy of the emitted clicks was higher in the target-present than in target-absent trials; on the contrary, mean click level, mean number of clicks per train, and overall train energy was slightly higher in the target-absent trials. Thus the data indicate that the hearing sensitivity of the whale varied depending on target presence or absence.

Sensing in a noisy world: lessons from auditory specialists, echolocating bats.

PubMed

Corcoran, Aaron J; Moss, Cynthia F

2017-12-15

All animals face the essential task of extracting biologically meaningful sensory information from the 'noisy' backdrop of their environments. Here, we examine mechanisms used by echolocating bats to localize objects, track small prey and communicate in complex and noisy acoustic environments. Bats actively control and coordinate both the emission and reception of sound stimuli through integrated sensory and motor mechanisms that have evolved together over tens of millions of years. We discuss how bats behave in different ecological scenarios, including detecting and discriminating target echoes from background objects, minimizing acoustic interference from competing conspecifics and overcoming insect noise. Bats tackle these problems by deploying a remarkable array of auditory behaviors, sometimes in combination with the use of other senses. Behavioral strategies such as ceasing sonar call production and active jamming of the signals of competitors provide further insight into the capabilities and limitations of echolocation. We relate these findings to the broader topic of how animals extract relevant sensory information in noisy environments. While bats have highly refined abilities for operating under noisy conditions, they face the same challenges encountered by many other species. We propose that the specialized sensory mechanisms identified in bats are likely to occur in analogous systems across the animal kingdom. © 2017. Published by The Company of Biologists Ltd.

Negative long term effects on harbour porpoises from a large scale offshore wind farm in the Baltic—evidence of slow recovery

NASA Astrophysics Data System (ADS)

Teilmann, Jonas; Carstensen, Jacob

2012-12-01

Offshore wind farms constitute a new and fast growing industry all over the world. This study investigates the long term impact on harbour porpoises, Phocoena phocoena, for more than 10 years (2001-12) from the first large scale offshore wind farm in the world, Nysted Offshore Wind Farm, in the Danish western Baltic Sea (72 × 2.3 MW turbines). The wind farm was brought into full operation in December 2003. At six stations, acoustic porpoise detectors (T-PODs) were placed inside the wind farm area and at a reference area 10 km to the east, to monitor porpoise echolocation activity as a proxy of porpoise presence. A modified statistical BACI design was applied to detect changes in porpoise presence before, during and after construction of the wind farm. The results show that the echolocation activity has significantly declined inside Nysted Offshore Wind Farm since the baseline in 2001-2 and has not fully recovered yet. The echolocation activity inside the wind farm has been gradually increasing (from 11% to 29% of the baseline level) since the construction of the wind farm, possibly due to habituation of the porpoises to the wind farm or enrichment of the environment due to reduced fishing and to artificial reef effects.

Calling louder and longer: how bats use biosonar under severe acoustic interference from other bats.

PubMed

Amichai, Eran; Blumrosen, Gaddi; Yovel, Yossi

2015-12-22

Active-sensing systems such as echolocation provide animals with distinct advantages in dark environments. For social animals, however, like many bat species, active sensing can present problems as well: when many individuals emit bio-sonar calls simultaneously, detecting and recognizing the faint echoes generated by one's own calls amid the general cacophony of the group becomes challenging. This problem is often termed 'jamming' and bats have been hypothesized to solve it by shifting the spectral content of their calls to decrease the overlap with the jamming signals. We tested bats' response in situations of extreme interference, mimicking a high density of bats. We played-back bat echolocation calls from multiple speakers, to jam flying Pipistrellus kuhlii bats, simulating a naturally occurring situation of many bats flying in proximity. We examined behavioural and echolocation parameters during search phase and target approach. Under severe interference, bats emitted calls of higher intensity and longer duration, and called more often. Slight spectral shifts were observed but they did not decrease the spectral overlap with jamming signals. We also found that pre-existing inter-individual spectral differences could allow self-call recognition. Results suggest that the bats' response aimed to increase the signal-to-noise ratio and not to avoid spectral overlap. © 2015 The Author(s).

A deterministic compressive sensing model for bat biosonar.

PubMed

Hague, David A; Buck, John R; Bilik, Igal

2012-12-01

The big brown bat (Eptesicus fuscus) uses frequency modulated (FM) echolocation calls to accurately estimate range and resolve closely spaced objects in clutter and noise. They resolve glints spaced down to 2 μs in time delay which surpasses what traditional signal processing techniques can achieve using the same echolocation call. The Matched Filter (MF) attains 10-12 μs resolution while the Inverse Filter (IF) achieves higher resolution at the cost of significantly degraded detection performance. Recent work by Fontaine and Peremans [J. Acoustic. Soc. Am. 125, 3052-3059 (2009)] demonstrated that a sparse representation of bat echolocation calls coupled with a decimating sensing method facilitates distinguishing closely spaced objects over realistic SNRs. Their work raises the intriguing question of whether sensing approaches structured more like a mammalian auditory system contains the necessary information for the hyper-resolution observed in behavioral tests. This research estimates sparse echo signatures using a gammatone filterbank decimation sensing method which loosely models the processing of the bat's auditory system. The decimated filterbank outputs are processed with [script-l](1) minimization. Simulations demonstrate that this model maintains higher resolution than the MF and significantly better detection performance than the IF for SNRs of 5-45 dB while undersampling the return signal by a factor of six.

Long-Term Monitoring of Dolphin Biosonar Activity in Deep Pelagic Waters of the Mediterranean Sea.

PubMed

Caruso, Francesco; Alonge, Giuseppe; Bellia, Giorgio; De Domenico, Emilio; Grammauta, Rosario; Larosa, Giuseppina; Mazzola, Salvatore; Riccobene, Giorgio; Pavan, Gianni; Papale, Elena; Pellegrino, Carmelo; Pulvirenti, Sara; Sciacca, Virginia; Simeone, Francesco; Speziale, Fabrizio; Viola, Salvatore; Buscaino, Giuseppa

2017-06-28

Dolphins emit short ultrasonic pulses (clicks) to acquire information about the surrounding environment, prey and habitat features. We investigated Delphinidae activity over multiple temporal scales through the detection of their echolocation clicks, using long-term Passive Acoustic Monitoring (PAM). The Istituto Nazionale di Fisica Nucleare operates multidisciplinary seafloor observatories in a deep area of the Central Mediterranean Sea. The Ocean noise Detection Experiment collected data offshore the Gulf of Catania from January 2005 to November 2006, allowing the study of temporal patterns of dolphin activity in this deep pelagic zone for the first time. Nearly 5,500 five-minute recordings acquired over two years were examined using spectrogram analysis and through development and testing of an automatic detection algorithm. Echolocation activity of dolphins was mostly confined to nighttime and crepuscular hours, in contrast with communicative signals (whistles). Seasonal variation, with a peak number of clicks in August, was also evident, but no effect of lunar cycle was observed. Temporal trends in echolocation corresponded to environmental and trophic variability known in the deep pelagic waters of the Ionian Sea. Long-term PAM and the continued development of automatic analysis techniques are essential to advancing the study of pelagic marine mammal distribution and behaviour patterns.

Acoustic mirror effect increases prey detection distance in trawling bats

NASA Astrophysics Data System (ADS)

Siemers, Björn M.; Baur, Eric; Schnitzler, Hans-Ulrich

2005-06-01

Many different and phylogenetically distant species of bats forage for insects above water bodies and take insects from and close to the surface; the so-called ‘trawling behaviour’. Detection of surface-based prey by echolocation is facilitated by acoustically smooth backgrounds such as water surfaces that reflect sound impinging at an acute angle away from the bat and thereby render a prey object acoustically conspicuous. Previous measurements had shown that the echo amplitude of a target on a smooth surface is higher than that of the same target in mid-air, due to an acoustic mirror effect. In behavioural experiments with three pond bats (Myotis dasycneme), we tested the hypothesis that the maximum distances at which bats can detect prey are larger for prey on smooth surfaces than for the same prey in an airborne situation. We determined the moment of prey detection from a change in echolocation behaviour and measured the detection distance in 3D space from IR-video recordings using stereo-photogrammetry. The bats showed the predicted increase in detection distance for prey on smooth surfaces. The acoustic mirror effect therefore increases search efficiency and contributes to the acoustic advantages encountered by echolocating bats when foraging at low heights above smooth water surfaces. These acoustic advantages may have favoured the repeated evolution of trawling behaviour.

The effect of climate on acoustic signals: does atmospheric sound absorption matter for bird song and bat echolocation?

PubMed

Snell-Rood, Emilie C

2012-02-01

The divergence of signals along ecological gradients may lead to speciation. The current research tests the hypothesis that variation in sound absorption selects for divergence in acoustic signals along climatic gradients, which has implications for understanding not only diversification, but also how organisms may respond to climate change. Because sound absorption varies with temperature, humidity, and the frequency of sound, individuals or species may vary signal structure with changes in climate over space or time. In particular, signals of lower frequency, narrower bandwidth, and longer duration should be more detectable in environments with high sound absorption. Using both North American wood warblers (Parulidae) and bats of the American Southwest, this work found evidence of associations between signal structure and sound absorption. Warbler species with higher mean absorption across their range were more likely to have narrow bandwidth songs. Bat species found in higher absorption habitats were more likely to have lower frequency echolocation calls. In addition, bat species changed echolocation call structure across seasons, using longer duration, lower frequency calls in the higher absorption rainy season. These results suggest that signals may diverge along climatic gradients due to variation in sound absorption, although the effects of absorption are modest. © 2012 Acoustical Society of America

Acoustic mirror effect increases prey detection distance in trawling bats.

PubMed

Siemers, Björn M; Baur, Eric; Schnitzler, Hans-Ulrich

2005-06-01

Many different and phylogenetically distant species of bats forage for insects above water bodies and take insects from and close to the surface; the so-called 'trawling behaviour'. Detection of surface-based prey by echolocation is facilitated by acoustically smooth backgrounds such as water surfaces that reflect sound impinging at an acute angle away from the bat and thereby render a prey object acoustically conspicuous. Previous measurements had shown that the echo amplitude of a target on a smooth surface is higher than that of the same target in mid-air, due to an acoustic mirror effect. In behavioural experiments with three pond bats (Myotis dasycneme), we tested the hypothesis that the maximum distances at which bats can detect prey are larger for prey on smooth surfaces than for the same prey in an airborne situation. We determined the moment of prey detection from a change in echolocation behaviour and measured the detection distance in 3D space from IR-video recordings using stereo-photogrammetry. The bats showed the predicted increase in detection distance for prey on smooth surfaces. The acoustic mirror effect therefore increases search efficiency and contributes to the acoustic advantages encountered by echolocating bats when foraging at low heights above smooth water surfaces. These acoustic advantages may have favoured the repeated evolution of trawling behaviour.

Effect of echolocation behavior-related constant frequency-frequency modulation sound on the frequency tuning of inferior collicular neurons in Hipposideros armiger.

PubMed

Tang, Jia; Fu, Zi-Ying; Wei, Chen-Xue; Chen, Qi-Cai

2015-08-01

In constant frequency-frequency modulation (CF-FM) bats, the CF-FM echolocation signals include both CF and FM components, yet the role of such complex acoustic signals in frequency resolution by bats remains unknown. Using CF and CF-FM echolocation signals as acoustic stimuli, the responses of inferior collicular (IC) neurons of Hipposideros armiger were obtained by extracellular recordings. We tested the effect of preceding CF or CF-FM sounds on the shape of the frequency tuning curves (FTCs) of IC neurons. Results showed that both CF-FM and CF sounds reduced the number of FTCs with tailed lower-frequency-side of IC neurons. However, more IC neurons experienced such conversion after adding CF-FM sound compared with CF sound. We also found that the Q 20 value of the FTC of IC neurons experienced the largest increase with the addition of CF-FM sound. Moreover, only CF-FM sound could cause an increase in the slope of the neurons' FTCs, and such increase occurred mainly in the lower-frequency edge. These results suggested that CF-FM sound could increase the accuracy of frequency analysis of echo and cut-off low-frequency elements from the habitat of bats more than CF sound.

The echolocation transmission beam of free-ranging Indo-Pacific humpback dolphins (Sousa chinensis).

PubMed

Fang, Liang; Wu, Yuping; Wang, Kexiong; Pine, Matthew K; Wang, Ding; Li, Songhai

2017-08-01

While the transmission beam of odontocetes has been described in a number of studies, the majority of them that have measured the transmission beam in two dimensions were focused on captive animals. Within the current study, a dedicated cross hydrophone array with nine elements was used to investigate the echolocation transmission beam of free-ranging Indo-Pacific humpback dolphins. A total of 265 on-axis clicks were analyzed, from which the apparent peak to peak source levels ranged between 168 to 207 dB (mean 184.5 dB ± 6.6 dB). The 3-dB beam width along the horizontal and vertical plane was 9.6° and 7.4°, respectively. Measured separately, the directivity index of the horizontal and vertical plane was 12.6 and 13.5 dB, respectively, and the overall directivity index (both planes combined) was 29.5 dB. The beam shape was slightly asymmetrical along the horizontal and vertical axis. Compared to other species, the characteristics of the transmitting beam of Indo-Pacific humpback dolphins were relatively close to the bottlenose dolphin (Tursiops truncatus), likely due to the similarity in the peak frequency and waveform of echolocation clicks and comparable body sizes of the two species.

Discriminating features of echolocation clicks of melon-headed whales (Peponocephala electra), bottlenose dolphins (Tursiops truncatus), and Gray's spinner dolphins (Stenella longirostris longirostris).

PubMed

Baumann-Pickering, Simone; Wiggins, Sean M; Hildebrand, John A; Roch, Marie A; Schnitzler, Hans-Ulrich

2010-10-01

Spectral parameters were used to discriminate between echolocation clicks produced by three dolphin species at Palmyra Atoll: melon-headed whales (Peponocephala electra), bottlenose dolphins (Tursiops truncatus) and Gray's spinner dolphins (Stenella longirostris longirostris). Single species acoustic behavior during daytime observations was recorded with a towed hydrophone array sampling at 192 and 480 kHz. Additionally, an autonomous, bottom moored High-frequency Acoustic Recording Package (HARP) collected acoustic data with a sampling rate of 200 kHz. Melon-headed whale echolocation clicks had the lowest peak and center frequencies, spinner dolphins had the highest frequencies and bottlenose dolphins were nested in between these two species. Frequency differences were significant. Temporal parameters were not well suited for classification. Feature differences were enhanced by reducing variability within a set of single clicks by calculating mean spectra for groups of clicks. Median peak frequencies of averaged clicks (group size 50) of melon-headed whales ranged between 24.4 and 29.7 kHz, of bottlenose dolphins between 26.7 and 36.7 kHz, and of spinner dolphins between 33.8 and 36.0 kHz. Discriminant function analysis showed the ability to correctly discriminate between 93% of melon-headed whales, 75% of spinner dolphins and 54% of bottlenose dolphins.

Evolutionary changes of the importance of olfaction in cetaceans based on the olfactory marker protein gene.

PubMed

Kishida, Takushi; Thewissen, J G M

2012-01-25

Odontocetes and mysticetes are two extant suborders of cetaceans. It is reported that the former have no sense of olfaction, while the latter can smell in air. To explain the ecological reason why mysticetes still retain their sense of smell, two hypotheses have been proposed - the echolocation-priority hypothesis, which assumes that the acquisition of echolocation causes the reduction of the importance of olfaction, and the filter-feeder hypothesis, which assumes that olfactory ability is important for filter-feeders to locate their prey because clouds of plankton give off a peculiar odor. The olfactory marker protein (OMP) is almost exclusively expressed in vertebrate olfactory receptor neurons, and is considered to play important roles in olfactory systems. In this study, full-length open reading frames of OMP genes were identified in 6 cetacean species and we analyzed the nonsynonymous to synonymous substitution rate ratio based on the maximum likelihood method. The evolutionary changes of the selective pressures on OMP genes did fit better to the filter-feeder hypothesis than to the echolocation-priority hypothesis. In addition, no pseudogenization mutations are found in all five odontocetes OMP genes investigated in this study. It may suggest that OMP retains some function even in 'anosmic' odontocetes. Copyright © 2011 Elsevier B.V. All rights reserved.

[Echolocation calls of free-flying Himalayan swiftlets (Aerodramus brevirostris)].

PubMed

Wang, Bin; Ma, Jian-Zhang; Chen, Yi; Tan, Liang-Jing; Liu, Qi; Shen, Qi-Qi; Liao, Qing-Yi; Zhang, Li-Biao

2013-02-01

Here, we present our findings of free-flying echolocation calls of Himalayan swiftlets (Aerodramus brevirostris), which were recorded in Shenjing Cave, Hupingshan National Reserve, Shimen County, Hunan Province in June 2012, using Avisoft-UltraSoundGate 116(e). We noted that after foraging at dusk, the Himalayan swiftlets flew fast into the cave without clicks, and then slowed down in dark area in the cave, but with sounds. The echolocation sounds of Himalayan swiftlets are broadband, double noise burst clicks, separated by a short pause. The inter-pulse intervals between double clicks (99.3±3.86 ms)were longer than those within double clicks (6.6±0.42 ms) (P<0.01). With the exception of peak frequency, between 6.2±0.08 kHz and 6.2±0.10 kHz, (P>0.05) and pulse duration 2.9±0.12 ms, 3.2±0.17 ms, (P>0.05) between the first and second, other factors-maximum frequency, minimum frequency, frequency bandwidth, and power-were significantly different between the clicks. The maximum frequency of the first pulse (20.1±1.10 kHz) was higher than that of second (15.4±0.98 kHz) (P<0.01), while the minimum frequency of the first pulse (3.7±0.12 kHz) was lower than that of second (4.0±0.09 kHz) (P<0.05); resulting in the frequency bandwidth of the first pulse (16.5±1.17 kHz) longer than that of second (11.4±1.01 kHz) (P<0.01). The power of the first pulse (-32.5±0.60 dB) was higher than that of second (-35.2±0.94 dB) (P<0.05). More importantly, we found that Himalayan swiftlets emitted echolocation pulses including ultrasonic sound, with a maximum frequency reaching 33.2 kHz.

The Dolphin Sonar: Excellent Capabilities In Spite of Some Mediocre Properties

NASA Astrophysics Data System (ADS)

Au, Whitlow W. L.

2004-11-01

Dolphin sonar research has been conducted for several decades and much has been learned about the capabilities of echolocating dolphins to detect, discriminate and recognize underwater targets. The results of these research projects suggest that dolphins possess the most sophisticated of all sonar for short ranges and shallow water where reverberation and clutter echoes are high. The critical feature of the dolphin sonar is the capability of discriminating and recognizing complex targets in a highly reverberant and noisy environment. The dolphin's detection threshold in reverberation occurs at a echo-to reverberation ratio of approximately 4 dB. Echolocating dolphins also have the capability to make fine discriminate of target properties such as wall thickness difference of water-filled cylinders and material differences in metallic plates. The high-resolution property of the animal's echolocation signals and the high dynamic range of its auditory system are important factors in their outstanding discrimination capabilities. In the wall thickness discrimination of cylinder experiment, time differences between echo highlights at small as 500-600 ns can be resolved by echolocating dolphins. Measurements of the targets used in the metallic plate composition experiment suggest that dolphins attended to echo components that were 20-30 dB below the maximum level for a specific target. It is interesting to realize that some of the properties of the dolphin sonar system are fairly mediocre, yet the total performance of the system is often outstanding. When compared to some technological sonar, the energy content of the dolphin sonar signal is not very high, the transmission and receiving beamwidths are fairly large, and the auditory filters are not very narrow. Yet the dolphin sonar has demonstrated excellent capabilities in spite the mediocre features of its "hardware." Reasons why dolphins can perform complex sonar task will be discussed in light of the "equipment" they possess.

Following a Foraging Fish-Finder: Diel Habitat Use of Blainville's Beaked Whales Revealed by Echolocation

PubMed Central

Arranz, Patricia; de Soto, Natacha Aguilar; Madsen, Peter T.; Brito, Alberto; Bordes, Fernando; Johnson, Mark P.

2011-01-01

Simultaneous high resolution sampling of predator behavior and habitat characteristics is often difficult to achieve despite its importance in understanding the foraging decisions and habitat use of predators. Here we tap into the biosonar system of Blainville's beaked whales, Mesoplodon densirostris, using sound and orientation recording tags to uncover prey-finding cues available to echolocating predators in the deep-sea. Echolocation sounds indicate where whales search and encounter prey, as well as the altitude of whales above the sea-floor and the density of organisms around them, providing a link between foraging activity and the bio-physical environment. Tagged whales (n = 9) hunted exclusively at depth, investing most of their search time either in the lower part of the deep scattering layer (DSL) or near the sea-floor with little diel change. At least 43% (420/974) of recorded prey-capture attempts were performed within the benthic boundary layer despite a wide range of dive depths, and many dives included both meso- and bentho-pelagic foraging. Blainville's beaked whales only initiate searching when already deep in the descent and encounter prey suitable for capture within 2 min of the start of echolocation, suggesting that these whales are accessing prey in reliable vertical strata. Moreover, these prey resources are sufficiently dense to feed the animals in what is effectively four hours of hunting per day enabling a strategy in which long dives to exploit numerous deep-prey with low nutritional value require protracted recovery periods (average 1.5 h) between dives. This apparent searching efficiency maybe aided by inhabiting steep undersea slopes with access to both the DSL and the sea-floor over small spatial scales. Aggregations of prey in these biotopes are located using biosonar-derived landmarks and represent stable and abundant resources for Blainville's beaked whales in the otherwise food-limited deep-ocean. PMID:22163295

Extremely high frequency sensitivity in a ‘simple’ ear

PubMed Central

Moir, Hannah M.; Jackson, Joseph C.; Windmill, James F. C.

2013-01-01

An evolutionary war is being played out between the bat, which uses ultrasonic calls to locate insect prey, and the moth, which uses microscale ears to listen for the approaching bat. While the highest known frequency of bat echolocation calls is 212 kHz, the upper limit of moth hearing is considered much lower. Here, we show that the greater wax moth, Galleria mellonella, is capable of hearing ultrasonic frequencies approaching 300 kHz; the highest frequency sensitivity of any animal. With auditory frequency sensitivity that is unprecedented in the animal kingdom, the greater wax moth is ready and armed for any echolocation call adaptations made by the bat in the on-going bat–moth evolutionary war. PMID:23658005

Bat noseleaf model: echolocation function, design considerations, and experimental verification.

PubMed

Kuc, Roman

2011-05-01

This paper describes a possible bat noseleaf echolocation function that improves target elevation resolution. Bats with a protruding noseleaf can rotate the lancet to act as an acoustic mirror that reflects the nostril emission, modeled as a virtual nostril that produces a delayed emission. The cancellation of the nostril and virtual nostril components at a target produces a sharp spectral notch whose frequency location relates to target elevation. This notch can be observed directly from the swept-frequency emission waveform, suggesting cochlear processing capabilities. Physical acoustic principles indicate the design considerations and trade-offs that a bat can accomplish through noseleaf shape and emission characteristics. An experimental model verifies the analysis and exhibits an elevation versus notch frequency sensitivity of approximately 1°/kHz.

Source parameter estimates of echolocation clicks from wild pygmy killer whales (Feresa attenuata) (L)

NASA Astrophysics Data System (ADS)

Madsen, P. T.; Kerr, I.; Payne, R.

2004-10-01

Pods of the little known pygmy killer whale (Feresa attenuata) in the northern Indian Ocean were recorded with a vertical hydrophone array connected to a digital recorder sampling at 320 kHz. Recorded clicks were directional, short (25 μs) transients with estimated source levels between 197 and 223 dB re. 1 μPa (pp). Spectra of clicks recorded close to or on the acoustic axis were bimodal with peak frequencies between 45 and 117 kHz, and with centroid frequencies between 70 and 85 kHz. The clicks share characteristics of echolocation clicks from similar sized, whistling delphinids, and have properties suited for the detection and classification of prey targeted by this odontocete. .

Active listening for spatial orientation in a complex auditory scene.

PubMed

Moss, Cynthia F; Bohn, Kari; Gilkenson, Hannah; Surlykke, Annemarie

2006-04-01

To successfully negotiate a complex environment, an animal must control the timing of motor behaviors in coordination with dynamic sensory information. Here, we report on adaptive temporal control of vocal-motor behavior in an echolocating bat, Eptesicus fuscus, as it captured tethered insects close to background vegetation. Recordings of the bat's sonar vocalizations were synchronized with high-speed video images that were used to reconstruct the bat's three-dimensional flight path and the positions of target and vegetation. When the bat encountered the difficult task of taking insects as close as 10-20 cm from the vegetation, its behavior changed significantly from that under open room conditions. Its success rate decreased by about 50%, its time to initiate interception increased by a factor of ten, and its high repetition rate "terminal buzz" decreased in duration by a factor of three. Under all conditions, the bat produced prominent sonar "strobe groups," clusters of echolocation pulses with stable intervals. In the final stages of insect capture, the bat produced strobe groups at a higher incidence when the insect was positioned near clutter. Strobe groups occurred at all phases of the wingbeat (and inferred respiration) cycle, challenging the hypothesis of strict synchronization between respiration and sound production in echolocating bats. The results of this study provide a clear demonstration of temporal vocal-motor control that directly impacts the signals used for perception.

Active Listening for Spatial Orientation in a Complex Auditory Scene

PubMed Central

Bohn, Kari; Gilkenson, Hannah; Surlykke, Annemarie

2006-01-01

To successfully negotiate a complex environment, an animal must control the timing of motor behaviors in coordination with dynamic sensory information. Here, we report on adaptive temporal control of vocal–motor behavior in an echolocating bat, Eptesicus fuscus, as it captured tethered insects close to background vegetation. Recordings of the bat's sonar vocalizations were synchronized with high-speed video images that were used to reconstruct the bat's three-dimensional flight path and the positions of target and vegetation. When the bat encountered the difficult task of taking insects as close as 10–20 cm from the vegetation, its behavior changed significantly from that under open room conditions. Its success rate decreased by about 50%, its time to initiate interception increased by a factor of ten, and its high repetition rate “terminal buzz” decreased in duration by a factor of three. Under all conditions, the bat produced prominent sonar “strobe groups,” clusters of echolocation pulses with stable intervals. In the final stages of insect capture, the bat produced strobe groups at a higher incidence when the insect was positioned near clutter. Strobe groups occurred at all phases of the wingbeat (and inferred respiration) cycle, challenging the hypothesis of strict synchronization between respiration and sound production in echolocating bats. The results of this study provide a clear demonstration of temporal vocal–motor control that directly impacts the signals used for perception. PMID:16509770

Passive Acoustic Monitoring the Diel, Lunar, Seasonal and Tidal Patterns in the Biosonar Activity of the Indo-Pacific Humpback Dolphins (Sousa chinensis) in the Pearl River Estuary, China.

PubMed

Wang, Zhi-Tao; Nachtigall, Paul E; Akamatsu, Tomonari; Wang, Ke-Xiong; Wu, Yu-Ping; Liu, Jian-Chang; Duan, Guo-Qin; Cao, Han-Jiang; Wang, Ding

2015-01-01

A growing demand for sustainable energy has led to an increase in construction of offshore windfarms. Guishan windmill farm will be constructed in the Pearl River Estuary, China, which sustains the world's largest known population of Indo-Pacific humpback dolphins (Sousa chinensis). Dolphin conservation is an urgent issue in this region. By using passive acoustic monitoring, a baseline distribution of data on this species in the Pearl River Estuary during pre-construction period had been collected. Dolphin biosonar detection and its diel, lunar, seasonal and tidal patterns were examined using a Generalized Linear Model. Significant higher echolocation detections at night than during the day, in winter-spring than in summer-autumn, at high tide than at flood tide were recognized. Significant higher echolocation detections during the new moon were recognized at night time. The diel, lunar and seasonal patterns for the echolocation encounter duration also significantly varied. These patterns could be due to the spatial-temporal variability of dolphin prey and illumination conditions. The baseline information will be useful for driving further effective action on the conservation of this species and in facilitating later assessments of the effects of the offshore windfarm on the dolphins by comparing the baseline to post construction and post mitigation efforts.

Echolocating bats rely on audiovocal feedback to adapt sonar signal design.

PubMed

Luo, Jinhong; Moss, Cynthia F

2017-10-10

Many species of bat emit acoustic signals and use information carried by echoes reflecting from nearby objects to navigate and forage. It is widely documented that echolocating bats adjust the features of sonar calls in response to echo feedback; however, it remains unknown whether audiovocal feedback contributes to sonar call design. Audiovocal feedback refers to the monitoring of one's own vocalizations during call production and has been intensively studied in nonecholocating animals. Audiovocal feedback not only is a necessary component of vocal learning but also guides the control of the spectro-temporal structure of vocalizations. Here, we show that audiovocal feedback is directly involved in the echolocating bat's control of sonar call features. As big brown bats tracked targets from a stationary position, we played acoustic jamming signals, simulating calls of another bat, timed to selectively perturb audiovocal feedback or echo feedback. We found that the bats exhibited the largest call-frequency adjustments when the jamming signals occurred during vocal production. By contrast, bats did not show sonar call-frequency adjustments when the jamming signals coincided with the arrival of target echoes. Furthermore, bats rapidly adapted sonar call design in the first vocalization following the jamming signal, revealing a response latency in the range of 66 to 94 ms. Thus, bats, like songbirds and humans, rely on audiovocal feedback to structure sonar signal design.

Determinants of echolocation call frequency variation in the Formosan lesser horseshoe bat (Rhinolophus monoceros)

PubMed Central

Chen, Shiang-Fan; Jones, Gareth; Rossiter, Stephen J.

2009-01-01

The origin and maintenance of intraspecific variation in vocal signals is important for population divergence and speciation. Where vocalizations are transmitted by vertical cultural inheritance, similarity will reflect co-ancestry, and thus vocal divergence should reflect genetic structure. Horseshoe bats are characterized by echolocation calls dominated by a constant frequency component that is partly determined by maternal imprinting. Although previous studies showed that constant frequency calls are also influenced by some non-genetic factors, it is not known how frequency relates to genetic structure. To test this, we related constant frequency variation to genetic and non-genetic variables in the Formosan lesser horseshoe bat (Rhinolophus monoceros). Recordings of bats from across Taiwan revealed that females called at higher frequencies than males; however, we found no effect of environmental or morphological factors on call frequency. By comparison, variation showed clear population structure, with frequencies lower in the centre and east, and higher in the north and south. Within these regions, frequency divergence was directional and correlated with geographical distance, suggesting that call frequencies are subject to cultural drift. However, microsatellite clustering analysis showed that broad differences in constant frequency among populations corresponded to discontinuities in allele frequencies resulting from vicariant events. Our results provide evidence that the processes shaping genetic subdivision have concomitant consequences for divergence in echolocation call frequency. PMID:19692399

Matched Behavioral and Neural Adaptations for Low Sound Level Echolocation in a Gleaning Bat, Antrozous pallidus.

PubMed

Measor, Kevin R; Leavell, Brian C; Brewton, Dustin H; Rumschlag, Jeffrey; Barber, Jesse R; Razak, Khaleel A

2017-01-01

In active sensing, animals make motor adjustments to match sensory inputs to specialized neural circuitry. Here, we describe an active sensing system for sound level processing. The pallid bat uses downward frequency-modulated (FM) sweeps as echolocation calls for general orientation and obstacle avoidance. The bat's auditory cortex contains a region selective for these FM sweeps (FM sweep-selective region, FMSR). We show that the vast majority of FMSR neurons are sensitive and strongly selective for relatively low levels (30-60 dB SPL). Behavioral testing shows that when a flying bat approaches a target, it reduces output call levels to keep echo levels between ∼30 and 55 dB SPL. Thus, the pallid bat behaviorally matches echo levels to an optimized neural representation of sound levels. FMSR neurons are more selective for sound levels of FM sweeps than tones, suggesting that across-frequency integration enhances level tuning. Level-dependent timing of high-frequency sideband inhibition in the receptive field shapes increased level selectivity for FM sweeps. Together with previous studies, these data indicate that the same receptive field properties shape multiple filters (sweep direction, rate, and level) for FM sweeps, a sound common in multiple vocalizations, including human speech. The matched behavioral and neural adaptations for low-intensity echolocation in the pallid bat will facilitate foraging with reduced probability of acoustic detection by prey.

Stimulation of the basal and central amygdala in the mustached bat triggers echolocation and agonistic vocalizations within multimodal output

PubMed Central

Ma, Jie; Kanwal, Jagmeet S.

2014-01-01

The neural substrate for the perception of vocalizations is relatively well described, but how their timing and specificity are tightly coupled with accompanying physiological changes and context-appropriate behaviors remains unresolved. We hypothesized that temporally integrated vocal and emotive responses, especially the expression of fear, vigilance and aggression, originate within the amygdala. To test this hypothesis, we performed electrical microstimulation at 461 highly restricted loci within the basal and central amygdala in awake mustached bats. At a subset of these sites, high frequency stimulation with weak constant current pulses presented at near-threshold levels triggered vocalization of either echolocation pulses or social calls. At the vast majority of locations, microstimulation produced a constellation of changes in autonomic and somatomotor outputs. These changes included widespread co-activation of significant tachycardia and hyperventilation and/or rhythmic ear pinna movements (PMs). In a few locations, responses were constrained to vocalization and/or PMs despite increases in the intensity of stimulation. The probability of eliciting echolocation pulses vs. social calls decreased in a medial-posterior to anterolateral direction within the centrobasal amygdala. Microinjections of kainic acid (KA) at stimulation sites confirmed the contribution of cellular activity rather than fibers-of-passage in the control of multimodal outputs. The results suggest that localized clusters of neurons may simultaneously modulate the activity of multiple central pattern generators (CPGs) present within the brainstem. PMID:24624089

Stimulation of the basal and central amygdala in the mustached bat triggers echolocation and agonistic vocalizations within multimodal output.

PubMed

Ma, Jie; Kanwal, Jagmeet S

2014-01-01

The neural substrate for the perception of vocalizations is relatively well described, but how their timing and specificity are tightly coupled with accompanying physiological changes and context-appropriate behaviors remains unresolved. We hypothesized that temporally integrated vocal and emotive responses, especially the expression of fear, vigilance and aggression, originate within the amygdala. To test this hypothesis, we performed electrical microstimulation at 461 highly restricted loci within the basal and central amygdala in awake mustached bats. At a subset of these sites, high frequency stimulation with weak constant current pulses presented at near-threshold levels triggered vocalization of either echolocation pulses or social calls. At the vast majority of locations, microstimulation produced a constellation of changes in autonomic and somatomotor outputs. These changes included widespread co-activation of significant tachycardia and hyperventilation and/or rhythmic ear pinna movements (PMs). In a few locations, responses were constrained to vocalization and/or PMs despite increases in the intensity of stimulation. The probability of eliciting echolocation pulses vs. social calls decreased in a medial-posterior to anterolateral direction within the centrobasal amygdala. Microinjections of kainic acid (KA) at stimulation sites confirmed the contribution of cellular activity rather than fibers-of-passage in the control of multimodal outputs. The results suggest that localized clusters of neurons may simultaneously modulate the activity of multiple central pattern generators (CPGs) present within the brainstem.

Passive Acoustic Monitoring the Diel, Lunar, Seasonal and Tidal Patterns in the Biosonar Activity of the Indo-Pacific Humpback Dolphins (Sousa chinensis) in the Pearl River Estuary, China

PubMed Central

Wang, Zhi-Tao; Nachtigall, Paul E.; Akamatsu, Tomonari; Wang, Ke-Xiong; Wu, Yu-Ping; Liu, Jian-Chang; Duan, Guo-Qin; Cao, Han-Jiang; Wang, Ding

2015-01-01

A growing demand for sustainable energy has led to an increase in construction of offshore windfarms. Guishan windmill farm will be constructed in the Pearl River Estuary, China, which sustains the world’s largest known population of Indo-Pacific humpback dolphins (Sousa chinensis). Dolphin conservation is an urgent issue in this region. By using passive acoustic monitoring, a baseline distribution of data on this species in the Pearl River Estuary during pre-construction period had been collected. Dolphin biosonar detection and its diel, lunar, seasonal and tidal patterns were examined using a Generalized Linear Model. Significant higher echolocation detections at night than during the day, in winter-spring than in summer-autumn, at high tide than at flood tide were recognized. Significant higher echolocation detections during the new moon were recognized at night time. The diel, lunar and seasonal patterns for the echolocation encounter duration also significantly varied. These patterns could be due to the spatial-temporal variability of dolphin prey and illumination conditions. The baseline information will be useful for driving further effective action on the conservation of this species and in facilitating later assessments of the effects of the offshore windfarm on the dolphins by comparing the baseline to post construction and post mitigation efforts. PMID:26580966

Bats adjust their mouth gape to zoom their biosonar field of view.

PubMed

Kounitsky, Pavel; Rydell, Jens; Amichai, Eran; Boonman, Arjan; Eitan, Ofri; Weiss, Anthony J; Yovel, Yossi

2015-05-26

Active sensing, where sensory acquisition is actively modulated, is an inherent component of almost all sensory systems. Echolocating bats are a prime example of active sensing. They can rapidly adjust many of their biosonar parameters to optimize sensory acquisition. They dynamically adjust pulse design, pulse duration, and pulse rate within dozens of milliseconds according to the sensory information that is required for the task that they are performing. The least studied and least understood degree of freedom in echolocation is emission beamforming--the ability to change the shape of the sonar sound beam in a functional way. Such an ability could have a great impact on the bat's control over its sensory perception. On the one hand, the bat could direct more energy into a narrow sector to zoom its biosonar field of view, and on the other hand, it could widen the beam to increase the space that it senses. We show that freely behaving bats constantly control their biosonar field of view in natural situations by rapidly adjusting their emitter aperture--the mouth gape. The bats dramatically narrowed the beam when entering a confined space, and they dramatically widened it within dozens of milliseconds when flying toward open space. Hence, mouth-emitting bats dynamically adjust their mouth gape to optimize the area that they sense with their echolocation system.

Object localization using a biosonar beam: how opening your mouth improves localization.

PubMed

Arditi, G; Weiss, A J; Yovel, Y

2015-08-01

Determining the location of a sound source is crucial for survival. Both predators and prey usually produce sound while moving, revealing valuable information about their presence and location. Animals have thus evolved morphological and neural adaptations allowing precise sound localization. Mammals rely on the temporal and amplitude differences between the sound signals arriving at their two ears, as well as on the spectral cues available in the signal arriving at a single ear to localize a sound source. Most mammals rely on passive hearing and are thus limited by the acoustic characteristics of the emitted sound. Echolocating bats emit sound to perceive their environment. They can, therefore, affect the frequency spectrum of the echoes they must localize. The biosonar sound beam of a bat is directional, spreading different frequencies into different directions. Here, we analyse mathematically the spatial information that is provided by the beam and could be used to improve sound localization. We hypothesize how bats could improve sound localization by altering their echolocation signal design or by increasing their mouth gape (the size of the sound emitter) as they, indeed, do in nature. Finally, we also reveal a trade-off according to which increasing the echolocation signal's frequency improves the accuracy of sound localization but might result in undesired large localization errors under low signal-to-noise ratio conditions.

Object localization using a biosonar beam: how opening your mouth improves localization

PubMed Central

Arditi, G.; Weiss, A. J.; Yovel, Y.

2015-01-01

Determining the location of a sound source is crucial for survival. Both predators and prey usually produce sound while moving, revealing valuable information about their presence and location. Animals have thus evolved morphological and neural adaptations allowing precise sound localization. Mammals rely on the temporal and amplitude differences between the sound signals arriving at their two ears, as well as on the spectral cues available in the signal arriving at a single ear to localize a sound source. Most mammals rely on passive hearing and are thus limited by the acoustic characteristics of the emitted sound. Echolocating bats emit sound to perceive their environment. They can, therefore, affect the frequency spectrum of the echoes they must localize. The biosonar sound beam of a bat is directional, spreading different frequencies into different directions. Here, we analyse mathematically the spatial information that is provided by the beam and could be used to improve sound localization. We hypothesize how bats could improve sound localization by altering their echolocation signal design or by increasing their mouth gape (the size of the sound emitter) as they, indeed, do in nature. Finally, we also reveal a trade-off according to which increasing the echolocation signal's frequency improves the accuracy of sound localization but might result in undesired large localization errors under low signal-to-noise ratio conditions. PMID:26361552

Bats adjust their mouth gape to zoom their biosonar field of view

PubMed Central

Rydell, Jens; Amichai, Eran; Boonman, Arjan; Eitan, Ofri; Weiss, Anthony J.; Yovel, Yossi

2015-01-01

Active sensing, where sensory acquisition is actively modulated, is an inherent component of almost all sensory systems. Echolocating bats are a prime example of active sensing. They can rapidly adjust many of their biosonar parameters to optimize sensory acquisition. They dynamically adjust pulse design, pulse duration, and pulse rate within dozens of milliseconds according to the sensory information that is required for the task that they are performing. The least studied and least understood degree of freedom in echolocation is emission beamforming—the ability to change the shape of the sonar sound beam in a functional way. Such an ability could have a great impact on the bat’s control over its sensory perception. On the one hand, the bat could direct more energy into a narrow sector to zoom its biosonar field of view, and on the other hand, it could widen the beam to increase the space that it senses. We show that freely behaving bats constantly control their biosonar field of view in natural situations by rapidly adjusting their emitter aperture—the mouth gape. The bats dramatically narrowed the beam when entering a confined space, and they dramatically widened it within dozens of milliseconds when flying toward open space. Hence, mouth-emitting bats dynamically adjust their mouth gape to optimize the area that they sense with their echolocation system. PMID:25941395

Matched Behavioral and Neural Adaptations for Low Sound Level Echolocation in a Gleaning Bat, Antrozous pallidus

PubMed Central

Measor, Kevin R.; Leavell, Brian C.; Brewton, Dustin H.; Rumschlag, Jeffrey; Barber, Jesse R.

2017-01-01

Abstract In active sensing, animals make motor adjustments to match sensory inputs to specialized neural circuitry. Here, we describe an active sensing system for sound level processing. The pallid bat uses downward frequency-modulated (FM) sweeps as echolocation calls for general orientation and obstacle avoidance. The bat’s auditory cortex contains a region selective for these FM sweeps (FM sweep-selective region, FMSR). We show that the vast majority of FMSR neurons are sensitive and strongly selective for relatively low levels (30-60 dB SPL). Behavioral testing shows that when a flying bat approaches a target, it reduces output call levels to keep echo levels between ∼30 and 55 dB SPL. Thus, the pallid bat behaviorally matches echo levels to an optimized neural representation of sound levels. FMSR neurons are more selective for sound levels of FM sweeps than tones, suggesting that across-frequency integration enhances level tuning. Level-dependent timing of high-frequency sideband inhibition in the receptive field shapes increased level selectivity for FM sweeps. Together with previous studies, these data indicate that the same receptive field properties shape multiple filters (sweep direction, rate, and level) for FM sweeps, a sound common in multiple vocalizations, including human speech. The matched behavioral and neural adaptations for low-intensity echolocation in the pallid bat will facilitate foraging with reduced probability of acoustic detection by prey. PMID:28275715

Description and clustering of echolocation signals of Commerson's dolphins (Cephalorhynchus commersonii) in Bahía San Julián, Argentina.

PubMed

Reyes Reyes, M Vanesa; Iñíguez, Miguel A; Hevia, Marta; Hildebrand, John A; Melcón, Mariana L

2015-10-01

Commerson's dolphins (Cephalorhynchus commersonii) inhabit coastal waters of Southern South America and Kerguelen Islands. Limited information exists about the acoustic repertoire of this species in the wild. Here, echolocation signals from free-ranging Commerson's dolphins were recorded in Bahía San Julián, Argentina. Signal parameters were calculated and a cluster analysis was made on 3180 regular clicks. Three clusters were obtained based on peak frequency (129, 137, and 173 kHz) and 3 dB bandwidth (8, 6, and 5 kHz). The 428 buzz clicks were analyzed separately. They consisted of clicks emitted with a median inter-click interval of 3.5 ms, peak frequency at 131 kHz, 3 dB bandwidth of 9 kHz, 10 dB bandwidth of 18 kHz, and duration of 56 μs. Buzz clicks were significantly shorter and with a lower peak frequency and a broader bandwidth than most of the regular clicks. This study provided the first description of different echolocation signals, including on- and off-axis signals, recorded from Commerson's dolphins in the wild, most likely as a result of animals at several distances and orientations to the recording device. This information could be useful while doing passive acoustic monitoring.

Echolocation signals of free-ranging Indo-Pacific humpback dolphins (Sousa chinensis) in Sanniang Bay, China.

PubMed

Fang, Liang; Li, Songhai; Wang, Kexiong; Wang, Zhitao; Shi, Wenjing; Wang, Ding

2015-09-01

While the low-frequency communication sounds of Indo-Pacific humpback dolphins (Sousa chinensis) have been reported in a number of papers, the high-frequency echolocation signals of Sousa chinensis, especially those living in the wild, have been less studied. In the current study, echolocation signals of humpback dolphins were recorded in Sanniang Bay, Guangxi Province, China, using a cross-type hydrophone array with five elements. In total, 77 candidate on-axis clicks from 77 scans were selected for analysis. The results showed that the varied peak-to-peak source levels ranged from 177.1 to 207.3 dB, with an average of 187.7 dB re: 1 μPa. The mean peak frequency was 109.0 kHz with a -3-dB bandwidth of 50.3 kHz and 95% energy duration of 22 μs. The -3-dB bandwidth was much broader than the root mean square bandwidth and exhibited a bimodal distribution. The center frequency exhibited a positive relationship with the peak-to-peak source level. The clicks of the wild Indo-Pacific humpback dolphins were short-duration, broadband, ultrasonic pulses, similar to those produced by other whistling dolphins of similar body size. However, the click source levels of the Indo-Pacific humpback dolphin appear to be lower than those of other whistling dolphins.

The detection of 'virtual' objects using echoes by humans: Spectral cues.

PubMed

Rowan, Daniel; Papadopoulos, Timos; Archer, Lauren; Goodhew, Amanda; Cozens, Hayley; Lopez, Ricardo Guzman; Edwards, David; Holmes, Hannah; Allen, Robert

2017-07-01

Some blind people use echoes to detect discrete, silent objects to support their spatial orientation/navigation, independence, safety and wellbeing. The acoustical features that people use for this are not well understood. Listening to changes in spectral shape due to the presence of an object could be important for object detection and avoidance, especially at short range, although it is currently not known whether it is possible with echolocation-related sounds. Bands of noise were convolved with recordings of binaural impulse responses of objects in an anechoic chamber to create 'virtual objects', which were analysed and played to sighted and blind listeners inexperienced in echolocation. The sounds were also manipulated to remove cues unrelated to spectral shape. Most listeners could accurately detect hard flat objects using changes in spectral shape. The useful spectral changes for object detection occurred above approximately 3 kHz, as with object localisation. However, energy in the sounds below 3 kHz was required to exploit changes in spectral shape for object detection, whereas energy below 3 kHz impaired object localisation. Further recordings showed that the spectral changes were diminished by room reverberation. While good high-frequency hearing is generally important for echolocation, the optimal echo-generating stimulus will probably depend on the task. Copyright © 2017 The Authors. Published by Elsevier B.V. All rights reserved.

Sound localization by echolocating bats

NASA Astrophysics Data System (ADS)

Aytekin, Murat

Echolocating bats emit ultrasonic vocalizations and listen to echoes reflected back from objects in the path of the sound beam to build a spatial representation of their surroundings. Important to understanding the representation of space through echolocation are detailed studies of the cues used for localization, the sonar emission patterns and how this information is assembled. This thesis includes three studies, one on the directional properties of the sonar receiver, one on the directional properties of the sonar transmitter, and a model that demonstrates the role of action in building a representation of auditory space. The general importance of this work to a broader understanding of spatial localization is discussed. Investigations of the directional properties of the sonar receiver reveal that interaural level difference and monaural spectral notch cues are both dependent on sound source azimuth and elevation. This redundancy allows flexibility that an echolocating bat may need when coping with complex computational demands for sound localization. Using a novel method to measure bat sonar emission patterns from freely behaving bats, I show that the sonar beam shape varies between vocalizations. Consequently, the auditory system of a bat may need to adapt its computations to accurately localize objects using changing acoustic inputs. Extra-auditory signals that carry information about pinna position and beam shape are required for auditory localization of sound sources. The auditory system must learn associations between extra-auditory signals and acoustic spatial cues. Furthermore, the auditory system must adapt to changes in acoustic input that occur with changes in pinna position and vocalization parameters. These demands on the nervous system suggest that sound localization is achieved through the interaction of behavioral control and acoustic inputs. A sensorimotor model demonstrates how an organism can learn space through auditory-motor contingencies. The model also reveals how different aspects of sound localization, such as experience-dependent acquisition, adaptation, and extra-auditory influences, can be brought together under a comprehensive framework. This thesis presents a foundation for understanding the representation of auditory space that builds upon acoustic cues, motor control, and learning dynamic associations between action and auditory inputs.

Enhanced echolocation via robust statistics and super-resolution of sonar images

NASA Astrophysics Data System (ADS)

Kim, Kio

Echolocation is a process in which an animal uses acoustic signals to exchange information with environments. In a recent study, Neretti et al. have shown that the use of robust statistics can significantly improve the resiliency of echolocation against noise and enhance its accuracy by suppressing the development of sidelobes in the processing of an echo signal. In this research, the use of robust statistics is extended to problems in underwater explorations. The dissertation consists of two parts. Part I describes how robust statistics can enhance the identification of target objects, which in this case are cylindrical containers filled with four different liquids. Particularly, this work employs a variation of an existing robust estimator called an L-estimator, which was first suggested by Koenker and Bassett. As pointed out by Au et al.; a 'highlight interval' is an important feature, and it is closely related with many other important features that are known to be crucial for dolphin echolocation. A varied L-estimator described in this text is used to enhance the detection of highlight intervals, which eventually leads to a successful classification of echo signals. Part II extends the problem into 2 dimensions. Thanks to the advances in material and computer technology, various sonar imaging modalities are available on the market. By registering acoustic images from such video sequences, one can extract more information on the region of interest. Computer vision and image processing allowed application of robust statistics to the acoustic images produced by forward looking sonar systems, such as Dual-frequency Identification Sonar and ProViewer. The first use of robust statistics for sonar image enhancement in this text is in image registration. Random Sampling Consensus (RANSAC) is widely used for image registration. The registration algorithm using RANSAC is optimized for sonar image registration, and the performance is studied. The second use of robust statistics is in fusing the images. It is shown that the maximum a posteriori fusion method can be formulated in a Kalman filter-like manner, and also that the resulting expression is identical to a W-estimator with a specific weight function.

Place recognition using batlike sonar.

PubMed

Vanderelst, Dieter; Steckel, Jan; Boen, Andre; Peremans, Herbert; Holderied, Marc W

2016-08-02

Echolocating bats have excellent spatial memory and are able to navigate to salient locations using bio-sonar. Navigating and route-following require animals to recognize places. Currently, it is mostly unknown how bats recognize places using echolocation. In this paper, we propose template based place recognition might underlie sonar-based navigation in bats. Under this hypothesis, bats recognize places by remembering their echo signature - rather than their 3D layout. Using a large body of ensonification data collected in three different habitats, we test the viability of this hypothesis assessing two critical properties of the proposed echo signatures: (1) they can be uniquely classified and (2) they vary continuously across space. Based on the results presented, we conclude that the proposed echo signatures satisfy both criteria. We discuss how these two properties of the echo signatures can support navigation and building a cognitive map.

A Device for Human Ultrasonic Echolocation.

PubMed

Sohl-Dickstein, Jascha; Teng, Santani; Gaub, Benjamin M; Rodgers, Chris C; Li, Crystal; DeWeese, Michael R; Harper, Nicol S

2015-06-01

We present a device that combines principles of ultrasonic echolocation and spatial hearing to provide human users with environmental cues that are 1) not otherwise available to the human auditory system, and 2) richer in object and spatial information than the more heavily processed sonar cues of other assistive devices. The device consists of a wearable headset with an ultrasonic emitter and stereo microphones with affixed artificial pinnae. The goal of this study is to describe the device and evaluate the utility of the echoic information it provides. The echoes of ultrasonic pulses were recorded and time stretched to lower their frequencies into the human auditory range, then played back to the user. We tested performance among naive and experienced sighted volunteers using a set of localization experiments, in which the locations of echo-reflective surfaces were judged using these time-stretched echoes. Naive subjects were able to make laterality and distance judgments, suggesting that the echoes provide innately useful information without prior training. Naive subjects were generally unable to make elevation judgments from recorded echoes. However, trained subjects demonstrated an ability to judge elevation as well. This suggests that the device can be used effectively to examine the environment and that the human auditory system can rapidly adapt to these artificial echolocation cues. Interpreting and interacting with the external world constitutes a major challenge for persons who are blind or visually impaired. This device has the potential to aid blind people in interacting with their environment.

Digital cranial endocast of Hyopsodus (Mammalia, "Condylarthra"): a case of paleogene terrestrial echolocation?

PubMed

Orliac, Maeva J; Argot, Christine; Gilissen, Emmanuel

2012-01-01

We here describe the endocranial cast of the Eocene archaic ungulate Hyopsodus lepidus AMNH 143783 (Bridgerian, North America) reconstructed from X-ray computed microtomography data. This represents the first complete cranial endocast known for Hyopsodontinae. The Hyopsodus endocast is compared to other known "condylarthran" endocasts, i. e. those of Pleuraspidotherium (Pleuraspidotheriidae), Arctocyon (Arctocyonidae), Meniscotherium (Meniscotheriidae), Phenacodus (Phenacodontidae), as well as to basal perissodactyls (Hyracotherium) and artiodactyls (Cebochoerus, Homacodon). Hyopsodus presents one of the highest encephalization quotients of archaic ungulates and shows an "advanced version" of the basal ungulate brain pattern, with a mosaic of archaic characters such as large olfactory bulbs, weak ventral expansion of the neopallium, and absence of neopallium fissuration, as well as more specialized ones such as the relative reduction of the cerebellum compared to cerebrum or the enlargement of the inferior colliculus. As in other archaic ungulates, Hyopsodus midbrain exposure is important, but it exhibits a dorsally protruding largely developed inferior colliculus, a feature unique among "Condylarthra". A potential correlation between the development of the inferior colliculus in Hyopsodus and the use of terrestrial echolocation as observed in extant tenrecs and shrews is discussed. The detailed analysis of the overall morphology of the postcranial skeleton of Hyopsodus indicates a nimble, fast moving animal that likely lived in burrows. This would be compatible with terrestrial echolocation used by the animal to investigate subterranean habitat and/or to minimize predation during nocturnal exploration of the environment.

A device for human ultrasonic echolocation

PubMed Central

Gaub, Benjamin M.; Rodgers, Chris C.; Li, Crystal; DeWeese, Michael R.; Harper, Nicol S.

2015-01-01

Objective We present a device that combines principles of ultrasonic echolocation and spatial hearing to provide human users with environmental cues that are 1) not otherwise available to the human auditory system and 2) richer in object, and spatial information than the more heavily processed sonar cues of other assistive devices. The device consists of a wearable headset with an ultrasonic emitter and stereo microphones with affixed artificial pinnae. The goal of this study is to describe the device and evaluate the utility of the echoic information it provides. Methods The echoes of ultrasonic pulses were recorded and time-stretched to lower their frequencies into the human auditory range, then played back to the user. We tested performance among naive and experienced sighted volunteers using a set of localization experiments in which the locations of echo-reflective surfaces were judged using these time stretched echoes. Results Naive subjects were able to make laterality and distance judgments, suggesting that the echoes provide innately useful information without prior training. Naive subjects were generally unable to make elevation judgments from recorded echoes. However trained subjects demonstrated an ability to judge elevation as well. Conclusion This suggests that the device can be used effectively to examine the environment and that the human auditory system can rapidly adapt to these artificial echolocation cues. Significance Interpreting and interacting with the external world constitutes a major challenge for persons who are blind or visually impaired. This device has the potential to aid blind people in interacting with their environment. PMID:25608301

Vespertilionid bats control the width of their biosonar sound beam dynamically during prey pursuit

PubMed Central

Jakobsen, Lasse; Surlykke, Annemarie

2010-01-01

Animals using sound for communication emit directional signals, focusing most acoustic energy in one direction. Echolocating bats are listening for soft echoes from insects. Therefore, a directional biosonar sound beam greatly increases detection probability in the forward direction and decreases off-axis echoes. However, high directionality has context-specific disadvantages: at close range the detection space will be vastly reduced, making a broad beam favorable. Hence, a flexible system would be very advantageous. We investigated whether bats can dynamically change directionality of their biosonar during aerial pursuit of insects. We trained five Myotis daubentonii and one Eptesicus serotinus to capture tethered mealworms and recorded their echolocation signals with a multimicrophone array. The results show that the bats broaden the echolocation beam drastically in the terminal phase of prey pursuit. M. daubentonii increased the half-amplitude angle from approximately 40° to approximately 90° horizontally and from approximately 45° to more than 90° vertically. The increase in beam width is achieved by lowering the frequency by roughly one octave from approximately 55 kHz to approximately 27.5 kHz. The E. serotinus showed beam broadening remarkably similar to that of M. daubentonii. Our results demonstrate dynamic control of beam width in both species. Hence, we propose directionality as an explanation for the frequency decrease observed in the buzz of aerial hawking vespertilionid bats. We predict that future studies will reveal dynamic control of beam width in a broad range of acoustically communicating animals. PMID:20643943

Active control of acoustic field-of-view in a biosonar system.

PubMed

Yovel, Yossi; Falk, Ben; Moss, Cynthia F; Ulanovsky, Nachum

2011-09-01

Active-sensing systems abound in nature, but little is known about systematic strategies that are used by these systems to scan the environment. Here, we addressed this question by studying echolocating bats, animals that have the ability to point their biosonar beam to a confined region of space. We trained Egyptian fruit bats to land on a target, under conditions of varying levels of environmental complexity, and measured their echolocation and flight behavior. The bats modulated the intensity of their biosonar emissions, and the spatial region they sampled, in a task-dependant manner. We report here that Egyptian fruit bats selectively change the emission intensity and the angle between the beam axes of sequentially emitted clicks, according to the distance to the target, and depending on the level of environmental complexity. In so doing, they effectively adjusted the spatial sector sampled by a pair of clicks-the "field-of-view." We suggest that the exact point within the beam that is directed towards an object (e.g., the beam's peak, maximal slope, etc.) is influenced by three competing task demands: detection, localization, and angular scanning-where the third factor is modulated by field-of-view. Our results suggest that lingual echolocation (based on tongue clicks) is in fact much more sophisticated than previously believed. They also reveal a new parameter under active control in animal sonar-the angle between consecutive beams. Our findings suggest that acoustic scanning of space by mammals is highly flexible and modulated much more selectively than previously recognized.

Vespertilionid bats control the width of their biosonar sound beam dynamically during prey pursuit.

PubMed

Jakobsen, Lasse; Surlykke, Annemarie

2010-08-03

Animals using sound for communication emit directional signals, focusing most acoustic energy in one direction. Echolocating bats are listening for soft echoes from insects. Therefore, a directional biosonar sound beam greatly increases detection probability in the forward direction and decreases off-axis echoes. However, high directionality has context-specific disadvantages: at close range the detection space will be vastly reduced, making a broad beam favorable. Hence, a flexible system would be very advantageous. We investigated whether bats can dynamically change directionality of their biosonar during aerial pursuit of insects. We trained five Myotis daubentonii and one Eptesicus serotinus to capture tethered mealworms and recorded their echolocation signals with a multimicrophone array. The results show that the bats broaden the echolocation beam drastically in the terminal phase of prey pursuit. M. daubentonii increased the half-amplitude angle from approximately 40 degrees to approximately 90 degrees horizontally and from approximately 45 degrees to more than 90 degrees vertically. The increase in beam width is achieved by lowering the frequency by roughly one octave from approximately 55 kHz to approximately 27.5 kHz. The E. serotinus showed beam broadening remarkably similar to that of M. daubentonii. Our results demonstrate dynamic control of beam width in both species. Hence, we propose directionality as an explanation for the frequency decrease observed in the buzz of aerial hawking vespertilionid bats. We predict that future studies will reveal dynamic control of beam width in a broad range of acoustically communicating animals.

The influence of flight speed on the ranging performance of bats using frequency modulated echolocation pulses

NASA Astrophysics Data System (ADS)

Boonman, Arjan M.; Parsons, Stuart; Jones, Gareth

2003-01-01

Many species of bat use ultrasonic frequency modulated (FM) pulses to measure the distance to objects by timing the emission and reception of each pulse. Echolocation is mainly used in flight. Since the flight speed of bats often exceeds 1% of the speed of sound, Doppler effects will lead to compression of the time between emission and reception as well as an elevation of the echo frequencies, resulting in a distortion of the perceived range. This paper describes the consequences of these Doppler effects on the ranging performance of bats using different pulse designs. The consequences of Doppler effects on ranging performance described in this paper assume bats to have a very accurate ranging resolution, which is feasible with a filterbank receiver. By modeling two receiver types, it was first established that the effects of Doppler compression are virtually independent of the receiver type. Then, used a cross-correlation model was used to investigate the effect of flight speed on Doppler tolerance and range-Doppler coupling separately. This paper further shows how pulse duration, bandwidth, function type, and harmonics influence Doppler tolerance and range-Doppler coupling. The influence of each signal parameter is illustrated using calls of several bat species. It is argued that range-Doppler coupling is a significant source of error in bat echolocation, and various strategies bats could employ to deal with this problem, including the use of range rate information are discussed.

Dynamics of hippocampal spatial representation in echolocating bats

PubMed Central

Ulanovsky, Nachum; Moss, Cynthia F.

2009-01-01

The ‘place fields‘ of hippocampal pyramidal neurons are not static. For example, upon a contextual change in the environment, place-fields may ‘remap‘ within typical timescales of ~1 minute. A few studies have shown more rapid dynamics in hippocampal activity, linked to internal processes, such as switches between spatial reference frames or changes within the theta cycle. However, little is known about rapid hippocampal place-field dynamics in response to external, sensory stimuli. Here, we studied this question in big brown bats, echolocating mammals in which we can readily measure rapid changes in sensory dynamics (sonar signals), as well as rapid behavioral switches between distal and proximal exploratory modes. First, we show that place-field size was modulated by the availability of sensory information, on a timescale of ~300-milliseconds: Bat hippocampal place-fields were smallest immediately after an echolocation call, but place-fields ‘diffused’ with the passage of time after the call, when echo information was no longer arriving. Second, we show rapid modulation of hippocampal place-fields as the animal switched between two exploratory modes. Third, we compared place fields and spatial-view fields of individual neurons and found that place tuning was much more pronounced than spatial-view tuning. In addition, dynamic fluctuations in spatial-view tuning were stronger than fluctuations in place tuning. Taken together, these results suggest that spatial representation in mammalian hippocampus can be very rapidly modulated by external sensory and behavioral events. PMID:20014379

Ambient noise induces independent shifts in call frequency and amplitude within the Lombard effect in echolocating bats

PubMed Central

Hage, Steffen R.; Jiang, Tinglei; Berquist, Sean W.; Feng, Jiang; Metzner, Walter

2013-01-01

The Lombard effect, an involuntary rise in call amplitude in response to masking ambient noise, represents one of the most efficient mechanisms to optimize signal-to-noise ratio. The Lombard effect occurs in birds and mammals, including humans, and is often associated with several other vocal changes, such as call frequency and duration. Most studies, however, have focused on noise-dependent changes in call amplitude. It is therefore still largely unknown how the adaptive changes in call amplitude relate to associated vocal changes such as frequency shifts, how the underlying mechanisms are linked, and if auditory feedback from the changing vocal output is needed. Here, we examined the Lombard effect and the associated changes in call frequency in a highly vocal mammal, echolocating horseshoe bats. We analyzed how bandpass-filtered noise (BFN; bandwidth 20 kHz) affected their echolocation behavior when BFN was centered on different frequencies within their hearing range. Call amplitudes increased only when BFN was centered on the dominant frequency component of the bats’ calls. In contrast, call frequencies increased for all but one BFN center frequency tested. Both amplitude and frequency rises were extremely fast and occurred in the first call uttered after noise onset, suggesting that no auditory feedback was required. The different effects that varying the BFN center frequency had on amplitude and frequency rises indicate different neural circuits and/or mechanisms underlying these changes. PMID:23431172

Digital Cranial Endocast of Hyopsodus (Mammalia, “Condylarthra”): A Case of Paleogene Terrestrial Echolocation?

PubMed Central

Orliac, Maeva J.; Argot, Christine; Gilissen, Emmanuel

2012-01-01

We here describe the endocranial cast of the Eocene archaic ungulate Hyopsodus lepidus AMNH 143783 (Bridgerian, North America) reconstructed from X-ray computed microtomography data. This represents the first complete cranial endocast known for Hyopsodontinae. The Hyopsodus endocast is compared to other known “condylarthran” endocasts, i. e. those of Pleuraspidotherium (Pleuraspidotheriidae), Arctocyon (Arctocyonidae), Meniscotherium (Meniscotheriidae), Phenacodus (Phenacodontidae), as well as to basal perissodactyls (Hyracotherium) and artiodactyls (Cebochoerus, Homacodon). Hyopsodus presents one of the highest encephalization quotients of archaic ungulates and shows an “advanced version” of the basal ungulate brain pattern, with a mosaic of archaic characters such as large olfactory bulbs, weak ventral expansion of the neopallium, and absence of neopallium fissuration, as well as more specialized ones such as the relative reduction of the cerebellum compared to cerebrum or the enlargement of the inferior colliculus. As in other archaic ungulates, Hyopsodus midbrain exposure is important, but it exhibits a dorsally protruding largely developed inferior colliculus, a feature unique among “Condylarthra”. A potential correlation between the development of the inferior colliculus in Hyopsodus and the use of terrestrial echolocation as observed in extant tenrecs and shrews is discussed. The detailed analysis of the overall morphology of the postcranial skeleton of Hyopsodus indicates a nimble, fast moving animal that likely lived in burrows. This would be compatible with terrestrial echolocation used by the animal to investigate subterranean habitat and/or to minimize predation during nocturnal exploration of the environment. PMID:22347998

Echolocation by the harbour porpoise: life in coastal waters.

PubMed

Miller, Lee A; Wahlberg, Magnus

2013-01-01

The harbor porpoise is one of the smallest and most widely spread of all toothed whales. They are found abundantly in coastal waters all around the northern hemisphere. They are among the 11 species known to use high frequency sonar of relative narrow bandwidth. Their narrow biosonar beam helps isolate echoes from prey among those from unwanted items and noise. Obtaining echoes from small objects like net mesh, net floats, and small prey is facilitated by the very high peak frequency around 130 kHz with a wavelength of about 12 mm. We argue that such echolocation signals and narrow band auditory filters give the harbor porpoise a selective advantage in a coastal environment. Predation by killer whales and a minimum noise region in the ocean around 130 kHz may have provided selection pressures for using narrow bandwidth high frequency biosonar signals.

Echolocation by the harbour porpoise: life in coastal waters

PubMed Central

Miller, Lee A.; Wahlberg, Magnus

2013-01-01

The harbor porpoise is one of the smallest and most widely spread of all toothed whales. They are found abundantly in coastal waters all around the northern hemisphere. They are among the 11 species known to use high frequency sonar of relative narrow bandwidth. Their narrow biosonar beam helps isolate echoes from prey among those from unwanted items and noise. Obtaining echoes from small objects like net mesh, net floats, and small prey is facilitated by the very high peak frequency around 130 kHz with a wavelength of about 12 mm. We argue that such echolocation signals and narrow band auditory filters give the harbor porpoise a selective advantage in a coastal environment. Predation by killer whales and a minimum noise region in the ocean around 130 kHz may have provided selection pressures for using narrow bandwidth high frequency biosonar signals. PMID:23596420

Place recognition using batlike sonar

PubMed Central

Vanderelst, Dieter; Steckel, Jan; Boen, Andre; Peremans, Herbert; Holderied, Marc W

2016-01-01

Echolocating bats have excellent spatial memory and are able to navigate to salient locations using bio-sonar. Navigating and route-following require animals to recognize places. Currently, it is mostly unknown how bats recognize places using echolocation. In this paper, we propose template based place recognition might underlie sonar-based navigation in bats. Under this hypothesis, bats recognize places by remembering their echo signature - rather than their 3D layout. Using a large body of ensonification data collected in three different habitats, we test the viability of this hypothesis assessing two critical properties of the proposed echo signatures: (1) they can be uniquely classified and (2) they vary continuously across space. Based on the results presented, we conclude that the proposed echo signatures satisfy both criteria. We discuss how these two properties of the echo signatures can support navigation and building a cognitive map. DOI: http://dx.doi.org/10.7554/eLife.14188.001 PMID:27481189

Fine-tuned echolocation and capture-flight of Myotis capaccinii when facing different-sized insect and fish prey.

PubMed

Aizpurua, Ostaizka; Aihartza, Joxerra; Alberdi, Antton; Baagøe, Hans J; Garin, Inazio

2014-09-15

Formerly thought to be a strictly insectivorous trawling bat, recent studies have shown that Myotis capaccinii also preys on fish. To determine whether differences exist in bat flight behaviour, prey handling and echolocation characteristics when catching fish and insects of different size, we conducted a field experiment focused on the last stage of prey capture. We used synchronized video and ultrasound recordings to measure several flight and dip features as well as echolocation characteristics, focusing on terminal buzz phase I, characterized by a call rate exceeding 100 Hz, and buzz phase II, characterized by a drop in the fundamental well below 20 kHz and a repetition rate exceeding 150 Hz. When capturing insects, bats used both parts of the terminal phase to the same extent, and performed short and superficial drags on the water surface. In contrast, when preying on fish, buzz I was longer and buzz II shorter, and the bats made longer and deeper dips. These variations suggest that lengthening buzz I and shortening buzz II when fishing is beneficial, probably because buzz I gives better discrimination ability and the broader sonar beam provided by buzz II is useless when no evasive flight of the prey is expected. Additionally, bats continued emitting calls beyond the theoretical signal-overlap zone, suggesting that they might obtain information even when they have surpassed that threshold, at least initially. This study shows that M. capaccinii can regulate the temporal components of its feeding buzzes and modify prey capture technique according to the target. © 2014. Published by The Company of Biologists Ltd.

Tongue-driven sonar beam steering by a lingual-echolocating fruit bat

PubMed Central

Falk, Benjamin; Chiu, Chen; Krishnan, Anand; Arbour, Jessica H.; Moss, Cynthia F.

2017-01-01

Animals enhance sensory acquisition from a specific direction by movements of head, ears, or eyes. As active sensing animals, echolocating bats also aim their directional sonar beam to selectively “illuminate” a confined volume of space, facilitating efficient information processing by reducing echo interference and clutter. Such sonar beam control is generally achieved by head movements or shape changes of the sound-emitting mouth or nose. However, lingual-echolocating Egyptian fruit bats, Rousettus aegyptiacus, which produce sound by clicking their tongue, can dramatically change beam direction at very short temporal intervals without visible morphological changes. The mechanism supporting this capability has remained a mystery. Here, we measured signals from free-flying Egyptian fruit bats and discovered a systematic angular sweep of beam focus across increasing frequency. This unusual signal structure has not been observed in other animals and cannot be explained by the conventional and widely-used “piston model” that describes the emission pattern of other bat species. Through modeling, we show that the observed beam features can be captured by an array of tongue-driven sound sources located along the side of the mouth, and that the sonar beam direction can be steered parsimoniously by inducing changes to the pattern of phase differences through moving tongue location. The effects are broadly similar to those found in a phased array—an engineering design widely found in human-made sonar systems that enables beam direction changes without changes in the physical transducer assembly. Our study reveals an intriguing parallel between biology and human engineering in solving problems in fundamentally similar ways. PMID:29244805

Tongue-driven sonar beam steering by a lingual-echolocating fruit bat.

PubMed

Lee, Wu-Jung; Falk, Benjamin; Chiu, Chen; Krishnan, Anand; Arbour, Jessica H; Moss, Cynthia F

2017-12-01

Animals enhance sensory acquisition from a specific direction by movements of head, ears, or eyes. As active sensing animals, echolocating bats also aim their directional sonar beam to selectively "illuminate" a confined volume of space, facilitating efficient information processing by reducing echo interference and clutter. Such sonar beam control is generally achieved by head movements or shape changes of the sound-emitting mouth or nose. However, lingual-echolocating Egyptian fruit bats, Rousettus aegyptiacus, which produce sound by clicking their tongue, can dramatically change beam direction at very short temporal intervals without visible morphological changes. The mechanism supporting this capability has remained a mystery. Here, we measured signals from free-flying Egyptian fruit bats and discovered a systematic angular sweep of beam focus across increasing frequency. This unusual signal structure has not been observed in other animals and cannot be explained by the conventional and widely-used "piston model" that describes the emission pattern of other bat species. Through modeling, we show that the observed beam features can be captured by an array of tongue-driven sound sources located along the side of the mouth, and that the sonar beam direction can be steered parsimoniously by inducing changes to the pattern of phase differences through moving tongue location. The effects are broadly similar to those found in a phased array-an engineering design widely found in human-made sonar systems that enables beam direction changes without changes in the physical transducer assembly. Our study reveals an intriguing parallel between biology and human engineering in solving problems in fundamentally similar ways.

Bats in Agroecosytems around California's Central Coast

NASA Astrophysics Data System (ADS)

Wayne, A.

2014-12-01

Bats in agroecosystems around California's Central Coast: A full quarter of California's land area is farmland. Crops account for 32.5 billion of California's GDP. Insect control is a big problem for farmers, and California bats eat only insects, saving farmers an estimated 3 to $53 billion a year. As farmers maximize crop yield, they use more pesticides, herbicides, and fertilizers, which contaminate runoff streams that bats drink from. Also, pesticide use kills bats' sole food source: insects. My research objective was to find out how farm management practices and landscape complexity affect bat diversity and activity, and to see which one affects bat activity more. We monitored 18 sites, including conventional, organic, and low and high-complexity landscapes. We noted more bat activity at sites with high complexity landscapes and organic practices than at sites with either low-complexity landscapes or conventional farming practices. I captured and processed bats and recorded data. I also classified insects collected from light traps. I learned how to handle bats and measure forearm length and weight, as well as how to indentify their gender. I took hair clippings and fecal samples, which yield data about the bats' diet. Their diet, in turn, gives us data about which pests they eat and therefore help control. I also learned about bats' echolocation: they have a special muscle over their ears that closes when they echolocate so that they don't burst their own eardrum. Also, some insects have evolved a special call that will disrupt bats echolocation so bats can't track it.

Bat use of a high-plains urban wildlife refuge

USGS Publications Warehouse

Everette, A.L.; O'Shea, T.J.; Ellison, L.E.; Stone, L.A.; McCance, J.L.

2001-01-01

Bats are significant components of mammalian diversity and in many areas are of management concern. However, little attention has been given to bats in urban or prairie landscapes. In 1997 and 1998, we determined species richness, relative abundance, roosting habits, and echolocation activity of bats at Rocky Mountain Arsenal National Wildlife Refuge (RMA), the largest urban unit in the United States refuge system, located on the high plains near Denver, Colorado. An inventory using mist nets revealed 3 species foraging at this site: big brown bats (Eptesicus fuscus), hoary bats (Lasiurus cinereus), and silver-haired bats (Lasionycteris noctivagans). Big brown bats comprised 86% of captures (n=176). This pattern was consistent with continental-scale predictions of bat species richness and evenness based on availability of potential roosts. Relative abundance based on captures was similar to that revealed by echolocation detector surveys, except that the latter revealed the likely presence of at least 2 additional species (Myotis spp. and red bats [Lasiurus borealis]). Echolocation activity was significantly greater (P=0.009) in areas with tree or water habitat edges than in open prairie, suggesting that maintaining such features is important for bats. Big brown bats commuted greater distances (9.2-18.8 km) from roosts in urban core areas to foraging sites on the refuge than typically reported for this species elsewhere, emphasizing the value of the site to these bats. Urban refuges can provide habitat of importance to bat populations, but may be characterized by abundant bats that roost in buildings if a variety of other kinds of roosting habitats are unavailable.

Unique Turbinal Morphology in Horseshoe Bats (Chiroptera: Rhinolophidae).

PubMed

Curtis, Abigail A; Simmons, Nancy B

2017-02-01

The mammalian nasal fossa contains a set of delicate and often structurally complex bones called turbinals. Turbinals and associated mucosae function in regulating respiratory heat and water loss, increasing surface area for olfactory tissue, and directing airflow within the nasal fossa. We used high-resolution micro-CT scanning to investigate a unique maxilloturbinal morphology in 37 species from the bat family Rhinolophidae, which we compared with those of families Hipposideridae, Megadermatidae, and Pteropodidae. Rhinolophids exhibit numerous structural modifications along the nasopharyngeal tract associated with emission of high duty cycle echolocation calls via the nostrils. In rhinolophids, we found that the maxilloturbinals and a portion of ethmoturbinal I form a pair of strand-like bony structures on each side of the nasal chamber. These structures project anteriorly from the transverse lamina and complete a hairpin turn to project posteriorly down the nasopharyngeal duct, and vary in length among species. The strand-like maxilloturbinals in Rhinolophidae were not observed in our outgroups and represent a synapomorphy for this family, and are unique in form among mammals. Within Rhinolophidae, maxilloturbinal size and cross-sectional shape were correlated with phylogeny. We hypothesize that strand-shaped maxilloturbinals may function to reduce respiratory heat and water loss without greatly impacting echolocation call transmission since they provide increased mucosal surface area for heat and moisture exchange but occupy minimal space. Alternatively, they may play a role in transmission of echolocation calls since they are located directly along the path sound travels between the larynx and nostrils during call emission. Anat Rec, 300:309-325, 2017. © 2016 Wiley Periodicals, Inc. © 2016 Wiley Periodicals, Inc.

Evolutionary escalation: the bat-moth arms race.

PubMed

Ter Hofstede, Hannah M; Ratcliffe, John M

2016-06-01

Echolocation in bats and high-frequency hearing in their insect prey make bats and insects an ideal system for studying the sensory ecology and neuroethology of predator-prey interactions. Here, we review the evolutionary history of bats and eared insects, focusing on the insect order Lepidoptera, and consider the evidence for antipredator adaptations and predator counter-adaptations. Ears evolved in a remarkable number of body locations across insects, with the original selection pressure for ears differing between groups. Although cause and effect are difficult to determine, correlations between hearing and life history strategies in moths provide evidence for how these two variables influence each other. We consider life history variables such as size, sex, circadian and seasonal activity patterns, geographic range and the composition of sympatric bat communities. We also review hypotheses on the neural basis for anti-predator behaviours (such as evasive flight and sound production) in moths. It is assumed that these prey adaptations would select for counter-adaptations in predatory bats. We suggest two levels of support for classifying bat traits as counter-adaptations: traits that allow bats to eat more eared prey than expected based on their availability in the environment provide a low level of support for counter-adaptations, whereas traits that have no other plausible explanation for their origination and maintenance than capturing defended prey constitute a high level of support. Specific predator counter-adaptations include calling at frequencies outside the sensitivity range of most eared prey, changing the pattern and frequency of echolocation calls during prey pursuit, and quiet, or 'stealth', echolocation. © 2016. Published by The Company of Biologists Ltd.

Active Control of Acoustic Field-of-View in a Biosonar System

PubMed Central

Yovel, Yossi; Falk, Ben; Moss, Cynthia F.; Ulanovsky, Nachum

2011-01-01

Active-sensing systems abound in nature, but little is known about systematic strategies that are used by these systems to scan the environment. Here, we addressed this question by studying echolocating bats, animals that have the ability to point their biosonar beam to a confined region of space. We trained Egyptian fruit bats to land on a target, under conditions of varying levels of environmental complexity, and measured their echolocation and flight behavior. The bats modulated the intensity of their biosonar emissions, and the spatial region they sampled, in a task-dependant manner. We report here that Egyptian fruit bats selectively change the emission intensity and the angle between the beam axes of sequentially emitted clicks, according to the distance to the target, and depending on the level of environmental complexity. In so doing, they effectively adjusted the spatial sector sampled by a pair of clicks—the “field-of-view.” We suggest that the exact point within the beam that is directed towards an object (e.g., the beam's peak, maximal slope, etc.) is influenced by three competing task demands: detection, localization, and angular scanning—where the third factor is modulated by field-of-view. Our results suggest that lingual echolocation (based on tongue clicks) is in fact much more sophisticated than previously believed. They also reveal a new parameter under active control in animal sonar—the angle between consecutive beams. Our findings suggest that acoustic scanning of space by mammals is highly flexible and modulated much more selectively than previously recognized. PMID:21931535

Neurodynamics for auditory stream segregation: tracking sounds in the mustached bat's natural environment.

PubMed

Kanwal, Jagmeet S; Medvedev, Andrei V; Micheyl, Christophe

2003-08-01

During navigation and the search phase of foraging, mustached bats emit approximately 25 ms long echolocation pulses (at 10-40 Hz) that contain multiple harmonics of a constant frequency (CF) component followed by a short (3 ms) downward frequency modulation. In the context of auditory stream segregation, therefore, bats may either perceive a coherent pulse-echo sequence (PEPE...), or segregated pulse and echo streams (P-P-P... and E-E-E...). To identify the neural mechanisms for stream segregation in bats, we developed a simple yet realistic neural network model with seven layers and 420 nodes. Our model required recurrent and lateral inhibition to enable output nodes in the network to 'latch-on' to a single tone (corresponding to a CF component in either the pulse or echo), i.e., exhibit differential suppression by the alternating two tones presented at a high rate (> 10 Hz). To test the applicability of our model to echolocation, we obtained neurophysiological data from the primary auditory cortex of awake mustached bats. Event-related potentials reliably reproduced the latching behaviour observed at output nodes in the network. Pulse as well as nontarget (clutter) echo CFs facilitated this latching. Individual single unit responses were erratic, but when summed over several recording sites, they also exhibited reliable latching behaviour even at 40 Hz. On the basis of these findings, we propose that a neural correlate of auditory stream segregation is present within localized synaptic activity in the mustached bat's auditory cortex and this mechanism may enhance the perception of echolocation sounds in the natural environment.

Bats aggregate to improve prey search but might be impaired when their density becomes too high.

PubMed

Cvikel, Noam; Egert Berg, Katya; Levin, Eran; Hurme, Edward; Borissov, Ivailo; Boonman, Arjan; Amichai, Eran; Yovel, Yossi

2015-01-19

Social foraging is a very common yet extremely complex behavior. Numerous studies attempted to model it with little supporting evidence. Studying it in the wild is difficult because it requires monitoring the animal's movement, its foraging success, and its interactions with conspecifics. We present a novel system that enables full night ultrasonic recording of freely foraging bats, in addition to GPS tracking. As they rely on echolocation, audio recordings of bats allow tapping into their sensory acquisition of the world. Rapid changes in echolocation allowed us to reveal the bats' dynamic reactions in response to prey or conspecifics—two key behaviors that are extremely difficult to assess in most animals. We found that bats actively aggregate and forage as a group. However, we also found that when the group became too dense, bats were forced to devote sensory attention to conspecifics that frequently entered their biosonar "field of view," impairing the bats' prey detection performance. Why then did bats fly in such high densities? By emitting echolocation calls, bats constantly provide public information about their detection of prey. Bats could therefore benefit from intentionally flying at a distance that enables eavesdropping on conspecifics. Group foraging, therefore, probably allowed bats to effectively operate as an array of sensors, increasing their searching efficiency. We suggest that two opposing forces are at play in determining the efficient foraging density: on the one hand, higher densities improve prey detection, but on the other hand, they increase conspecific interference. Copyright © 2015 Elsevier Ltd. All rights reserved.

Coordinated Control of Acoustical Field of View and Flight in Three-Dimensional Space for Consecutive Capture by Echolocating Bats during Natural Foraging.

PubMed

Sumiya, Miwa; Fujioka, Emyo; Motoi, Kazuya; Kondo, Masaru; Hiryu, Shizuko

2017-01-01

Echolocating bats prey upon small moving insects in the dark using sophisticated sonar techniques. The direction and directivity pattern of the ultrasound broadcast of these bats are important factors that affect their acoustical field of view, allowing us to investigate how the bats control their acoustic attention (pulse direction) for advanced flight maneuvers. The purpose of this study was to understand the behavioral strategies of acoustical sensing of wild Japanese house bats Pipistrellus abramus in three-dimensional (3D) space during consecutive capture flights. The results showed that when the bats successively captured multiple airborne insects in short time intervals (less than 1.5 s), they maintained not only the immediate prey but also the subsequent one simultaneously within the beam widths of the emitted pulses in both horizontal and vertical planes before capturing the immediate one. This suggests that echolocating bats maintain multiple prey within their acoustical field of view by a single sensing using a wide directional beam while approaching the immediate prey, instead of frequently shifting acoustic attention between multiple prey. We also numerically simulated the bats' flight trajectories when approaching two prey successively to investigate the relationship between the acoustical field of view and the prey direction for effective consecutive captures. This simulation demonstrated that acoustically viewing both the immediate and the subsequent prey simultaneously increases the success rate of capturing both prey, which is considered to be one of the basic axes of efficient route planning for consecutive capture flight. The bat's wide sonar beam can incidentally cover multiple prey while the bat forages in an area where the prey density is high. Our findings suggest that the bats then keep future targets within their acoustical field of view for effective foraging. In addition, in both the experimental results and the numerical simulations, the acoustic sensing and flights of the bats showed narrower vertical ranges than horizontal ranges. This suggests that the bats control their acoustic sensing according to different schemes in the horizontal and vertical planes according to their surroundings. These findings suggest that echolocating bats coordinate their control of the acoustical field of view and flight for consecutive captures in 3D space during natural foraging.

Coordinated Control of Acoustical Field of View and Flight in Three-Dimensional Space for Consecutive Capture by Echolocating Bats during Natural Foraging

PubMed Central

Sumiya, Miwa; Fujioka, Emyo; Motoi, Kazuya; Kondo, Masaru; Hiryu, Shizuko

2017-01-01

Echolocating bats prey upon small moving insects in the dark using sophisticated sonar techniques. The direction and directivity pattern of the ultrasound broadcast of these bats are important factors that affect their acoustical field of view, allowing us to investigate how the bats control their acoustic attention (pulse direction) for advanced flight maneuvers. The purpose of this study was to understand the behavioral strategies of acoustical sensing of wild Japanese house bats Pipistrellus abramus in three-dimensional (3D) space during consecutive capture flights. The results showed that when the bats successively captured multiple airborne insects in short time intervals (less than 1.5 s), they maintained not only the immediate prey but also the subsequent one simultaneously within the beam widths of the emitted pulses in both horizontal and vertical planes before capturing the immediate one. This suggests that echolocating bats maintain multiple prey within their acoustical field of view by a single sensing using a wide directional beam while approaching the immediate prey, instead of frequently shifting acoustic attention between multiple prey. We also numerically simulated the bats’ flight trajectories when approaching two prey successively to investigate the relationship between the acoustical field of view and the prey direction for effective consecutive captures. This simulation demonstrated that acoustically viewing both the immediate and the subsequent prey simultaneously increases the success rate of capturing both prey, which is considered to be one of the basic axes of efficient route planning for consecutive capture flight. The bat’s wide sonar beam can incidentally cover multiple prey while the bat forages in an area where the prey density is high. Our findings suggest that the bats then keep future targets within their acoustical field of view for effective foraging. In addition, in both the experimental results and the numerical simulations, the acoustic sensing and flights of the bats showed narrower vertical ranges than horizontal ranges. This suggests that the bats control their acoustic sensing according to different schemes in the horizontal and vertical planes according to their surroundings. These findings suggest that echolocating bats coordinate their control of the acoustical field of view and flight for consecutive captures in 3D space during natural foraging. PMID:28085936

Whales: Incredible Ocean Mammals.

ERIC Educational Resources Information Center

Devona, Henry

1992-01-01

Describes an integrated thematic unit for children from kindergarten to third grade. Explains that the unit incorporates reading, speaking, writing, science, mathematics, social studies, and art into the study of whales. Suggests learning activities on echolocation, migration, measurement, scrimshaw, history, and human interaction with the…

Guide to the BATS Resource Trunk.

ERIC Educational Resources Information Center

Arizona Game and Fish Dept., Phoenix.

This guide provides detailed information, resources, and activities to teach students about the bats of Arizona. Chapters include: (1) "What is a Bat?"; (2) "Megabat or Microbat?"; (3) "Bat Anatomy"; (4) Diet and Feeding"; (5) Echolocation"; (6) Reproduction and Lifespan"; (7) "Flight"; (8)…

Bidirectional Echolocation in the Bat Barbastella barbastellus: Different Signals of Low Source Level Are Emitted Upward through the Nose and Downward through the Mouth

PubMed Central

Seibert, Anna-Maria; Koblitz, Jens C.; Denzinger, Annette; Schnitzler, Hans-Ulrich

2015-01-01

The Barbastelle bat (Barbastella barbastellus) preys almost exclusively on tympanate moths. While foraging, this species alternates between two different signal types. We investigated whether these signals differ in emission direction or source level (SL) as assumed from earlier single microphone recordings. We used two different settings of a 16-microphone array to determine SL and sonar beam direction at various locations in the field. Both types of search signals had low SLs (81 and 82 dB SPL rms re 1 m) as compared to other aerial-hawking bats. These two signal types were emitted in different directions; type 1 signals were directed downward and type 2 signals upward. The angle between beam directions was approximately 70°. Barbastelle bats are able to emit signals through both the mouth and the nostrils. As mouth and nostrils are roughly perpendicular to each other, we conclude that type 1 signals are emitted through the mouth while type 2 signals and approach signals are emitted through the nose. We hypothesize that the “stealth” echolocation system of B. barbastellus is bifunctional. The more upward directed nose signals may be mainly used for search and localization of prey. Their low SL prevents an early detection by eared moths but comes at the expense of a strongly reduced detection range for the environment below the bat. The more downward directed mouth signals may have evolved to compensate for this disadvantage and may be mainly used for spatial orientation. We suggest that the possibly bifunctional echolocation system of B. barbastellus has been adapted to the selective foraging of eared moths and is an excellent example of a sophisticated sensory arms race between predator and prey. PMID:26352271

Testing the Sensory Drive Hypothesis: Geographic variation in echolocation frequencies of Geoffroy's horseshoe bat (Rhinolophidae: Rhinolophus clivosus)

PubMed Central

Catto, Sarah; Mutumi, Gregory L.; Finger, Nikita; Webala, Paul W.

2017-01-01

Geographic variation in sensory traits is usually influenced by adaptive processes because these traits are involved in crucial life-history aspects including orientation, communication, lineage recognition and mate choice. Studying this variation can therefore provide insights into lineage diversification. According to the Sensory Drive Hypothesis, lineage diversification may be driven by adaptation of sensory systems to local environments. It predicts that acoustic signals vary in association with local climatic conditions so that atmospheric attenuation is minimized and transmission of the signals maximized. To test this prediction, we investigated the influence of climatic factors (specifically relative humidity and temperature) on geographic variation in the resting frequencies of the echolocation pulses of Geoffroy’s horseshoe bat, Rhinolophus clivosus. If the evolution of phenotypic variation in this lineage tracks climate variation, human induced climate change may lead to decreases in detection volumes and a reduction in foraging efficiency. A complex non-linear interaction between relative humidity and temperature affects atmospheric attenuation of sound and principal components composed of these correlated variables were, therefore, used in a linear mixed effects model to assess their contribution to observed variation in resting frequencies. A principal component composed predominantly of mean annual temperature (factor loading of -0.8455) significantly explained a proportion of the variation in resting frequency across sites (P < 0.05). Specifically, at higher relative humidity (around 60%) prevalent across the distribution of R. clivosus, increasing temperature had a strong negative effect on resting frequency. Climatic factors thus strongly influence acoustic signal divergence in this lineage, supporting the prediction of the Sensory Drive Hypothesis. The predicted future increase in temperature due to climate change is likely to decrease the detection volume in echolocating bats and adversely impact their foraging efficiency. PMID:29186147

The role of ecological factors in shaping bat cone opsin evolution.

PubMed

Gutierrez, Eduardo de A; Schott, Ryan K; Preston, Matthew W; Loureiro, Lívia O; Lim, Burton K; Chang, Belinda S W

2018-04-11

Bats represent one of the largest and most striking nocturnal mammalian radiations, exhibiting many visual system specializations for performance in light-limited environments. Despite representing the greatest ecological diversity and species richness in Chiroptera, Neotropical lineages have been undersampled in molecular studies, limiting the potential for identifying signatures of selection on visual genes associated with differences in bat ecology. Here, we investigated how diverse ecological pressures mediate long-term shifts in selection upon long-wavelength ( Lws ) and short-wavelength ( Sws1 ) opsins, photosensitive cone pigments that form the basis of colour vision in most mammals, including bats. We used codon-based likelihood clade models to test whether ecological variables associated with reliance on visual information (e.g. echolocation ability and diet) or exposure to varying light environments (e.g. roosting behaviour and foraging habitat) mediated shifts in evolutionary rates in bat cone opsin genes. Using additional cone opsin sequences from newly sequenced eye transcriptomes of six Neotropical bat species, we found significant evidence for different ecological pressures influencing the evolution of the cone opsins. While Lws is evolving under significantly lower constraint in highly specialized high-duty cycle echolocating lineages, which have enhanced sonar ability to detect and track targets, variation in Sws1 constraint was significantly associated with foraging habitat, exhibiting elevated rates of evolution in species that forage among vegetation. This suggests that increased reliance on echolocation as well as the spectral environment experienced by foraging bats may differentially influence the evolution of different cone opsins. Our study demonstrates that different ecological variables may underlie contrasting evolutionary patterns in bat visual opsins, and highlights the suitability of clade models for testing ecological hypotheses of visual evolution. © 2018 The Author(s).

Pliocene bats (Chiroptera) from Kanapoi, Turkana Basin, Kenya.

PubMed

Gunnell, Gregg F; Manthi, Fredrick K

2018-04-05

Fossil bats from the Pliocene of Africa are extremely rare, especially in East Africa where meager records have been reported only from two localities in the Omo River Basin Shungura Formation and from a scattering of localities in the Afar Depression, both in Ethiopia. Here we report on a diverse assemblage of bats from Kanapoi in the Turkana Basin that date to approximately 4.19 million years ago. The Kanapoi bat community consists of four different species of fruit bats including a new genus and two new species as well as five species of echolocating bats, the most common of which are two new species of the molossid genus Mops. Additionally, among the echolocating bats, a new species of the emballonurid Saccolaimus is documented at Kanapoi along with an additional Saccolaimus species and a potentially new species of the nycterid Nycteris. Compared to other East African Pliocene bat assemblages, the Kanapoi bat community is unique in preserving molossids and curiously lacks any evidence of cave dwelling bats like rhinolophids or hipposiderids, which are both common at other East African sites. The bats making up the Kanapoi community all typically roost in trees, with some preferring deeper forests and larger trees (molossids), while the others (pteropodids, nycterids and emballonurids) roost in trees near open areas. Living fruit bats that are related to Kanapoi species typically forage for fruits along the margins of forests and in open savannah. The echolocating forms from Kanapoi consist of groups that aerially hawk for insects in open areas between patches of forest and along water courses. The habitats preferred by living relatives of the Kanapoi bats are in agreement with those constructed for Kanapoi based on other lines of evidence. Copyright © 2018. Published by Elsevier Ltd.

Testing the Sensory Drive Hypothesis: Geographic variation in echolocation frequencies of Geoffroy's horseshoe bat (Rhinolophidae: Rhinolophus clivosus).

PubMed

Jacobs, David S; Catto, Sarah; Mutumi, Gregory L; Finger, Nikita; Webala, Paul W

2017-01-01

Geographic variation in sensory traits is usually influenced by adaptive processes because these traits are involved in crucial life-history aspects including orientation, communication, lineage recognition and mate choice. Studying this variation can therefore provide insights into lineage diversification. According to the Sensory Drive Hypothesis, lineage diversification may be driven by adaptation of sensory systems to local environments. It predicts that acoustic signals vary in association with local climatic conditions so that atmospheric attenuation is minimized and transmission of the signals maximized. To test this prediction, we investigated the influence of climatic factors (specifically relative humidity and temperature) on geographic variation in the resting frequencies of the echolocation pulses of Geoffroy's horseshoe bat, Rhinolophus clivosus. If the evolution of phenotypic variation in this lineage tracks climate variation, human induced climate change may lead to decreases in detection volumes and a reduction in foraging efficiency. A complex non-linear interaction between relative humidity and temperature affects atmospheric attenuation of sound and principal components composed of these correlated variables were, therefore, used in a linear mixed effects model to assess their contribution to observed variation in resting frequencies. A principal component composed predominantly of mean annual temperature (factor loading of -0.8455) significantly explained a proportion of the variation in resting frequency across sites (P < 0.05). Specifically, at higher relative humidity (around 60%) prevalent across the distribution of R. clivosus, increasing temperature had a strong negative effect on resting frequency. Climatic factors thus strongly influence acoustic signal divergence in this lineage, supporting the prediction of the Sensory Drive Hypothesis. The predicted future increase in temperature due to climate change is likely to decrease the detection volume in echolocating bats and adversely impact their foraging efficiency.

Scaling of echolocation call parameters in bats.

PubMed

Jones, G

1999-12-01

I investigated the scaling of echolocation call parameters (frequency, duration and repetition rate) in bats in a functional context. Low-duty-cycle bats operate with search phase cycles of usually less than 20 %. They process echoes in the time domain and are therefore intolerant of pulse-echo overlap. High-duty-cycle (>30 %) species use Doppler shift compensation, and they separate pulse and echo in the frequency domain. Call frequency scales negatively with body mass in at least five bat families. Pulse duration scales positively with mass in low-duty-cycle quasi-constant-frequency (QCF) species because the large aerial-hawking species that emit these signals fly fast in open habitats. They therefore detect distant targets and experience pulse-echo overlap later than do smaller bats. Pulse duration also scales positively with mass in the Hipposideridae, which show at least partial Doppler shift compensation. Pulse repetition rate corresponds closely with wingbeat frequency in QCF bat species that fly relatively slowly. Larger, fast-flying species often skip pulses when detecting distant targets. There is probably a trade-off between call intensity and repetition rate because 'whispering' bats (and hipposiderids) produce several calls per predicted wingbeat and because batches of calls are emitted per wingbeat during terminal buzzes. Severe atmospheric attenuation at high frequencies limits the range of high-frequency calls. Low-duty-cycle bats that call at high frequencies must therefore use short pulses to avoid pulse-echo overlap. Rhinolophids escape this constraint by Doppler shift compensation and, importantly, can exploit advantages associated with the emission of both high-frequency and long-duration calls. Low frequencies are unsuited for the detection of small prey, and low repetition rates may limit prey detection rates. Echolocation parameters may therefore constrain maximum body size in aerial-hawking bats.

Biosonar performance of foraging beaked whales (Mesoplodon densirostris).

PubMed

Madsen, P T; Johnson, M; de Soto, N Aguilar; Zimmer, W M X; Tyack, P

2005-01-01

Toothed whales (Cetacea, odontoceti) emit sound pulses to probe their surroundings by active echolocation. Non-invasive, acoustic Dtags were placed on deep-diving Blainville's beaked whales (Mesoplodon densirostris) to record their ultrasonic clicks and the returning echoes from prey items, providing a unique view on how a whale operates its biosonar during foraging in the wild. The process of echolocation during prey capture in this species can be divided into search, approach and terminal phases, as in echolocating bats. The approach phase, defined by the onset of detectable echoes recorded on the tag for click sequences terminated by a buzz, has interclick intervals (ICI) of 300-400 ms. These ICIs are more than a magnitude longer than the decreasing two-way travel time to the targets, showing that ICIs are not given by the two-way-travel times plus a fixed, short lag time. During the approach phase, the received echo energy increases by 10.4(+/-2) dB when the target range is halved, demonstrating that the whales do not employ range-compensating gain control of the transmitter, as has been implicated for some bats and dolphins. The terminal/buzz phase with ICIs of around 10 ms is initiated when one or more targets are within approximately a body length of the whale (2-5 m), so that strong echo returns in the approach phase are traded for rapid updates in the terminal phase. It is suggested that stable ICIs in the search and approach phases facilitate auditory scene analysis in a complex multi-target environment, and that a concomitant low click rate allows the whales to maintain high sound pressure outputs for prey detection and discrimination with a pneumatically driven, bi-modal sound generator.

Blind people are more sensitive than sighted people to binaural sound-location cues, particularly inter-aural level differences.

PubMed

Nilsson, Mats E; Schenkman, Bo N

2016-02-01

Blind people use auditory information to locate sound sources and sound-reflecting objects (echolocation). Sound source localization benefits from the hearing system's ability to suppress distracting sound reflections, whereas echolocation would benefit from "unsuppressing" these reflections. To clarify how these potentially conflicting aspects of spatial hearing interact in blind versus sighted listeners, we measured discrimination thresholds for two binaural location cues: inter-aural level differences (ILDs) and inter-aural time differences (ITDs). The ILDs or ITDs were present in single clicks, in the leading component of click pairs, or in the lagging component of click pairs, exploiting processes related to both sound source localization and echolocation. We tested 23 blind (mean age = 54 y), 23 sighted-age-matched (mean age = 54 y), and 42 sighted-young (mean age = 26 y) listeners. The results suggested greater ILD sensitivity for blind than for sighted listeners. The blind group's superiority was particularly evident for ILD-lag-click discrimination, suggesting not only enhanced ILD sensitivity in general but also increased ability to unsuppress lagging clicks. This may be related to the blind person's experience of localizing reflected sounds, for which ILDs may be more efficient than ITDs. On the ITD-discrimination tasks, the blind listeners performed better than the sighted age-matched listeners, but not better than the sighted young listeners. ITD sensitivity declines with age, and the equal performance of the blind listeners compared to a group of substantially younger listeners is consistent with the notion that blind people's experience may offset age-related decline in ITD sensitivity. Copyright © 2015 The Authors. Published by Elsevier B.V. All rights reserved.

Adaptive evolution of the myo6 gene in old world fruit bats (family: pteropodidae).

PubMed

Shen, Bin; Han, Xiuqun; Jones, Gareth; Rossiter, Stephen J; Zhang, Shuyi

2013-01-01

Myosin VI (encoded by the Myo6 gene) is highly expressed in the inner and outer hair cells of the ear, retina, and polarized epithelial cells such as kidney proximal tubule cells and intestinal enterocytes. The Myo6 gene is thought to be involved in a wide range of physiological functions such as hearing, vision, and clathrin-mediated endocytosis. Bats (Chiroptera) represent one of the most fascinating mammal groups for molecular evolutionary studies of the Myo6 gene. A diversity of specialized adaptations occur among different bat lineages, such as echolocation and associated high-frequency hearing in laryngeal echolocating bats, large eyes and a strong dependence on vision in Old World fruit bats (Pteropodidae), and specialized high-carbohydrate but low-nitrogen diets in both Old World and New World fruit bats (Phyllostomidae). To investigate what role(s) the Myo6 gene might fulfill in bats, we sequenced the coding region of the Myo6 gene in 15 bat species and used molecular evolutionary analyses to detect evidence of positive selection in different bat lineages. We also conducted real-time PCR assays to explore the expression levels of Myo6 in a range of tissues from three representative bat species. Molecular evolutionary analyses revealed that the Myo6 gene, which was widely considered as a hearing gene, has undergone adaptive evolution in the Old World fruit bats which lack laryngeal echolocation and associated high-frequency hearing. Real-time PCR showed the highest expression level of the Myo6 gene in the kidney among ten tissues examined in three bat species, indicating an important role for this gene in kidney function. We suggest that Myo6 has undergone adaptive evolution in Old World fruit bats in relation to receptor-mediated endocytosis for the preservation of protein and essential nutrients.

Bat mortality and activity at a Northern Iowa wind resource area

USGS Publications Warehouse

Jain, A.A.; Koford, Rolf R.; Hancock, A.W.; Zenner, G.G.

2011-01-01

We examined bat collision mortality, activity and species composition at an 89-turbine wind resource area in farmland of north-central Iowa from mid-Apr. to mid-Dec., 2003 and mid-Mar. to mid-Dec., 2004. We found 30 bats beneath turbines on cleared ground and gravel access areas in 2003 and 45 bats in 2004. After adjusting for search probability, search efficiency and scavenging rate, we estimated total bat mortality at 396 ?? 72 (95 ci) in 2003 and 636 ?? 112 (95 ci) in 2004. Although carcasses were mostly migratory tree bats, we found a considerable proportion of little brown bats (Myotis lucifugus). We recorded 1465 bat echolocation call files at turbine sites ( 34.88 call files/detector-night) and 1536 bat call files at adjacent non-turbine sites ( 36.57 call files/detector-night). Bat activity did not differ significantly between turbine and non-turbine sites. A large proportion of recorded call files were made by Myotis sp. but this may be because we detected activity at ground level only. There was no relationship between types of turbine lights and either collision mortality or echolocation activity. The highest levels of bat echolocation activity and collision mortality were recorded during Jul. and Aug. during the autumn dispersal and migration period. The fatality rates for bats in general and little brown bats in particular were higher at the Top of Iowa Wind Resource Area than at other, comparable studies in the region. Future efforts to study behavior of bats in flight around turbines as well as cumulative impact studies should not ignore non-tree dwelling bats, generally regarded as minimally affected. ?? 2011, American Midland Naturalist.

Bidirectional Echolocation in the Bat Barbastella barbastellus: Different Signals of Low Source Level Are Emitted Upward through the Nose and Downward through the Mouth.

PubMed

Seibert, Anna-Maria; Koblitz, Jens C; Denzinger, Annette; Schnitzler, Hans-Ulrich

2015-01-01

The Barbastelle bat (Barbastella barbastellus) preys almost exclusively on tympanate moths. While foraging, this species alternates between two different signal types. We investigated whether these signals differ in emission direction or source level (SL) as assumed from earlier single microphone recordings. We used two different settings of a 16-microphone array to determine SL and sonar beam direction at various locations in the field. Both types of search signals had low SLs (81 and 82 dB SPL rms re 1 m) as compared to other aerial-hawking bats. These two signal types were emitted in different directions; type 1 signals were directed downward and type 2 signals upward. The angle between beam directions was approximately 70°. Barbastelle bats are able to emit signals through both the mouth and the nostrils. As mouth and nostrils are roughly perpendicular to each other, we conclude that type 1 signals are emitted through the mouth while type 2 signals and approach signals are emitted through the nose. We hypothesize that the "stealth" echolocation system of B. barbastellus is bifunctional. The more upward directed nose signals may be mainly used for search and localization of prey. Their low SL prevents an early detection by eared moths but comes at the expense of a strongly reduced detection range for the environment below the bat. The more downward directed mouth signals may have evolved to compensate for this disadvantage and may be mainly used for spatial orientation. We suggest that the possibly bifunctional echolocation system of B. barbastellus has been adapted to the selective foraging of eared moths and is an excellent example of a sophisticated sensory arms race between predator and prey.

Genome-wide signatures of convergent evolution in echolocating mammals

PubMed Central

Parker, Joe; Tsagkogeorga, Georgia; Cotton, James A.; Liu, Yuan; Provero, Paolo; Stupka, Elia; Rossiter, Stephen J.

2013-01-01

Evolution is typically thought to proceed through divergence of genes, proteins, and ultimately phenotypes1-3. However, similar traits might also evolve convergently in unrelated taxa due to similar selection pressures4,5. Adaptive phenotypic convergence is widespread in nature, and recent results from a handful of genes have suggested that this phenomenon is powerful enough to also drive recurrent evolution at the sequence level6-9. Where homoplasious substitutions do occur these have long been considered the result of neutral processes. However, recent studies have demonstrated that adaptive convergent sequence evolution can be detected in vertebrates using statistical methods that model parallel evolution9,10 although the extent to which sequence convergence between genera occurs across genomes is unknown. Here we analyse genomic sequence data in mammals that have independently evolved echolocation and show for the first time that convergence is not a rare process restricted to a handful of loci but is instead widespread, continuously distributed and commonly driven by natural selection acting on a small number of sites per locus. Systematic analyses of convergent sequence evolution in 805,053 amino acids within 2,326 orthologous coding gene sequences compared across 22 mammals (including four new bat genomes) revealed signatures consistent with convergence in nearly 200 loci. Strong and significant support for convergence among bats and the dolphin was seen in numerous genes linked to hearing or deafness, consistent with an involvement in echolocation. Surprisingly we also found convergence in many genes linked to vision: the convergent signal of many sensory genes was robustly correlated with the strength of natural selection. This first attempt to detect genome-wide convergent sequence evolution across divergent taxa reveals the phenomenon to be much more pervasive than previously recognised. PMID:24005325

Dynamic representation of 3D auditory space in the midbrain of the free-flying echolocating bat

PubMed Central

2018-01-01

Essential to spatial orientation in the natural environment is a dynamic representation of direction and distance to objects. Despite the importance of 3D spatial localization to parse objects in the environment and to guide movement, most neurophysiological investigations of sensory mapping have been limited to studies of restrained subjects, tested with 2D, artificial stimuli. Here, we show for the first time that sensory neurons in the midbrain superior colliculus (SC) of the free-flying echolocating bat encode 3D egocentric space, and that the bat’s inspection of objects in the physical environment sharpens tuning of single neurons, and shifts peak responses to represent closer distances. These findings emerged from wireless neural recordings in free-flying bats, in combination with an echo model that computes the animal’s instantaneous stimulus space. Our research reveals dynamic 3D space coding in a freely moving mammal engaged in a real-world navigation task. PMID:29633711

Comparing phase-sensitive and phase-insensitive echolocation target images using a monaural audible sonar.

PubMed

Kuc, Roman

2018-04-01

This paper describes phase-sensitive and phase-insensitive processing of monaural echolocation waveforms to generate target maps. Composite waveforms containing both the emission and echoes are processed to estimate the target impulse response using an audible sonar. Phase-sensitive processing yields the composite signal envelope, while phase-insensitive processing that starts with the composite waveform power spectrum yields the envelope of the autocorrelation function. Analysis and experimental verification show that multiple echoes form an autocorrelation function that produces near-range phantom-reflector artifacts. These artifacts interfere with true target echoes when the first true echo occurs at a time that is less than the total duration of the target echoes. Initial comparison of phase-sensitive and phase-insensitive maps indicates that both display important target features, indicating that phase is not vital. A closer comparison illustrates the improved resolution of phase-sensitive processing, the near-range phantom-reflectors produced by phase-insensitive processing, and echo interference and multiple reflection artifacts that were independent of the processing.

The evolution of sensory divergence in the context of limited gene flow in the bumblebee bat

PubMed Central

Puechmaille, Sébastien J.; Gouilh, Meriadeg Ar; Piyapan, Piyathip; Yokubol, Medhi; Mie, Khin Mie; Bates, Paul J.; Satasook, Chutamas; Nwe, Tin; Bu, Si Si Hla; Mackie, Iain J.; Petit, Eric J.; Teeling, Emma C.

2011-01-01

The sensory drive theory of speciation predicts that populations of the same species inhabiting different environments can differ in sensory traits, and that this sensory difference can ultimately drive speciation. However, even in the best-known examples of sensory ecology driven speciation, it is uncertain whether the variation in sensory traits is the cause or the consequence of a reduction in levels of gene flow. Here we show strong genetic differentiation, no gene flow and large echolocation differences between the allopatric Myanmar and Thai populations of the world's smallest mammal, Craseonycteris thonglongyai, and suggest that geographic isolation most likely preceded sensory divergence. Within the geographically continuous Thai population, we show that geographic distance has a primary role in limiting gene flow rather than echolocation divergence. In line with sensory-driven speciation models, we suggest that in C. thonglongyai, limited gene flow creates the suitable conditions that favour the evolution of sensory divergence via local adaptation. PMID:22146392

Biosonar behaviour of free-ranging porpoises.

PubMed

Akamatsu, Tomonari; Wang, Ding; Wang, Kexiong; Naito, Yasuhiko

2005-04-22

Detecting objects in their paths is a fundamental perceptional function of moving organisms. Potential risks and rewards, such as prey, predators, conspecifics or non-biological obstacles, must be detected so that an animal can modify its behaviour accordingly. However, to date few studies have considered how animals in the wild focus their attention. Dolphins and porpoises are known to actively use sonar or echolocation. A newly developed miniature data logger attached to a porpoise allows for individual recording of acoustical search efforts and inspection distance based on echolocation. In this study, we analysed the biosonar behaviour of eight free-ranging finless porpoises (Neophocaena phocaenoides) and demonstrated that these animals inspect the area ahead of them before swimming silently into it. The porpoises inspected distances up to 77 m, whereas their swimming distance without using sonar was less than 20 m. The inspection distance was long enough to ensure a wide safety margin before facing real risks or rewards. Once a potential prey item was detected, porpoises adjusted their inspection distance from the remote target throughout their approach.

Range-dependent flexibility in the acoustic field of view of echolocating porpoises (Phocoena phocoena)

PubMed Central

Wisniewska, Danuta M; Ratcliffe, John M; Beedholm, Kristian; Christensen, Christian B; Johnson, Mark; Koblitz, Jens C; Wahlberg, Magnus; Madsen, Peter T

2015-01-01

Toothed whales use sonar to detect, locate, and track prey. They adjust emitted sound intensity, auditory sensitivity and click rate to target range, and terminate prey pursuits with high-repetition-rate, low-intensity buzzes. However, their narrow acoustic field of view (FOV) is considered stable throughout target approach, which could facilitate prey escape at close-range. Here, we show that, like some bats, harbour porpoises can broaden their biosonar beam during the terminal phase of attack but, unlike bats, maintain the ability to change beamwidth within this phase. Based on video, MRI, and acoustic-tag recordings, we propose this flexibility is modulated by the melon and implemented to accommodate dynamic spatial relationships with prey and acoustic complexity of surroundings. Despite independent evolution and different means of sound generation and transmission, whales and bats adaptively change their FOV, suggesting that beamwidth flexibility has been an important driver in the evolution of echolocation for prey tracking. DOI: http://dx.doi.org/10.7554/eLife.05651.001 PMID:25793440

Range-dependent flexibility in the acoustic field of view of echolocating porpoises (Phocoena phocoena).

PubMed

Wisniewska, Danuta M; Ratcliffe, John M; Beedholm, Kristian; Christensen, Christian B; Johnson, Mark; Koblitz, Jens C; Wahlberg, Magnus; Madsen, Peter T

2015-03-20

Toothed whales use sonar to detect, locate, and track prey. They adjust emitted sound intensity, auditory sensitivity and click rate to target range, and terminate prey pursuits with high-repetition-rate, low-intensity buzzes. However, their narrow acoustic field of view (FOV) is considered stable throughout target approach, which could facilitate prey escape at close-range. Here, we show that, like some bats, harbour porpoises can broaden their biosonar beam during the terminal phase of attack but, unlike bats, maintain the ability to change beamwidth within this phase. Based on video, MRI, and acoustic-tag recordings, we propose this flexibility is modulated by the melon and implemented to accommodate dynamic spatial relationships with prey and acoustic complexity of surroundings. Despite independent evolution and different means of sound generation and transmission, whales and bats adaptively change their FOV, suggesting that beamwidth flexibility has been an important driver in the evolution of echolocation for prey tracking.

Sound-conducting mechanisms for echolocation hearing of a dolphin

NASA Astrophysics Data System (ADS)

Ryabov, Vyacheslav A.

2005-09-01

The morphological study of the lower jaw of a dolphin (Tursiops truncatus p.), and the modeling and calculation of its structures from the acoustic point of view have been conducted. It was determined that the cross-sectional area of the mandibular canal (MC) increases exponentially. The MC represents the acoustical horn. The mental foramens (MFs) is positioned in the horn throat, representing the nonequidistant array of waveguide delay lines (NAWDL). The acoustical horn ensures the traveling wave conditions inside the MC and intensifies sonar echoes up to 1514 times. This ``ideal'' traveling wave antenna is created by nature, representing the combination of the NAWDL and the acoustical horn. The dimensions and sequence of morphological structures of the lower jaw are optimal both for reception and forming the beam pattern, and for the amplification and transmission of sonar echoes up to the bulla tympani. Morphological structures of the lower jaw are considered as components of the peripheral section of the dolphin echolocation hearing.

Behavioural and neurobiological implications of linear and non-linear features in larynx phonations of horseshoe bats

PubMed Central

Kobayasi, Kohta I.; Hage, Steffen R.; Berquist, Sean; Feng, Jiang; Zhang, Shuyi; Metzner, Walter

2012-01-01

Mammalian vocalizations exhibit large variations in their spectrotemporal features, although it is still largely unknown which result from intrinsic biomechanical properties of the larynx and which are under direct neuromuscular control. Here we show that mere changes in laryngeal air flow yield several non-linear effects on sound production, in an isolated larynx preparation from horseshoe bats. Most notably, there are sudden jumps between two frequency bands used for either echolocation or communication in natural vocalizations. These jumps resemble changes in “registers” as in yodelling. In contrast, simulated contractions of the main larynx muscle produce linear frequency changes, but are limited to echolocation or communication frequencies. Only by combining non-linear and linear properties can this larynx therefore produce sounds covering the entire frequency range of natural calls. This may give behavioural meaning to yodelling-like vocal behaviour and reshape our thinking about how the brain controls the multitude of spectral vocal features in mammals. PMID:23149729

Cranial asymmetry in Eocene archaeocete whales and the evolution of directional hearing in water

PubMed Central

Fahlke, Julia M.; Gingerich, Philip D.; Welsh, Robert C.; Wood, Aaron R.

2011-01-01

Eocene archaeocete whales gave rise to all modern toothed and baleen whales (Odontoceti and Mysticeti) during or near the Eocene-Oligocene transition. Odontocetes have asymmetrical skulls, with asymmetry linked to high-frequency sound production and echolocation. Mysticetes are generally assumed to have symmetrical skulls and lack high-frequency hearing. Here we show that protocetid and basilosaurid archaeocete skulls are distinctly and directionally asymmetrical. Archaeocete asymmetry involves curvature and axial torsion of the cranium, but no telescoping. Cranial asymmetry evolved in Eocene archaeocetes as part of a complex of traits linked to directional hearing (such as pan-bone thinning of the lower jaws, mandibular fat pads, and isolation of the ear region), probably enabling them to hear the higher sonic frequencies of sound-producing fish on which they preyed. Ultrasonic echolocation evolved in Oligocene odontocetes, enabling them to find silent prey. Asymmetry and much of the sonic-frequency range of directional hearing were lost in Oligocene mysticetes during the shift to low-frequency hearing and bulk-straining predation. PMID:21873217

Echolocation: A Study of Auditory Functioning in Blind and Sighted Subjects.

ERIC Educational Resources Information Center

Arias, C.; And Others

1993-01-01

This study evaluated the peripheral and central auditory functioning (and thus the potential to perceive obstacles through reflected sound) of eight totally blind persons and eight sighted persons. The blind subjects were able to process auditory information faster than the control group. (Author/DB)

Bat Facts and Fun.

ERIC Educational Resources Information Center

McKee, Judith A.

1992-01-01

Describes a unit of study for elementary school science on bats. Students investigate the different types of bats; examine their behavior; find facts that other students are unlikely to know; write stories about bats; and examine the concept of echolocation, the means by which bats navigate. Suggests integrated activities for mathematics…

Physics and Size in Biological Systems.

ERIC Educational Resources Information Center

Barnes, George

1989-01-01

Described is the subject of biological scaling for physics teachers including examples and in-depth reading. Topics are elements of scaling, terminal velocities, Lilliputian and Brobdingnagian, brain evolution, dolphin echolocation, surface tension, gravity change, food and oxygen, and seeing. Ten references on physics and size, and ten questions…

Sensory Biology: Acoustic Reflectors Attract Bats to Roost in Pitcher Plants.

PubMed

Jones, Gareth

2015-07-20

A new study shows that a carnivorous plant attracts bats by possessing modified pitfall taps that increase the reflectivity of echolocation calls. Bats benefit by finding roosting sites, and the plants gain by receiving nitrogen from guano. Copyright © 2015 Elsevier Ltd. All rights reserved.

RESEARCH BULLETIN. NUMBER 4.

ERIC Educational Resources Information Center

CLARK, LESLIE L., ED.

THE FOLLOWING FOUR RESEARCH REPORTS ON ORIENTATION AND MOBILITY FOR THE BLIND BASED ON AURAL INFORMATION ARE CONTAINED IN THIS BULLETIN--"A PSYCHOACOUSTIC STUDY OF FACTORS AFFECTING HUMAN ECHOLOCATION" BY JOHN R. WELCH, "ORIENTATION BY AURAL CLUES" BY IVO KOHLER, "SONAR SYSTEM OF THE BLIND" BY WINTHROP N. KELLOGG, AND "TRIAL OF AN ACOUSTIC BLIND…

Marine Mammals: Hearing and Echolocation at Coconut Island

DTIC Science & Technology

2010-01-01

biosonar discrimination capability which may result in reduced ability to forage A formerly stranded long finned pilot whale and a recently stranded...control in odontocete biosonar : an evoked-potential study. J Acous Soc Am Yuen, M.E., Nachtigall, P.E., and Supin, A.Ya., and Breese, M. (2005

Pursuit, Avoidance, and Cohesion in Flight: Multi-Purpose Control Laws and Neuromorphic VLSI

DTIC Science & Technology

2010-10-01

34 Binaural Spectral Cues for Ultrasonic Localization," Proc. International Symposium on Circuits and Systems, pp. 2110 - 2113, 2008 (DOI:10.1109/ISCAS...T. K. Horiuchi, C. Bansal, and T. M. Massoud (2009), " Binaural Intensity Comparison in the Echolocating Bat Using Synaptic Conductance," Proc

78 FR 11844 - Taking of Marine Mammals Incidental to Specified Activities; Bremerton Ferry Terminal Wingwall...

Federal Register 2010, 2011, 2012, 2013, 2014

2013-02-20

... high- intensity, noise could cause masking at particular frequencies for marine mammals that utilize... mostly concentrated at low frequency ranges, it may have less effect on high frequency echolocation... reduce the communication space of animals (e.g., Clark et al. 2009) and cause increased stress levels (e...

Basic Hearing and Echolocation Mechanisms of Marine Mammals: Measured Auditory Evoked Potential and Behavioral Experiments FY 2007

DTIC Science & Technology

2007-01-01

Breese, M. (2007) Evoked-potential recovery during double click stimulation in a whale: A possibility of biosonar automatic gain control. Journal of...Yokohama Japan (published) Supin A.Ya, Nachtigall, P.E., and Breese, M. Source level to sensation level ratio of transmitted biosonar pulses in an

Beaked Whale Presence, Habitat, and Sound Production in the North Pacific

DTIC Science & Technology

2011-09-30

Schnitzler, and J. A. Hildebrand. 2010. Echolocation signals of a beaked whale at Palmyra Atoll . Journal of the Acoustical Society of America 127:3790...McDonald, M. A., J. A. Hildebrand, S. M. Wiggins, D. W. Johnston , and J. J. Polovina. 2009. An acoustic survey of beaked whales at Cross Seamount near

Some recollections of D. R. Griffin as a young man

NASA Astrophysics Data System (ADS)

Galambos, Robert

2004-05-01

In 1939 Don Griffin invited me to join him in his earliest bat echolocation experiments. I will tell a few stories about what we two graduate students did together, and show the sound movie in which, for the first time, we recorded their cries as they flew and avoided obstacles.

Sperm whale long-range echolocation sounds revealed by ANTARES, a deep-sea neutrino telescope.

PubMed

André, M; Caballé, A; van der Schaar, M; Solsona, A; Houégnigan, L; Zaugg, S; Sánchez, A M; Castell, J V; Solé, M; Vila, F; Djokic, D; Adrián-Martínez, S; Albert, A; Anghinolfi, M; Anton, G; Ardid, M; Aubert, J-J; Avgitas, T; Baret, B; Barrios-Martí, J; Basa, S; Bertin, V; Biagi, S; Bormuth, R; Bouwhuis, M C; Bruijn, R; Brunner, J; Busto, J; Capone, A; Caramete, L; Carr, J; Celli, S; Chiarusi, T; Circella, M; Coleiro, A; Coniglione, R; Costantini, H; Coyle, P; Creusot, A; Deschamps, A; De Bonis, G; Distefano, C; Di Palma, I; Donzaud, C; Dornic, D; Drouhin, D; Eberl, T; El Bojaddaini, I; Elsässer, D; Enzenhöfer, A; Fehn, K; Felis, I; Fusco, L A; Galatà, S; Gay, P; Geißelsöder, S; Geyer, K; Giordano, V; Gleixner, A; Glotin, H; Gracia-Ruiz, R; Graf, K; Hallmann, S; van Haren, H; Heijboer, A J; Hello, Y; Hernandez-Rey, J J; Hößl, J; Hofestädt, J; Hugon, C; Illuminati, G; James, C W; de Jong, M; Jongen, M; Kadler, M; Kalekin, O; Katz, U; Kießling, D; Kouchner, A; Kreter, M; Kreykenbohm, I; Kulikovskiy, V; Lachaud, C; Lahmann, R; Lefèvre, D; Leonora, E; Loucatos, S; Marcelin, M; Margiotta, A; Marinelli, A; Martínez-Mora, J A; Mathieu, A; Melis, K; Michael, T; Migliozzi, P; Moussa, A; Mueller, C; Nezri, E; Păvălaş, G E; Pellegrino, C; Perrina, C; Piattelli, P; Popa, V; Pradier, T; Racca, C; Riccobene, G; Roensch, K; Saldaña, M; Samtleben, D F E; Sanguineti, M; Sapienza, P; Schnabel, J; Schüssler, F; Seitz, T; Sieger, C; Spurio, M; Stolarczyk, Th; Sánchez-Losa, A; Taiuti, M; Trovato, A; Tselengidou, M; Turpin, D; Tönnis, C; Vallage, B; Vallée, C; Van Elewyck, V; Vivolo, D; Wagner, S; Wilms, J; Zornoza, J D; Zuñiga, J

2017-04-12

Despite dedicated research has been carried out to adequately map the distribution of the sperm whale in the Mediterranean Sea, unlike other regions of the world, the species population status is still presently uncertain. The analysis of two years of continuous acoustic data provided by the ANTARES neutrino telescope revealed the year-round presence of sperm whales in the Ligurian Sea, probably associated with the availability of cephalopods in the region. The presence of the Ligurian Sea sperm whales was demonstrated through the real-time analysis of audio data streamed from a cabled-to-shore deep-sea observatory that allowed the hourly tracking of their long-range echolocation behaviour on the Internet. Interestingly, the same acoustic analysis indicated that the occurrence of surface shipping noise would apparently not condition the foraging behaviour of the sperm whale in the area, since shipping noise was almost always present when sperm whales were acoustically detected. The continuous presence of the sperm whale in the region confirms the ecological value of the Ligurian sea and the importance of ANTARES to help monitoring its ecosystems.

Localization and tracking of moving objects in two-dimensional space by echolocation.

PubMed

Matsuo, Ikuo

2013-02-01

Bats use frequency-modulated echolocation to identify and capture moving objects in real three-dimensional space. Experimental evidence indicates that bats are capable of locating static objects with a range accuracy of less than 1 μs. A previously introduced model estimates ranges of multiple, static objects using linear frequency modulation (LFM) sound and Gaussian chirplets with a carrier frequency compatible with bat emission sweep rates. The delay time for a single object was estimated with an accuracy of about 1.3 μs by measuring the echo at a low signal-to-noise ratio (SNR). The range accuracy was dependent not only on the SNR but also the Doppler shift, which was dependent on the movements. However, it was unclear whether this model could estimate the moving object range at each timepoint. In this study, echoes were measured from the rotating pole at two receiving points by intermittently emitting LFM sounds. The model was shown to localize moving objects in two-dimensional space by accurately estimating the object's range at each timepoint.

Acoustic characteristics of biosonar sounds of free-ranging botos (Inia geoffrensis) and tucuxis (Sotalia fluviatilis) in the Negro River, Amazon, Brazil.

PubMed

Yamamoto, Yukiko; Akamatsu, Tomonari; da Silva, Vera M F; Yoshida, Yayoi; Kohshima, Shiro

2015-08-01

Odontoceti emit broadband high-frequency clicks on echolocation for orientation or prey detection. In the Amazon Basin, two odontoceti species, boto (Amazon River dolphin, Inia geoffrensis) and tucuxi (Sotalia fluviatilis), live sympatrically. The acoustic characteristics of the echolocation clicks of free-ranging botos and tucuxis were measured with a hydrophone array consisting of a full-band and an acoustic event recorder (A-tag). The clicks of the two species were short-duration broadband signals. The apparent source level was 201 dB 1 μPa peak-to-peak at 1 m in the botos and 181 dB 1 μPa peak-to-peak at 1 m in the tucuxis, and the centroid frequency was 82.3 kHz in the botos and 93.1 kHz in the tucuxis. The high apparent source level and low centroid frequency are possibly due to the difference in body size or sound production organs, especially the nasal structure, the sound source of clicks in odontoceti.

Stereotypical rapid source level regulation in the harbour porpoise biosonar

NASA Astrophysics Data System (ADS)

Linnenschmidt, Meike; Kloepper, Laura N.; Wahlberg, Magnus; Nachtigall, Paul E.

2012-09-01

Some odontocetes and bats vary both click intensity and receiver sensitivity during echolocation, depending on target range. It is not known how this so-called automatic gain control is regulated by the animal. The source level of consecutive echolocation clicks from a harbour porpoise was measured with a hydrophone array while the animal detected an aluminium cylinder at 2, 4 or 8 m distance in a go/no-go paradigm. On-axis clicks had source levels of 145-174 dB re 1 μPa peak-to-peak. During target-present trials the click trains reached comparable source levels independent of the range to the target after three clicks. After an additional click, the source level was reduced for the 2 and 4 m trials until it equalled the one-way transmission loss. During target-absent trials, the source level remained high throughout the entire click train. Given typical values of harbour porpoise inter-click intervals, the source level reduction commenced within a few 100 ms from the first click in the click train. This may indicate a sub-cortically regulated source level regulation in the harbour porpoise.

Hawkmoths produce anti-bat ultrasound

PubMed Central

Barber, Jesse R.; Kawahara, Akito Y.

2013-01-01

Bats and moths have been engaged in aerial warfare for nearly 65 Myr. This arms race has produced a suite of counter-adaptations in moths, including bat-detecting ears. One set of defensive strategies involves the active production of sound; tiger moths' ultrasonic replies to bat attack have been shown to startle bats, warn the predators of bad taste and jam their biosonar. Here, we report that hawkmoths in the Choerocampina produce entirely ultrasonic sounds in response to tactile stimulation and the playback of biosonar attack sequences. Males do so by grating modified scraper scales on the outer surface of the genital valves against the inner margin of the last abdominal tergum. Preliminary data indicate that females also produce ultrasound to touch and playback of echolocation attack, but they do so with an entirely different mechanism. The anti-bat function of these sounds is unknown but might include startling, cross-family acoustic mimicry, warning of unprofitability or physical defence and/or jamming of echolocation. Hawkmoths present a novel and tractable system to study both the function and evolution of anti-bat defences. PMID:23825084

Biosonar behaviour of free-ranging porpoises

PubMed Central

Akamatsu, Tomonari; Wang, Ding; Wang, Kexiong; Naito, Yasuhiko

2005-01-01

Detecting objects in their paths is a fundamental perceptional function of moving organisms. Potential risks and rewards, such as prey, predators, conspecifics or non-biological obstacles, must be detected so that an animal can modify its behaviour accordingly. However, to date few studies have considered how animals in the wild focus their attention. Dolphins and porpoises are known to actively use sonar or echolocation. A newly developed miniature data logger attached to a porpoise allows for individual recording of acoustical search efforts and inspection distance based on echolocation. In this study, we analysed the biosonar behaviour of eight free-ranging finless porpoises (Neophocaena phocaenoides) and demonstrated that these animals inspect the area ahead of them before swimming silently into it. The porpoises inspected distances up to 77 m, whereas their swimming distance without using sonar was less than 20 m. The inspection distance was long enough to ensure a wide safety margin before facing real risks or rewards. Once a potential prey item was detected, porpoises adjusted their inspection distance from the remote target throughout their approach. PMID:15888412

Doppler-shift compensation in the Taiwanese leaf-nosed bat (Hipposideros terasensis) recorded with a telemetry microphone system during flight

NASA Astrophysics Data System (ADS)

Hiryu, Shizuko; Katsura, Koji; Lin, Liang-Kong; Riquimaroux, Hiroshi; Watanabe, Yoshiaki

2005-12-01

Biosonar behavior was examined in Taiwanese leaf-nosed bats (Hipposideros terasensis; CF-FM bats) during flight. Echolocation sounds were recorded using a telemetry microphone mounted on the bat's head. Flight speed and three-dimensional trajectory of the bat were reconstructed from images taken with a dual high-speed video camera system. Bats were observed to change the intensity and emission rate of pulses depending on the distance from the landing site. Frequencies of the dominant second harmonic constant frequency component (CF2) of calls estimated from the bats' flight speed agreed strongly with observed values. Taiwanese leaf-nosed bats changed CF2 frequencies depending on flight speed, which caused the CF2 frequencies of the Doppler-shifted echoes to remain constant. Pulse frequencies were also estimated using echoes returning directly ahead of the bat and from its sides for two different flight conditions: landing and U-turn. Bats in flight may periodically alter their attended angles from the front to the side when emitting echolocation pulses.

Correlated evolution between hearing sensitivity and social calls in bats

PubMed Central

Bohn, Kirsten M; Moss, Cynthia F; Wilkinson, Gerald S

2006-01-01

Echolocating bats are auditory specialists, with exquisite hearing that spans several octaves. In the ultrasonic range, bat audiograms typically show highest sensitivity in the spectral region of their species-specific echolocation calls. Well-developed hearing in the audible range has been commonly attributed to a need to detect sounds produced by prey. However, bat pups often emit isolation calls with low-frequency components that facilitate mother–young reunions. In this study, we examine whether low-frequency hearing in bats exhibits correlated evolution with (i) body size; (ii) high-frequency hearing sensitivity or (iii) pup isolation call frequency. Using published audiograms, we found that low-frequency hearing sensitivity is not dependent on body size but is related to high-frequency hearing. After controlling for high-frequency hearing, we found that low-frequency hearing exhibits correlated evolution with isolation call frequency. We infer that detection and discrimination of isolation calls have favoured enhanced low-frequency hearing because accurate parental investment is critical: bats have low reproductive rates, non-volant altricial young and must often identify their pups within large crèches. PMID:17148288

Neural Processing of Target Distance by Echolocating Bats: Functional Roles of the Auditory Midbrain

PubMed Central

Wenstrup, Jeffrey J.; Portfors, Christine V.

2011-01-01

Using their biological sonar, bats estimate distance to avoid obstacles and capture moving prey. The primary distance cue is the delay between the bat's emitted echolocation pulse and the return of an echo. The mustached bat's auditory midbrain (inferior colliculus, IC) is crucial to the analysis of pulse-echo delay. IC neurons are selective for certain delays between frequency modulated (FM) elements of the pulse and echo. One role of the IC is to create these “delay-tuned”, “FM-FM” response properties through a series of spectro-temporal integrative interactions. A second major role of the midbrain is to project target distance information to many parts of the brain. Pathways through auditory thalamus undergo radical reorganization to create highly ordered maps of pulse-echo delay in auditory cortex, likely contributing to perceptual features of target distance analysis. FM-FM neurons in IC also project strongly to pre-motor centers including the pretectum and the pontine nuclei. These pathways may contribute to rapid adjustments in flight, body position, and sonar vocalizations that occur as a bat closes in on a target. PMID:21238485

Echolocation with bat buzz emissions: model and biomimetic sonar for elevation estimation.

PubMed

Kuc, Roman

2012-01-01

Just prior to capture the Buzz II emissions of some mouth-emitting bats, such as Eptesicus fuscus, are observed to exhibit spectra having multiple peaks. This paper proposes an echolocation strategy that uses such spectra with energy concentrated in specific frequency bands for determining target elevation. A biomimetic sonar was implemented to produce a tri-modal spectrum by driving a speaker with a signal rich in harmonics. The emission magnitudes at these harmonic frequencies measured as a function of elevation in the zero-azimuth plane form distinct beams. A template was formed from the ratio of the first harmonic and fundamental magnitudes to determine elevation. The elevation estimator exhibited a sub-degree accuracy (SD = 0.4° over a 20° interval centered at the elevation at which these two beams intersect in the zero-azimuth plane. Spectral cues from -40° to +10° elevation allow a qualitative non-linear control of sonar orientation to drive the target to the beam-intersection point where quantitative elevation estimates are available. © 2012 Acoustical Society of America.

Sperm whale long-range echolocation sounds revealed by ANTARES, a deep-sea neutrino telescope

NASA Astrophysics Data System (ADS)

André, M.; Caballé, A.; van der Schaar, M.; Solsona, A.; Houégnigan, L.; Zaugg, S.; Sánchez, A. M.; Castell, J. V.; Solé, M.; Vila, F.; Djokic, D.; Adrián-Martínez, S.; Albert, A.; Anghinolfi, M.; Anton, G.; Ardid, M.; Aubert, J.-J.; Avgitas, T.; Baret, B.; Barrios-Martí, J.; Basa, S.; Bertin, V.; Biagi, S.; Bormuth, R.; Bouwhuis, M. C.; Bruijn, R.; Brunner, J.; Busto, J.; Capone, A.; Caramete, L.; Carr, J.; Celli, S.; Chiarusi, T.; Circella, M.; Coleiro, A.; Coniglione, R.; Costantini, H.; Coyle, P.; Creusot, A.; Deschamps, A.; de Bonis, G.; Distefano, C.; di Palma, I.; Donzaud, C.; Dornic, D.; Drouhin, D.; Eberl, T.; El Bojaddaini, I.; Elsässer, D.; Enzenhöfer, A.; Fehn, K.; Felis, I.; Fusco, L. A.; Galatà, S.; Gay, P.; Geißelsöder, S.; Geyer, K.; Giordano, V.; Gleixner, A.; Glotin, H.; Gracia-Ruiz, R.; Graf, K.; Hallmann, S.; van Haren, H.; Heijboer, A. J.; Hello, Y.; Hernandez-Rey, J. J.; Hößl, J.; Hofestädt, J.; Hugon, C.; Illuminati, G.; James, C. W.; de Jong, M.; Jongen, M.; Kadler, M.; Kalekin, O.; Katz, U.; Kießling, D.; Kouchner, A.; Kreter, M.; Kreykenbohm, I.; Kulikovskiy, V.; Lachaud, C.; Lahmann, R.; Lefèvre, D.; Leonora, E.; Loucatos, S.; Marcelin, M.; Margiotta, A.; Marinelli, A.; Martínez-Mora, J. A.; Mathieu, A.; Melis, K.; Michael, T.; Migliozzi, P.; Moussa, A.; Mueller, C.; Nezri, E.; Păvălaş, G. E.; Pellegrino, C.; Perrina, C.; Piattelli, P.; Popa, V.; Pradier, T.; Racca, C.; Riccobene, G.; Roensch, K.; Saldaña, M.; Samtleben, D. F. E.; Sanguineti, M.; Sapienza, P.; Schnabel, J.; Schüssler, F.; Seitz, T.; Sieger, C.; Spurio, M.; Stolarczyk, Th.; Sánchez-Losa, A.; Taiuti, M.; Trovato, A.; Tselengidou, M.; Turpin, D.; Tönnis, C.; Vallage, B.; Vallée, C.; van Elewyck, V.; Vivolo, D.; Wagner, S.; Wilms, J.; Zornoza, J. D.; Zuñiga, J.

2017-04-01

Despite dedicated research has been carried out to adequately map the distribution of the sperm whale in the Mediterranean Sea, unlike other regions of the world, the species population status is still presently uncertain. The analysis of two years of continuous acoustic data provided by the ANTARES neutrino telescope revealed the year-round presence of sperm whales in the Ligurian Sea, probably associated with the availability of cephalopods in the region. The presence of the Ligurian Sea sperm whales was demonstrated through the real-time analysis of audio data streamed from a cabled-to-shore deep-sea observatory that allowed the hourly tracking of their long-range echolocation behaviour on the Internet. Interestingly, the same acoustic analysis indicated that the occurrence of surface shipping noise would apparently not condition the foraging behaviour of the sperm whale in the area, since shipping noise was almost always present when sperm whales were acoustically detected. The continuous presence of the sperm whale in the region confirms the ecological value of the Ligurian sea and the importance of ANTARES to help monitoring its ecosystems.

Sperm whale long-range echolocation sounds revealed by ANTARES, a deep-sea neutrino telescope

PubMed Central

André, M.; Caballé, A.; van der Schaar, M.; Solsona, A.; Houégnigan, L.; Zaugg, S.; Sánchez, A. M.; Castell, J. V.; Solé, M.; Vila, F.; Djokic, D.; Adrián-Martínez, S.; Albert, A.; Anghinolfi, M.; Anton, G.; Ardid, M.; Aubert, J.-J.; Avgitas, T.; Baret, B.; Barrios-Martí, J.; Basa, S.; Bertin, V.; Biagi, S.; Bormuth, R.; Bouwhuis, M. C.; Bruijn, R.; Brunner, J.; Busto, J.; Capone, A.; Caramete, L.; Carr, J.; Celli, S.; Chiarusi, T.; Circella, M.; Coleiro, A.; Coniglione, R.; Costantini, H.; Coyle, P.; Creusot, A.; Deschamps, A.; De Bonis, G.; Distefano, C.; Di Palma, I.; Donzaud, C.; Dornic, D.; Drouhin, D.; Eberl, T.; El Bojaddaini, I.; Elsässer, D.; Enzenhöfer, A.; Fehn, K.; Felis, I.; Fusco, L. A.; Galatà, S.; Gay, P.; Geißelsöder, S.; Geyer, K.; Giordano, V.; Gleixner, A.; Glotin, H.; Gracia-Ruiz, R.; Graf, K.; Hallmann, S.; van Haren, H.; Heijboer, A. J.; Hello, Y.; Hernandez-Rey, J. J.; Hößl, J.; Hofestädt, J.; Hugon, C.; Illuminati, G.; James, C. W.; de Jong, M.; Jongen, M.; Kadler, M.; Kalekin, O.; Katz, U.; Kießling, D.; Kouchner, A.; Kreter, M.; Kreykenbohm, I.; Kulikovskiy, V.; Lachaud, C.; Lahmann, R.; Lefèvre, D.; Leonora, E.; Loucatos, S.; Marcelin, M.; Margiotta, A.; Marinelli, A.; Martínez-Mora, J. A.; Mathieu, A.; Melis, K.; Michael, T.; Migliozzi, P.; Moussa, A.; Mueller, C.; Nezri, E.; Păvălaş, G. E.; Pellegrino, C.; Perrina, C.; Piattelli, P.; Popa, V.; Pradier, T.; Racca, C.; Riccobene, G.; Roensch, K.; Saldaña, M.; Samtleben, D. F. E.; Sanguineti, M.; Sapienza, P.; Schnabel, J.; Schüssler, F.; Seitz, T.; Sieger, C.; Spurio, M.; Stolarczyk, Th.; Sánchez-Losa, A.; Taiuti, M.; Trovato, A.; Tselengidou, M.; Turpin, D.; Tönnis, C.; Vallage, B.; Vallée, C.; Van Elewyck, V.; Vivolo, D.; Wagner, S.; Wilms, J.; Zornoza, J. D.; Zuñiga, J.

2017-01-01

Despite dedicated research has been carried out to adequately map the distribution of the sperm whale in the Mediterranean Sea, unlike other regions of the world, the species population status is still presently uncertain. The analysis of two years of continuous acoustic data provided by the ANTARES neutrino telescope revealed the year-round presence of sperm whales in the Ligurian Sea, probably associated with the availability of cephalopods in the region. The presence of the Ligurian Sea sperm whales was demonstrated through the real-time analysis of audio data streamed from a cabled-to-shore deep-sea observatory that allowed the hourly tracking of their long-range echolocation behaviour on the Internet. Interestingly, the same acoustic analysis indicated that the occurrence of surface shipping noise would apparently not condition the foraging behaviour of the sperm whale in the area, since shipping noise was almost always present when sperm whales were acoustically detected. The continuous presence of the sperm whale in the region confirms the ecological value of the Ligurian sea and the importance of ANTARES to help monitoring its ecosystems. PMID:28401960

Bottlenose dolphins perceive object features through echolocation.

PubMed

Harley, Heidi E; Putman, Erika A; Roitblat, Herbert L

2003-08-07

How organisms (including people) recognize distant objects is a fundamental question. The correspondence between object characteristics (distal stimuli), like visual shape, and sensory characteristics (proximal stimuli), like retinal projection, is ambiguous. The view that sensory systems are 'designed' to 'pick up' ecologically useful information is vague about how such mechanisms might work. In echolocating dolphins, which are studied as models for object recognition sonar systems, the correspondence between echo characteristics and object characteristics is less clear. Many cognitive scientists assume that object characteristics are extracted from proximal stimuli, but evidence for this remains ambiguous. For example, a dolphin may store 'sound templates' in its brain and identify whole objects by listening for a particular sound. Alternatively, a dolphin's brain may contain algorithms, derived through natural endowments or experience or both, which allow it to identify object characteristics based on sounds. The standard method used to address this question in many species is indirect and has led to equivocal results with dolphins. Here we outline an appropriate method and test it to show that dolphins extract object characteristics directly from echoes.

The evaluation of marine traffic noise and its effect on the Chinese White Dolphins in Sha Chau and Lung Kwu Chau Marine Park, Hong Kong

NASA Astrophysics Data System (ADS)

Chan, T.

2016-12-01

The Hong Kong International Airport (HKIA) handles a large amount of air traffic, at 68.5 million passengers per year (approx. 40.1 million flights per year). Given that flight demand is forecast to reach 97 million passengers per year (approx 60.2 million flights per year) by 2030, the HKIA must optimise runway capacity in order to meet growing demand. Hence, the Airport Authority Hong Kong (AAHK) has put forward a proposal to expand the airport into a three-runway system by building a third runway. However, this presents many environmental constraints. These include major impacts on marine ecology; especially the Chinese White Dolphins (Sousa chinensis) that live around the Sha Chau and Lung Kwu Chau Marine Park north to the airport. Due to that the third runway will reclaim approximately 650 hectares of the se so re-routing of speed boats and ferries has been planned. This includes one route that passes through the marine nursery inside the park. This indirectly causes noise pollution that will affect marine life. Since dolphins rely on echolocation (often discerned by a "click" sound) to communicate and navigate, increasing marine traffic (re-routing) causes obscuring noises and acoustic communication interfering with echolocation, as well as collisions which lead to physical injuries. Underwater construction works involving techniques such as percussive piling may interfere with the dolphins' echolocation capability. The data has been collected using a hydrophone by observing the noise frequency changes within the marine park. The noises come from a number of sources, including fishing boats, and speed boats, krill and around 20 estuarine fish species. The evaluation of four years (2013-2016) has shown that noise pollution is increasing (as indicated by the increase of frequencies around 100 125 kHz) and that it has caused disruption in regular dolphin movement (irregular clicking sequence) .

Beaked whales respond to simulated and actual navy sonar.

PubMed

Tyack, Peter L; Zimmer, Walter M X; Moretti, David; Southall, Brandon L; Claridge, Diane E; Durban, John W; Clark, Christopher W; D'Amico, Angela; DiMarzio, Nancy; Jarvis, Susan; McCarthy, Elena; Morrissey, Ronald; Ward, Jessica; Boyd, Ian L

2011-03-14

Beaked whales have mass stranded during some naval sonar exercises, but the cause is unknown. They are difficult to sight but can reliably be detected by listening for echolocation clicks produced during deep foraging dives. Listening for these clicks, we documented Blainville's beaked whales, Mesoplodon densirostris, in a naval underwater range where sonars are in regular use near Andros Island, Bahamas. An array of bottom-mounted hydrophones can detect beaked whales when they click anywhere within the range. We used two complementary methods to investigate behavioral responses of beaked whales to sonar: an opportunistic approach that monitored whale responses to multi-day naval exercises involving tactical mid-frequency sonars, and an experimental approach using playbacks of simulated sonar and control sounds to whales tagged with a device that records sound, movement, and orientation. Here we show that in both exposure conditions beaked whales stopped echolocating during deep foraging dives and moved away. During actual sonar exercises, beaked whales were primarily detected near the periphery of the range, on average 16 km away from the sonar transmissions. Once the exercise stopped, beaked whales gradually filled in the center of the range over 2-3 days. A satellite tagged whale moved outside the range during an exercise, returning over 2-3 days post-exercise. The experimental approach used tags to measure acoustic exposure and behavioral reactions of beaked whales to one controlled exposure each of simulated military sonar, killer whale calls, and band-limited noise. The beaked whales reacted to these three sound playbacks at sound pressure levels below 142 dB re 1 µPa by stopping echolocation followed by unusually long and slow ascents from their foraging dives. The combined results indicate similar disruption of foraging behavior and avoidance by beaked whales in the two different contexts, at exposures well below those used by regulators to define disturbance.

Adaptive Evolution of the Myo6 Gene in Old World Fruit Bats (Family: Pteropodidae)

PubMed Central

Shen, Bin; Han, Xiuqun; Jones, Gareth; Rossiter, Stephen J.; Zhang, Shuyi

2013-01-01

Myosin VI (encoded by the Myo6 gene) is highly expressed in the inner and outer hair cells of the ear, retina, and polarized epithelial cells such as kidney proximal tubule cells and intestinal enterocytes. The Myo6 gene is thought to be involved in a wide range of physiological functions such as hearing, vision, and clathrin-mediated endocytosis. Bats (Chiroptera) represent one of the most fascinating mammal groups for molecular evolutionary studies of the Myo6 gene. A diversity of specialized adaptations occur among different bat lineages, such as echolocation and associated high-frequency hearing in laryngeal echolocating bats, large eyes and a strong dependence on vision in Old World fruit bats (Pteropodidae), and specialized high-carbohydrate but low-nitrogen diets in both Old World and New World fruit bats (Phyllostomidae). To investigate what role(s) the Myo6 gene might fulfill in bats, we sequenced the coding region of the Myo6 gene in 15 bat species and used molecular evolutionary analyses to detect evidence of positive selection in different bat lineages. We also conducted real-time PCR assays to explore the expression levels of Myo6 in a range of tissues from three representative bat species. Molecular evolutionary analyses revealed that the Myo6 gene, which was widely considered as a hearing gene, has undergone adaptive evolution in the Old World fruit bats which lack laryngeal echolocation and associated high-frequency hearing. Real-time PCR showed the highest expression level of the Myo6 gene in the kidney among ten tissues examined in three bat species, indicating an important role for this gene in kidney function. We suggest that Myo6 has undergone adaptive evolution in Old World fruit bats in relation to receptor-mediated endocytosis for the preservation of protein and essential nutrients. PMID:23620821

Amazon river dolphins (Inia geoffrensis) use a high-frequency short-range biosonar.

PubMed

Ladegaard, Michael; Jensen, Frants Havmand; de Freitas, Mafalda; Ferreira da Silva, Vera Maria; Madsen, Peter Teglberg

2015-10-01

Toothed whales produce echolocation clicks with source parameters related to body size; however, it may be equally important to consider the influence of habitat, as suggested by studies on echolocating bats. A few toothed whale species have fully adapted to river systems, where sonar operation is likely to result in higher clutter and reverberation levels than those experienced by most toothed whales at sea because of the shallow water and dense vegetation. To test the hypothesis that habitat shapes the evolution of toothed whale biosonar parameters by promoting simpler auditory scenes to interpret in acoustically complex habitats, echolocation clicks of wild Amazon river dolphins were recorded using a vertical seven-hydrophone array. We identified 404 on-axis biosonar clicks having a mean SLpp of 190.3 ± 6.1 dB re. 1 µPa, mean SLEFD of 132.1 ± 6.0 dB re. 1 µPa(2)s, mean Fc of 101.2 ± 10.5 kHz, mean BWRMS of 29.3 ± 4.3 kHz and mean ICI of 35.1 ± 17.9 ms. Piston fit modelling resulted in an estimated half-power beamwidth of 10.2 deg (95% CI: 9.6-10.5 deg) and directivity index of 25.2 dB (95% CI: 24.9-25.7 dB). These results support the hypothesis that river-dwelling toothed whales operate their biosonars at lower amplitude and higher sampling rates than similar-sized marine species without sacrificing high directivity, in order to provide high update rates in acoustically complex habitats and simplify auditory scenes through reduced clutter and reverberation levels. We conclude that habitat, along with body size, is an important evolutionary driver of source parameters in toothed whale biosonars. © 2015. Published by The Company of Biologists Ltd.

High levels of activity of bats at gold mining water bodies: implications for compliance with the International Cyanide Management Code.

PubMed

Griffiths, Stephen R; Donato, David B; Coulson, Graeme; Lumsden, Linda F

2014-06-01

Wildlife and livestock are known to visit and interact with tailings dam and other wastewater impoundments at gold mines. When cyanide concentrations within these water bodies exceed a critical toxicity threshold, significant cyanide-related mortality events can occur in wildlife. Highly mobile taxa such as birds are particularly susceptible to cyanide toxicosis. Nocturnally active bats have similar access to uncovered wastewater impoundments as birds; however, cyanide toxicosis risks to bats remain ambiguous. This study investigated activity of bats in the airspace above two water bodies at an Australian gold mine, to assess the extent to which bats use these water bodies and hence are at potential risk of exposure to cyanide. Bat activity was present on most nights sampled during the 16-month survey period, although it was highly variable across nights and months. Therefore, despite the artificial nature of wastewater impoundments at gold mines, these structures present attractive habitats to bats. As tailings slurry and supernatant pooling within the tailings dam were consistently well below the industry protective concentration limit of 50 mg/L weak acid dissociable (WAD) cyanide, wastewater solutions stored within the tailings dam posed a minimal risk of cyanide toxicosis for wildlife, including bats. This study showed that passively recorded bat echolocation call data provides evidence of the presence and relative activity of bats above water bodies at mine sites. Furthermore, echolocation buzz calls recorded in the airspace directly above water provide indirect evidence of foraging and/or drinking. Both echolocation monitoring and systematic sampling of cyanide concentration in open wastewater impoundments can be incorporated into a gold mine risk-assessment model in order to evaluate the risk of bat exposure to cyanide. In relation to risk minimisation management practices, the most effective mechanism for preventing cyanide toxicosis to wildlife, including bats, is capping the concentration of cyanide in tailings discharged to open impoundments at 50 mg/L WAD.

Beaked Whales Respond to Simulated and Actual Navy Sonar

PubMed Central

Tyack, Peter L.; Zimmer, Walter M. X.; Moretti, David; Southall, Brandon L.; Claridge, Diane E.; Durban, John W.; Clark, Christopher W.; D'Amico, Angela; DiMarzio, Nancy; Jarvis, Susan; McCarthy, Elena; Morrissey, Ronald; Ward, Jessica; Boyd, Ian L.

2011-01-01

Beaked whales have mass stranded during some naval sonar exercises, but the cause is unknown. They are difficult to sight but can reliably be detected by listening for echolocation clicks produced during deep foraging dives. Listening for these clicks, we documented Blainville's beaked whales, Mesoplodon densirostris, in a naval underwater range where sonars are in regular use near Andros Island, Bahamas. An array of bottom-mounted hydrophones can detect beaked whales when they click anywhere within the range. We used two complementary methods to investigate behavioral responses of beaked whales to sonar: an opportunistic approach that monitored whale responses to multi-day naval exercises involving tactical mid-frequency sonars, and an experimental approach using playbacks of simulated sonar and control sounds to whales tagged with a device that records sound, movement, and orientation. Here we show that in both exposure conditions beaked whales stopped echolocating during deep foraging dives and moved away. During actual sonar exercises, beaked whales were primarily detected near the periphery of the range, on average 16 km away from the sonar transmissions. Once the exercise stopped, beaked whales gradually filled in the center of the range over 2–3 days. A satellite tagged whale moved outside the range during an exercise, returning over 2–3 days post-exercise. The experimental approach used tags to measure acoustic exposure and behavioral reactions of beaked whales to one controlled exposure each of simulated military sonar, killer whale calls, and band-limited noise. The beaked whales reacted to these three sound playbacks at sound pressure levels below 142 dB re 1 µPa by stopping echolocation followed by unusually long and slow ascents from their foraging dives. The combined results indicate similar disruption of foraging behavior and avoidance by beaked whales in the two different contexts, at exposures well below those used by regulators to define disturbance. PMID:21423729

Effects of Orientation and Weatherproofing on the Detection of Bat Echolocation Calls

Treesearch

E. Britzke; B. Slack; M Armstrong; S. Loeb

2010-01-01

Ultrasonic detectors are powerful tools for the study of bat ecology. Many options are available for deploying acoustic detectors including various weatherproofing designs and microphone orientations, but the impacts of these options on the quantity and quality of the bat calls that are recorded are unknown. We compared the impacts of three microphone orientations (...

Proceedings of the International Conference on Sensory Devices for the Blind (London, England, June, 1966).

ERIC Educational Resources Information Center

Dufton, Richard, Ed.

The conference proceedings include papers on sensory aids for visually handicapped mobility and reading. Two papers each treat mobility as a general problem, sociocultural surveys on mobility and reading, and echolocation in man and bats. Five papers concern reports and evaluations of practical trials of the sonic monaural aid; one deals with…

Third International Conference on Acoustic Communication by Animals

DTIC Science & Technology

2011-09-30

communications Invited Speakers Peter Tyack cetacean communications Christopher Clark acoustic environment of whales Whitlow Au sound detection and...echolocation by dolphins Magnus Wahlberg sperm whale acoustics Robert Dooling bird hearing Ronald Hoy communication strategies in insects Peter Narins...frogs (6). Topics covered included cognition/language; song and call classification; rule learning; acoustic ecology; communication in noisy

Activation of the Hippocampal Complex during Tactile Maze Solving in Congenitally Blind Subjects

ERIC Educational Resources Information Center

Gagnon, Lea; Schneider, Fabien C.; Siebner, Hartwig R.; Paulson, Olaf B.; Kupers, Ron; Ptito, Maurice

2012-01-01

Despite their lack of vision, congenitally blind subjects are able to build and manipulate cognitive maps for spatial navigation. It is assumed that they thereby rely more heavily on echolocation, proprioceptive signals and environmental cues such as ambient temperature and audition to compensate for their lack of vision. Little is known, however,…

77 FR 19231 - Takes of Marine Mammals Incidental to Specified Activities; Navy Training Conducted at the Silver...

Federal Register 2010, 2011, 2012, 2013, 2014

2012-03-30

... break, presence of glassy foam, and/or perhaps scattered whitecaps). Applicable mitigation zones are... California based on passive acoustic detection of two distinct echolocation click patterns. No population... 10 1,020 1,220 (2) ZOI and Swim Speed-Time-Buffer Addition Based on acoustic propagation modeling and...

Development and Functions of Signature Whistles of Free-Ranging Bottlenose Dolphins, Tursiops truncatus

DTIC Science & Technology

1992-09-01

34 (Waser 1977), "loud calls" of lion tailed macaques (Green 1981), long calls of orangutans (Galdikas and Insley 1988), and gorilla "roars" (Marler and...whistles and echolocation clicks. Galdikas, B. M. F. and S. J. Insley. 1988. The fast call of the adult male orangutan . J. Mamm. 69: 371-375. Gould, E

Divergence of dim-light vision among bats (order: Chiroptera) as estimated by molecular and electrophysiological methods.

PubMed

Liu, He-Qun; Wei, Jing-Kuan; Li, Bo; Wang, Ming-Shan; Wu, Rui-Qi; Rizak, Joshua D; Zhong, Li; Wang, Lu; Xu, Fu-Qiang; Shen, Yong-Yi; Hu, Xin-Tian; Zhang, Ya-Ping

2015-06-23

Dim-light vision is present in all bats, but is divergent among species. Old-World fruit bats (Pteropodidae) have fully developed eyes; the eyes of insectivorous bats are generally degraded, and these bats rely on well-developed echolocation. An exception is the Emballonuridae, which are capable of laryngeal echolocation but prefer to use vision for navigation and have normal eyes. In this study, integrated methods, comprising manganese-enhanced magnetic resonance imaging (MEMRI), f-VEP and RNA-seq, were utilized to verify the divergence. The results of MEMRI showed that Pteropodidae bats have a much larger superior colliculus (SC)/ inferior colliculus (IC) volume ratio (3:1) than insectivorous bats (1:7). Furthermore, the absolute visual thresholds (log cd/m(2)•s) of Pteropodidae (-6.30 and -6.37) and Emballonuridae (-3.71) bats were lower than those of other insectivorous bats (-1.90). Finally, genes related to the visual pathway showed signs of positive selection, convergent evolution, upregulation and similar gene expression patterns in Pteropodidae and Emballonuridae bats. Different results imply that Pteropodidae and Emballonuridae bats have more developed vision than the insectivorous bats and suggest that further research on bat behavior is warranted.

Propagation of narrow-band-high-frequency clicks: measured and modeled transmission loss of porpoise-like clicks in porpoise habitats.

PubMed

DeRuiter, Stacy L; Hansen, Michael; Koopman, Heather N; Westgate, Andrew J; Tyack, Peter L; Madsen, Peter T

2010-01-01

Estimating the range at which harbor porpoises can detect prey items and environmental objects is integral to understanding their biosonar. Understanding the ranges at which they can use echolocation to detect and avoid obstacles is particularly important for strategies to reduce bycatch. Transmission loss (TL) during acoustic propagation is an important determinant of those detection ranges, and it also influences animal detection functions used in passive acoustic monitoring. However, common assumptions regarding TL have rarely been tested. Here, TL of synthetic porpoise clicks was measured in porpoise habitats in Canada and Denmark, and field data were compared with spherical spreading law and ray-trace (Bellhop) model predictions. Both models matched mean observations quite well in most cases, indicating that a spherical spreading law can usually provide an accurate first-order estimate of TL for porpoise sounds in porpoise habitat. However, TL varied significantly (+/-10 dB) between sites and over time in response to variability in seafloor characteristics, sound-speed profiles, and other short-timescale environmental fluctuations. Such variability should be taken into account in estimates of the ranges at which porpoises can communicate acoustically, detect echolocation targets, and be detected via passive acoustic monitoring.

Forward-masking based gain control in odontocete biosonar: an evoked-potential study.

PubMed

Supin, Alexander Ya; Nachtigall, Paul E; Breese, Marlee

2009-04-01

Auditory evoked potentials (AEPs) were recorded during echolocation in a false killer whale Pseudorca crassidens. An electronically synthesized and played-back ("phantom") echo was used. Each electronic echo was triggered by an emitted biosonar pulse. The echo had a spectrum similar to that of the emitted biosonar clicks, and its intensity was proportional to that of the emitted click. The attenuation of the echo relative to the emitted click and its delay was controlled by the experimenter. Four combinations of echo attenuation and delay were tested (-31 dB, 2 ms), (-40 dB, 4 ms), (-49 dB, 8 ms), and (-58 dB, 16 ms); thus, attenuation and delay were associated with a rate of 9 dB of increased attenuation per delay doubling. AEPs related to emitted clicks displayed a regular amplitude dependence on the click level. Echo-related AEPs did not feature amplitude dependence on echo attenuation or emitted click levels, except in a few combinations of the lowest values of these two variables. The results are explained by a hypothesis that partial forward masking of the echoes by the preceding emitted sonar pulses serves as a kind of automatic gain control in the auditory system of echolocating odontocetes.

Source-to-sensation level ratio of transmitted biosonar pulses in an echolocating false killer whale.

PubMed

Supin, Alexander Ya; Nachtigall, Paul E; Breese, Marlee

2006-07-01

Transmitted biosonar pulses, and the brain auditory evoked potentials (AEPs) associated with those pulses, were synchronously recorded in a false killer whale Pseudorca crassidens trained to accept suction-cup EEG electrodes and to detect targets by echolocation. AEP amplitude was investigated as a function of the transmitted biosonar pulse source level. For that, a few thousand of the individual AEP records were sorted according to the spontaneously varied amplitude of synchronously recorded biosonar pulses. In each of the sorting bins (in 5-dB steps) AEP records were averaged to extract AEP from noise; AEP amplitude was plotted as a function of the biosonar pulse source level. For comparison, AEPs were recorded to external (in free field) sound pulses of a waveform and spectrum similar to those of the biosonar pulses; amplitude of these AEPs was plotted as a function of sound pressure level. A comparison of these two functions has shown that, depending on the presence or absence of a target, the sensitivity of the whale's hearing to its own transmitted biosonar pulses was 30 to 45 dB lower than might be expected in a free acoustic field.

Stereotypical rapid source level regulation in the harbour porpoise biosonar.

PubMed

Linnenschmidt, Meike; Kloepper, Laura N; Wahlberg, Magnus; Nachtigall, Paul E

2012-09-01

Some odontocetes and bats vary both click intensity and receiver sensitivity during echolocation, depending on target range. It is not known how this so-called automatic gain control is regulated by the animal. The source level of consecutive echolocation clicks from a harbour porpoise was measured with a hydrophone array while the animal detected an aluminium cylinder at 2, 4 or 8 m distance in a go/no-go paradigm. On-axis clicks had source levels of 145-174 dB re 1 μPa peak-to-peak. During target-present trials the click trains reached comparable source levels independent of the range to the target after three clicks. After an additional click, the source level was reduced for the 2 and 4 m trials until it equalled the one-way transmission loss. During target-absent trials, the source level remained high throughout the entire click train. Given typical values of harbour porpoise inter-click intervals, the source level reduction commenced within a few 100 ms from the first click in the click train. This may indicate a sub-cortically regulated source level regulation in the harbour porpoise.

Trackline and point detection probabilities for acoustic surveys of Cuvier's and Blainville's beaked whales.

PubMed

Barlow, Jay; Tyack, Peter L; Johnson, Mark P; Baird, Robin W; Schorr, Gregory S; Andrews, Russel D; Aguilar de Soto, Natacha

2013-09-01

Acoustic survey methods can be used to estimate density and abundance using sounds produced by cetaceans and detected using hydrophones if the probability of detection can be estimated. For passive acoustic surveys, probability of detection at zero horizontal distance from a sensor, commonly called g(0), depends on the temporal patterns of vocalizations. Methods to estimate g(0) are developed based on the assumption that a beaked whale will be detected if it is producing regular echolocation clicks directly under or above a hydrophone. Data from acoustic recording tags placed on two species of beaked whales (Cuvier's beaked whale-Ziphius cavirostris and Blainville's beaked whale-Mesoplodon densirostris) are used to directly estimate the percentage of time they produce echolocation clicks. A model of vocal behavior for these species as a function of their diving behavior is applied to other types of dive data (from time-depth recorders and time-depth-transmitting satellite tags) to indirectly determine g(0) in other locations for low ambient noise conditions. Estimates of g(0) for a single instant in time are 0.28 [standard deviation (s.d.) = 0.05] for Cuvier's beaked whale and 0.19 (s.d. = 0.01) for Blainville's beaked whale.

Precocious hearing in harbour porpoise neonates.

PubMed

Wahlberg, Magnus; Delgado-García, Lara; Kristensen, Jakob H

2017-02-01

Hearing is the primary sensory modality for toothed whales, but it is not known at which age it is fully developed. For newborn calves, hearing could fill an important function in maintaining contact with the mother and to develop echolocation skills. We non-invasively measured the auditory brainstem response (ABR) in two neonate (age 1-4 days) and three adult harbour porpoises (Phocoena phocoena). The stimuli consisted of clicks centred at 130 kHz, which is within the frequency band used for echolocation and communication in this species. The temporal pattern of the neonate ABRs was indistinguishable to the adult ones. There were no significant differences between calves and adults regarding hearing thresholds and ABR latencies. The ABR amplitudes were up to more than an order of magnitude larger in newborns than in adults, most likely due to the neonates' smaller size. These results indicate that hearing is fully developed within a day after birth, which suggests that harbour porpoise neonates have the earliest hearing development of any mammal studied so far. This may be explained by the evolutionary pressures imposed by the aquatic environment for a rapid development of the key sensory system in harbour porpoises.

The role of tragus on echolocating bat, Eptesicus fuscus

NASA Astrophysics Data System (ADS)

Chiu, Chen; Moss, Cynthia

2005-04-01

Echolocating bats produce ultrasonic vocal signals and utilize the returning echoes to detect, localize and track prey, and also to avoid obstacles. The pinna and tragus, two major components of the bats external ears, play important roles in filtering returning echoes. The tragus is generally believed to play a role in vertical sound localization. The purpose of this study is to further examine how manipulation of the tragus affects a free-flying bat's prey capture and obstacle avoidance behavior. The first part of this study involved a prey capture experiment, and the bat was trained to catch the tethered mealworms in a large room. The second experiment involved obstacle avoidance, and the bat's task was to fly through the largest opening from a horizontal wire array without touching the wires. In both experiments, the bat performed the tasks under three different conditions: with intact tragus, tragus-deflection and recovery from tragus-deflection. Significantly lower performance was observed in both experiments when tragi were glued down. However, the bat adjusted quickly and returned to baseline performance a few days after the manipulation. The results suggest that tragus-deflection does have effects on both the prey capture and obstacle avoidance behavior. [Work supported by NSF.

Auditory fovea and Doppler shift compensation: adaptations for flutter detection in echolocating bats using CF-FM signals.

PubMed

Schnitzler, Hans-Ulrich; Denzinger, Annette

2011-05-01

Rhythmical modulations in insect echoes caused by the moving wings of fluttering insects are behaviourally relevant information for bats emitting CF-FM signals with a high duty cycle. Transmitter and receiver of the echolocation system in flutter detecting foragers are especially adapted for the processing of flutter information. The adaptations of the transmitter are indicated by a flutter induced increase in duty cycle, and by Doppler shift compensation (DSC) that keeps the carrier frequency of the insect echoes near a reference frequency. An adaptation of the receiver is the auditory fovea on the basilar membrane, a highly expanded frequency representation centred to the reference frequency. The afferent projections from the fovea lead to foveal areas with an overrepresentation of sharply tuned neurons with best frequencies near the reference frequency throughout the entire auditory pathway. These foveal neurons are very sensitive to stimuli with natural and simulated flutter information. The frequency range of the foveal areas with their flutter processing neurons overlaps exactly with the frequency range where DS compensating bats most likely receive echoes from fluttering insects. This tight match indicates that auditory fovea and DSC are adaptations for the detection and evaluation of insects flying in clutter.

Echolocation in Blainville's beaked whales (Mesoplodon densirostris).

PubMed

Madsen, P T; de Soto, N Aguilar; Arranz, P; Johnson, M

2013-06-01

Here we use sound and movement recording tags to study how deep-diving Blainville's beaked whales (Mesoplodon densirostris) use echolocation to forage in their natural mesopelagic habitat. These whales ensonify thousands of organisms per dive but select only about 25 prey for capture. They negotiate their cluttered environment by radiating sound in a narrow 20° field of view which they sample with 1.5-3 clicks per metre travelled requiring only some 60 clicks to locate, select and approach each prey. Sampling rates do not appear to be defined by the range to individual targets, but rather by the movement of the predator. Whales sample faster when they encounter patches of prey allowing them to search new water volumes while turning rapidly to stay within a patch. This implies that the Griffin search-approach-capture model of biosonar foraging must be expanded to account for sampling behaviours adapted to the overall prey distribution. Beaked whales can classify prey at more than 15 m range adopting stereotyped motor patterns when approaching some prey. This long detection range relative to swimming speed facilitates a deliberate mode of sensory-motor operation in which prey and capture tactics can be selected to optimize energy returns during long breath-hold dives.

Acoustic basis for recognition of aspect-dependent three-dimensional targets by an echolocating bottlenose dolphin.

PubMed

Helweg, D A; Au, W W; Roitblat, H L; Nachtigall, P E

1996-04-01

The relationships between acoustic features of target echoes and the cognitive representations of the target formed by an echolocating dolphin will influence the ease with which the dolphin can recognize a target. A blindfolded Atlantic bottlenose dolphin (Tursiops truncatus) learned to match aspect-dependent three-dimensional targets (such as a cube) at haphazard orientations, although with some difficulty. This task may have been difficult because aspect-dependent targets produce different echoes at different orientations, which required the dolphin to have some capability for object constancy across changes in echo characteristics. Significant target-related differences in echo amplitude, rms bandwidth, and distributions of interhighlight intervals were observed among echoes collected when the dolphin was performing the task. Targets could be classified using a combination of energy flux density and rms bandwidth by a linear discriminant analysis and a nearest centroid classifier. Neither statistical model could classify targets without amplitude information, but the highest accuracy required spectral information as well. This suggests that the dolphin recognized the targets using a multidimensional representation containing amplitude and spectral information and that dolphins can form stable representations of targets regardless of orientation based on varying sensory properties.

PRINCIPLES AND PATTERNS OF BAT MOVEMENTS: FROM AERODYNAMICS TO ECOLOGY

PubMed Central

Voigt, Christian C.; Frick, Winifred F.; Holderied, Marc W.; Holland, Richard; Kerth, Gerald; Mello, Marco A. R.; Plowright, Raina K.; Swartz, Sharon; Yovel, Yossi

2018-01-01

Movement ecology as an integrative discipline has advanced associated fields because it presents not only a conceptual framework for understanding movement principles but also helps formulate predictions about the consequences of movements for animals and their environments. Here, we synthesize recent studies on principles and patterns of bat movements in context of the movement ecology paradigm. The motion capacity of bats is defined by their highly articulated, flexible wings. Power production during flight follows a U-shaped curve in relation to speed in bats yet, in contrast to birds, bats use mostly exogenous nutrients for sustained flight. The navigation capacity of most bats is dominated by the echolocation system, yet other sensory modalities, including an iron-based magnetic sense, may contribute to navigation depending on a bat’s familiarity with the terrain. Patterns derived from these capacities relate to antagonistic and mutualistic interactions with food items. The navigation capacity of bats may influence their sociality, in particular, the extent of group foraging based on eavesdropping on conspecifics’ echolocation calls. We infer that understanding the movement ecology of bats within the framework of the movement ecology paradigm provides new insights into ecological processes mediated by bats, from ecosystem services to diseases. PMID:29861509

Remote sensing of chemical warfare agent by CO2 -lidar

NASA Astrophysics Data System (ADS)

Geiko, Pavel P.; Smirnov, Sergey S.

2014-11-01

The possibilities of remote sensing of chemical warfare agent by differential absorption method were analyzed. The CO2 - laser emission lines suitable for sounding of chemical warfare agent with provision for disturbing absorptions by water vapor were choose. The detection range of chemical warfare agents was estimated for a lidar based on CO2 - laser The other factors influencing upon echolocation range were analyzed.

Unsupervised Learning (Clustering) of Odontocete Echolocation Clicks

DTIC Science & Technology

2015-09-30

of their bandwidth. Results on Risso’s dolphins (Grampus griseus), Pacific white-sided dolphins (Lagenorhynchus obliquidens), and Cuvier’s beaked...acoustic encounters to see which ones appeared to be closely related to one another. We noted that some of the Pacific white-sided and Risso’s dolphin ...should be clusterable. The group of odontocetes that we cannot label reliably by their acoustic features, primarily common dolphins (Delphinus spp

Do you hear what I see? Vocalization relative to visual detection rates of Hawaiian hoary bats (Lasiurus cinereus semotus)

USGS Publications Warehouse

Gorresen, Paulo Marcos; Cryan, Paul; Montoya-Aiona, Kristina; Bonaccorso, Frank

2017-01-01

Bats vocalize during flight as part of the sensory modality called echolocation, but very little is known about whether flying bats consistently call. Occasional vocal silence during flight when bats approach prey or conspecifics has been documented for relatively few species and situations. Bats flying alone in clutter-free airspace are not known to forgo vocalization, yet prior observations suggested possible silent behavior in certain, unexpected situations. Determining when, why, and where silent behavior occurs in bats will help evaluate major assumptions of a primary monitoring method for bats used in ecological research, management, and conservation. In this study, we recorded flight activity of Hawaiian hoary bats (Lasiurus cinereus semotus) under seminatural conditions using both thermal video cameras and acoustic detectors. Simultaneous video and audio recordings from 20 nights of observation at 10 sites were analyzed for correspondence between detection methods, with a focus on video observations in three distance categories for which accompanying vocalizations were detected. Comparison of video and audio detections revealed that a high proportion of Hawaiian hoary bats “seen” on video were not simultaneously “heard.” On average, only about one in three visual detections within a night had an accompanying call detection, but this varied greatly among nights. Bats flying on curved flight paths and individuals nearer the cameras were more likely to be detected by both methods. Feeding and social calls were detected, but no clear pattern emerged from the small number of observations involving closely interacting bats. These results may indicate that flying Hawaiian hoary bats often forgo echolocation, or do not always vocalize in a way that is detectable with common sampling and monitoring methods. Possible reasons for the low correspondence between visual and acoustic detections range from methodological to biological and include a number of biases associated with the propagation and detection of sound, cryptic foraging strategies, or conspecific presence. Silent flight behavior may be more prevalent in echolocating bats than previously appreciated, has profound implications for ecological research, and deserves further characterization and study.

Topography of sound level representation in the FM sweep selective region of the pallid bat auditory cortex.

PubMed

Measor, Kevin; Yarrow, Stuart; Razak, Khaleel A

2018-05-26

Sound level processing is a fundamental function of the auditory system. To determine how the cortex represents sound level, it is important to quantify how changes in level alter the spatiotemporal structure of cortical ensemble activity. This is particularly true for echolocating bats that have control over, and often rapidly adjust, call level to actively change echo level. To understand how cortical activity may change with sound level, here we mapped response rate and latency changes with sound level in the auditory cortex of the pallid bat. The pallid bat uses a 60-30 kHz downward frequency modulated (FM) sweep for echolocation. Neurons tuned to frequencies between 30 and 70 kHz in the auditory cortex are selective for the properties of FM sweeps used in echolocation forming the FM sweep selective region (FMSR). The FMSR is strongly selective for sound level between 30 and 50 dB SPL. Here we mapped the topography of level selectivity in the FMSR using downward FM sweeps and show that neurons with more monotonic rate level functions are located in caudomedial regions of the FMSR overlapping with high frequency (50-60 kHz) neurons. Non-monotonic neurons dominate the FMSR, and are distributed across the entire region, but there is no evidence for amplitopy. We also examined how first spike latency of FMSR neurons change with sound level. The majority of FMSR neurons exhibit paradoxical latency shift wherein the latency increases with sound level. Moreover, neurons with paradoxical latency shifts are more strongly level selective and are tuned to lower sound level than neurons in which latencies decrease with level. These data indicate a clustered arrangement of neurons according to monotonicity, with no strong evidence for finer scale topography, in the FMSR. The latency analysis suggests mechanisms for strong level selectivity that is based on relative timing of excitatory and inhibitory inputs. Taken together, these data suggest how the spatiotemporal spread of cortical activity may represent sound level. Copyright © 2018. Published by Elsevier B.V.

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

PubMed

Ghose, Kaushik; Horiuchi, Timothy K; Krishnaprasad, P S; Moss, Cynthia F

2006-05-01

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.

Building a Virtual Model of a Baleen Whale: Phase 2

DTIC Science & Technology

2015-09-30

example, the biosonar signal generation mechanism and the formation of an acoustic transmission beam, or to measure the amplitude differences and...from simulations with published results from live animal biosonar research (Cranford and Krysl, 2013). Collectively, our research results butress the...2004). " Biosonar pulse production in odontocetes: The state of our knowledge," in Echolocation in bats and dolphins, edited by J. A. Thomas, C. F

Control of Biosonar Behavior by the Auditory Cortex

DTIC Science & Technology

1988-11-28

TITLE (include Security Classification) Control of Biosonar Behavior by the Auditory Cortex 12. PERSONAL AUTHOR(S) Nobuo Suga and Stephen Gaioni 13a...NOTATION 17. COSATI CODES IS SUBJECT TERMS (Continue on reverse if necessary and identify by block number) FIELD GROUP1 SUB-GROUP - biosonar ; echolocation...SLesion experiments were conducted to examine whether the functional organization of the mustached bat’s auditory cortex is related to biosonar

Building a Virtual Model of a Baleen Whale: Phase 2

DTIC Science & Technology

2013-09-30

valid by comparing results gleaned from live animals involved in biosonar tasks (Cranford and Krysl, 2013) and have begun the process of...Cranford, T. W., and Amundin, M. E. (2003). " Biosonar pulse production in odontocetes: The state of our knowledge," in Echolocation in bats and...Krysl, P. (2014). "Validation of a vibroacoustic finite element model using bottlenose dolphin simulations: The dolphin biosonar beam is focused in

The Metabolic Cost of Click Production in Bottlenose Dolphins

DTIC Science & Technology

2014-09-30

America 64, 411-422. Holt, M.M., Noren, D.P., Veirs, V., Emmons, C.K., Veirs, S. 2009. Speaking up: Killer whales (Orcinus orca) increase their call...2007. Recording and quantification of ultrasonic echolocation clicks from free-ranging toothed whales . Deep-Sea Research I 54, 1421-1444. 15...Individual right whales call louder in increased environmental noise. Biology Letters published online 7 July 2010 doi: 10.1098/rsbl.2010.0451

Advanced Methods for Passive Acoustic Detection, Classification, and Localization of Marine Mammals

DTIC Science & Technology

2012-09-30

floor 1176 Howell St Newport RI 02842 phone: (401) 832-5749 fax: (401) 832-4441 email: David.Moretti@navy.mil Steve W. Martin SPAWAR...multiclass support vector machine (SVM) classifier was previously developed ( Jarvis et al. 2008). This classifier both detects and classifies echolocation...whales. Here Moretti’s group, especially S. Jarvis , will improve the SVM classifier by resolving confusion between species whose clicks overlap in

Acoustic divergence in two cryptic Hipposideros species: a role for social selection?

PubMed Central

Kingston, T.; Lara, M. C.; Jones, G.; Akbar, Z.; Kunz, T. H.; Schneider, C. J.

2001-01-01

We present evidence that a relatively widespread and common bat from South East Asia comprises two morphologically cryptic but acoustically divergent species. A population of the bicoloured leaf-nosed bat (Hipposideros bicolor) from Peninsular Malaysia exhibits a bimodal distribution of echolocation call frequencies, with peaks in the frequency of maximum energy at ca. 131 and 142 kHz. The two phonic types are genetically distinct, with a cytochrome b sequence divergence of just under 7%. We consider the mechanisms by which acoustic divergence in these species might arise. Differences in call frequency are not likely to effect resource partitioning by detectable prey size or functional range. However, ecological segregation may be achieved by differences in microhabitat use; the 131kHz H. bicolor is characterized by significantly longer forearms, lower wing loading, a lower aspect ratio and a more rounded wingtip, features that are associated with greater manoeuvrability in flight that may enable it to forage in more cluttered environments relative to the 142 kHz phonic type. We suggest that acoustic divergence in these species is a consequence of social selection for a clear communication channel, which is mediated by the close link between the acoustic signal and receptor systems imposed by the highly specialized nature of the hipposiderid and rhinolophid echolocation system. PMID:11429138

Evolution of cranial telescoping in echolocating whales (Cetacea: Odontoceti).

PubMed

Churchill, Morgan; Geisler, Jonathan H; Beatty, Brian L; Goswami, Anjali

2018-05-01

Odontocete (echolocating whale) skulls exhibit extreme posterior displacement and overlapping of facial bones, here referred to as retrograde cranial telescoping. To examine retrograde cranial telescoping across 40 million years of whale evolution, we collected 3D scans of whale skulls spanning odontocete evolution. We used a sliding semilandmark morphometric approach with Procrustes superimposition and PCA to capture and describe the morphological variation present in the facial region, followed by Ancestral Character State Reconstruction (ACSR) and evolutionary model fitting on significant components to determine how retrograde cranial telescoping evolved. The first PC score explains the majority of variation associated with telescoping and reflects the posterior migration of the external nares and premaxilla alongside expansion of the maxilla and frontal. The earliest diverging fossil odontocetes were found to exhibit a lesser degree of cranial telescoping than later diverging but contemporary whale taxa. Major shifts in PC scores and centroid size are identified at the base of Odontoceti, and early burst and punctuated equilibrium models best fit the evolution of retrograde telescoping. This indicates that the Oligocene was a period of unusually high diversity and evolution in whale skull morphology, with little subsequent evolution in telescoping. © 2018 The Author(s). Evolution © 2018 The Society for the Study of Evolution.

Can the elongated hindwing tails of fluttering moths serve as false sonar targets to divert bat attacks?

PubMed

Lee, Wu-Jung; Moss, Cynthia F

2016-05-01

It has long been postulated that the elongated hindwing tails of many saturniid moths have evolved to create false sonar targets to divert the attack of echolocation-guided bat predators. However, rigorous echo-acoustic evidence to support this hypothesis has been lacking. In this study, fluttering luna moths (Actias luna), a species with elongated hindwing tails, were ensonified with frequency modulated chirp signals from all angles of orientation and across the wingbeat cycle. High-speed stereo videography was combined with pulse compression sonar processing to characterize the echo information available to foraging bats. Contrary to previous suggestions, the results show that the tail echoes are weak and do not dominate the sonar returns, compared to the large, planar wings and the moth body. However, the distinctive twisted morphology of the tails create persistent echoes across all angles of orientation, which may induce erroneous sonar target localization and disrupt accurate tracking by echolocating bats. These findings thus suggest a refinement of the false target hypothesis to emphasize sonar localization errors induced by the twisted tails, and highlight the importance of physics-based approaches to study the sensory information involved in the evolutionary arms race between moths and their bat predators.

Big brown bats (Eptesicus fuscus) reveal diverse strategies for sonar target tracking in clutter.

PubMed

Mao, Beatrice; Aytekin, Murat; Wilkinson, Gerald S; Moss, Cynthia F

2016-09-01

Bats actively adjust the acoustic features of their sonar calls to control echo information specific to a given task and environment. A previous study investigated how bats adapted their echolocation behavior when tracking a moving target in the presence of a stationary distracter at different distances and angular offsets. The use of only one distracter, however, left open the possibility that a bat could reduce the interference of the distracter by turning its head. Here, bats tracked a moving target in the presence of one or two symmetrically placed distracters to investigate adaptive echolocation behavior in a situation where vocalizing off-axis would result in increased interference from distracter echoes. Both bats reduced bandwidth and duration but increased sweep rate in more challenging distracter conditions, and surprisingly, made more head turns in the two-distracter condition compared to one, but only when distracters were placed at large angular offsets. However, for most variables examined, subjects showed distinct strategies to reduce clutter interference, either by (1) changing spectral or temporal features of their calls, or (2) producing large numbers of sonar sound groups and consistent head-turning behavior. The results suggest that individual bats can use different strategies for target tracking in cluttered environments.

Clutter and conspecifics: a comparison of their influence on echolocation and flight behaviour in Daubenton's bat, Myotis daubentonii.

PubMed

Fawcett, Kayleigh; Ratcliffe, John M

2015-03-01

We compared the influence of conspecifics and clutter on echolocation and flight speed in the bat Myotis daubentonii. In a large room, actual pairs of bats exhibited greater disparity in peak frequency (PF), minimum frequency (F MIN) and call period compared to virtual pairs of bats, each flying alone. Greater inter-individual disparity in PF and F MIN may reduce acoustic interference and/or increase signal self-recognition in the presence of conspecifics. Bats flying alone in a smaller flight room, to simulate a more cluttered habitat as compared to the large flight room, produced calls of shorter duration and call period, lower intensity, and flew at lower speeds. In cluttered space, shorter call duration should reduce masking, while shorter call period equals more updates to the bat's auditory scene. Lower intensity likely reflects reduced range detection requirements, reduced speed the demands of flying in clutter. Our results show that some changes (e.g. PF separation) are associated with conspecifics, others with closed habitat (e.g. reduced call intensity). However, we demonstrate that call duration, period, and flight speed appear similarly influenced by conspecifics and clutter. We suggest that some changes reduce conspecific interference and/or improve self-recognition, while others demonstrate that bats experience each other like clutter.

The Source Parameters of Echolocation Clicks from Captive and Free-Ranging Yangtze Finless Porpoises (Neophocaena asiaeorientalis asiaeorientalis).

PubMed

Fang, Liang; Wang, Ding; Li, Yongtao; Cheng, Zhaolong; Pine, Matthew K; Wang, Kexiong; Li, Songhai

2015-01-01

The clicks of Yangtze finless porpoises (Neophocaena asiaeorientalis asiaeorientalis) from 7 individuals in the tank of Baiji aquarium, 2 individuals in a netted pen at Shishou Tian-e-zhou Reserve and 4 free-ranging individuals at Tianxingzhou were recorded using a broadband digital recording system with four element hydrophones. The peak-to-peak apparent source level (ASL_pp) of clicks from individuals at the Baiji aquarium was 167 dB re 1 μPa with mean center frequency of 133 kHz, -3dB bandwidth of 18 kHz and -10 dB duration of 58 μs. The ASL_pp of clicks from individuals at the Shishou Tian-e-zhou Reserve was 180 dB re 1 μPa with mean center frequency of 128 kHz, -3dB bandwidth of 20 kHz and -10 dB duration of 39 μs. The ASL_pp of clicks from individuals at Tianxingzhou was 176 dB re 1 μPa with mean center frequency of 129 kHz, -3dB bandwidth of 15 kHz and -10 dB duration of 48 μs. Differences between the source parameters of clicks among the three groups of finless porpoises suggest these animals adapt to their echolocation signals depending on their surroundings.

Divergence of dim-light vision among bats (order: Chiroptera) as estimated by molecular and electrophysiological methods

PubMed Central

Liu, He-Qun; Wei, Jing-Kuan; Li, Bo; Wang, Ming-Shan; Wu, Rui-Qi; Rizak, Joshua D.; Zhong, Li; Wang, Lu; Xu, Fu-Qiang; Shen, Yong-Yi; Hu, Xin-Tian; Zhang, Ya-Ping

2015-01-01

Dim-light vision is present in all bats, but is divergent among species. Old-World fruit bats (Pteropodidae) have fully developed eyes; the eyes of insectivorous bats are generally degraded, and these bats rely on well-developed echolocation. An exception is the Emballonuridae, which are capable of laryngeal echolocation but prefer to use vision for navigation and have normal eyes. In this study, integrated methods, comprising manganese-enhanced magnetic resonance imaging (MEMRI), f-VEP and RNA-seq, were utilized to verify the divergence. The results of MEMRI showed that Pteropodidae bats have a much larger superior colliculus (SC)/ inferior colliculus (IC) volume ratio (3:1) than insectivorous bats (1:7). Furthermore, the absolute visual thresholds (log cd/m2•s) of Pteropodidae (−6.30 and −6.37) and Emballonuridae (−3.71) bats were lower than those of other insectivorous bats (−1.90). Finally, genes related to the visual pathway showed signs of positive selection, convergent evolution, upregulation and similar gene expression patterns in Pteropodidae and Emballonuridae bats. Different results imply that Pteropodidae and Emballonuridae bats have more developed vision than the insectivorous bats and suggest that further research on bat behavior is warranted. PMID:26100095

Interaction of emitted sonar pulses and simulated echoes in a false killer whale: an evoked-potential study.

PubMed

Supin, Alexander Ya; Nachtigall, Paul E; Breese, Marlee

2011-09-01

Auditory evoked potentials (AEP) were recorded during echolocation in a false killer whale Pseudorca crassidens. An electronically synthesized and played-back (simulated) echo was triggered by an emitted biosonar pulse, and its intensity was proportional to that of the emitted click. The delay and transfer factor of the echo relative to the emitted click was controlled by the operator. The echo delay varied from 2 to 16 ms (by two-fold steps), and the transfer factor varied within ranges from -45 to -30 dB at the 2-ms delay to -60 to -45 dB at the 16-ms delay. Echo-related AEPs featured amplitude dependence both on echo delay at a constant transfer factor (the longer the delay, the higher amplitude) and on echo transfer factor at a constant delay (the higher transfer factor, the higher amplitude). Conjunctional variation of the echo transfer factor and delay kept the AEP amplitude constant when the delay to transfer factor trade was from -7.1 to -8.4 dB per delay doubling. The results confirm the hypothesis that partial forward masking of the echoes by the preceding emitted sonar pulses serves as a time-varying automatic gain control in the auditory system of echolocating odontocetes. © 2011 Acoustical Society of America

The modulation rate transfer function of a harbour porpoise (Phocoena phocoena).

PubMed

Linnenschmidt, Meike; Wahlberg, Magnus; Damsgaard Hansen, Janni

2013-02-01

During echolocation, toothed whales produce ultrasonic clicks at extremely rapid rates and listen for the returning echoes. The auditory brainstem response (ABR) duration was evaluated in terms of latency between single peaks: 5.5 ms (from peak I to VII), 3.4 ms (I-VI), and 1.4 ms (II-IV). In comparison to the killer whale and the bottlenose dolphin, the ABR of the harbour porpoise has shorter intervals between the peaks and consequently a shorter ABR duration. This indicates that the ABR duration and peak latencies are possibly related to the relative size of the auditory structures of the central nervous system and thus to the animal's size. The ABR to a sinusoidal amplitude modulated stimulus at 125 kHz (sensitivity threshold 63 dB re 1 μPa rms) was evaluated to determine the modulation rate transfer function of a harbour porpoise. The ABR showed distinct envelope following responses up to a modulation rate of 1,900 Hz. The corresponding calculated equivalent rectangular duration of 263 μs indicates a good temporal resolution in the harbour porpoise auditory system similar to the one for the bottlenose dolphin. The results explain how the harbour porpoise can follow clicks and echoes during echolocation with very short inter click intervals.

Keeping returns optimal: gain control exerted through sensitivity adjustments in the harbour porpoise auditory system

PubMed Central

Linnenschmidt, Meike; Beedholm, Kristian; Wahlberg, Magnus; Højer-Kristensen, Jakob; Nachtigall, Paul E.

2012-01-01

Animals that use echolocation (biosonar) listen to acoustic signals with a large range of intensities, because echo levels vary with the fourth power of the animal's distance to the target. In man-made sonar, engineers apply automatic gain control to stabilize the echo energy levels, thereby rendering them independent of distance to the target. Both toothed whales and bats vary the level of their echolocation clicks to compensate for the distance-related energy loss. By monitoring the auditory brainstem response (ABR) during a psychophysical task, we found that a harbour porpoise (Phocoena phocoena), in addition to adjusting the sound level of the outgoing signals up to 5.4 dB, also reduces its ABR threshold by 6 dB when the target distance doubles. This self-induced threshold shift increases the dynamic range of the biosonar system and compensates for half of the variation of energy that is caused by changes in the distance to the target. In combination with an increased source level as a function of target range, this helps the porpoise to maintain a stable echo-evoked ABR amplitude irrespective of target range, and is therefore probably an important tool enabling porpoises to efficiently analyse and classify received echoes. PMID:22279169

Passive acoustic localization of the Atlantic bottlenose dolphin using whistles and echolocation clicks.

PubMed

Freitag, L E; Tyack, P L

1993-04-01

A method for localization and tracking of calling marine mammals was tested under realistic field conditions that include noise, multipath, and arbitrarily located sensors. Experiments were performed in two locations using four and six hydrophones with captive Atlantic bottlenose dolphins (Tursiops truncatus). Acoustic signals from the animals were collected in the field using a digital acoustic data acquisition system. The data were then processed off-line to determine relative hydrophone positions and the animal locations. Accurate hydrophone position estimates are achieved by pinging sequentially from each hydrophone to all the others. A two-step least-squares algorithm is then used to determine sensor locations from the calibration data. Animal locations are determined by estimating the time differences of arrival of the dolphin signals at the different sensors. The peak of a matched filter output or the first cycle of the observed waveform is used to determine arrival time of an echolocation click. Cross correlation between hydrophones is used to determine inter-sensor time delays of whistles. Calculation of source location using the time difference of arrival measurements is done using a least-squares solution to minimize error. These preliminary experimental results based on a small set of data show that realistic trajectories for moving animals may be generated from consecutive location estimates.

Life history constrains biochemical development in the highly specialized odontocete echolocation system

PubMed Central

Koopman, Heather N; Zahorodny, Zoey P

2008-01-01

The vertebrate head has undergone enormous modification from the features borne by early ancestors. The growth of skull bones has been well studied in many species, yet little is known about corresponding soft tissue development. Among mammals, some of the most unusual examples of cranial evolution exist in the toothed whales (odontocetes). Specialized fat bodies in toothed whale heads play important roles in sound transmission and reception. These fat bodies contain unique endogenous lipids, with favourable acoustic properties, arranged in highly organized, three-dimensional patterns. We link variation in developmental rates of acoustic fats with life-history strategy, using bottlenose dolphins and harbour porpoises. Porpoise acoustic fats attain adult configurations earlier (less than 1 year) and at a faster pace than dolphins. The accelerated lipid accumulation in porpoises reflects the earlier need for fully functional echolocation systems. Dolphins enjoy 3–6 years of maternal care; porpoises must achieve total independence by approximately nine months. Further, a stereotypic ‘blueprint’ for the spatial distribution of lipids is established prior to birth, demonstrating the highly conserved nature of the intricate biochemical arrangement in acoustic tissues. This system illustrates an unusual case of soft tissue development being constrained by life history, rather than the more commonly observed mechanistic or phyletic constraints. PMID:18611851

Target representation of naturalistic echolocation sequences in single unit responses from the inferior colliculus of big brown bats

NASA Astrophysics Data System (ADS)

Sanderson, Mark I.; Simmons, James A.

2005-11-01

Echolocating big brown bats (Eptesicus fuscus) emit trains of frequency-modulated (FM) biosonar signals whose duration, repetition rate, and sweep structure change systematically during interception of prey. When stimulated with a 2.5-s sequence of 54 FM pulse-echo pairs that mimic sounds received during search, approach, and terminal stages of pursuit, single neurons (N=116) in the bat's inferior colliculus (IC) register the occurrence of a pulse or echo with an average of <1 spike/sound. Individual IC neurons typically respond to only a segment of the search or approach stage of pursuit, with fewer neurons persisting to respond in the terminal stage. Composite peristimulus-time-histogram plots of responses assembled across the whole recorded population of IC neurons depict the delay of echoes and, hence, the existence and distance of the simulated biosonar target, entirely as on-response latencies distributed across time. Correlated changes in pulse duration, repetition rate, and pulse or echo amplitude do modulate the strength of responses (probability of the single spike actually occurring for each sound), but registration of the target itself remains confined exclusively to the latencies of single spikes across cells. Modeling of echo processing in FM biosonar should emphasize spike-time algorithms to explain the content of biosonar images.

An echolocation model for the restoration of an acoustic image from a single-emission echo

NASA Astrophysics Data System (ADS)

Matsuo, Ikuo; Yano, Masafumi

2004-12-01

Bats can form a fine acoustic image of an object using frequency-modulated echolocation sound. The acoustic image is an impulse response, known as a reflected-intensity distribution, which is composed of amplitude and phase spectra over a range of frequencies. However, bats detect only the amplitude spectrum due to the low-time resolution of their peripheral auditory system, and the frequency range of emission is restricted. It is therefore necessary to restore the acoustic image from limited information. The amplitude spectrum varies with the changes in the configuration of the reflected-intensity distribution, while the phase spectrum varies with the changes in its configuration and location. Here, by introducing some reasonable constraints, a method is proposed for restoring an acoustic image from the echo. The configuration is extrapolated from the amplitude spectrum of the restricted frequency range by using the continuity condition of the amplitude spectrum at the minimum frequency of the emission and the minimum phase condition. The determination of the location requires extracting the amplitude spectra, which vary with its location. For this purpose, the Gaussian chirplets with a carrier frequency compatible with bat emission sweep rates were used. The location is estimated from the temporal changes of the amplitude spectra. .

Wind turbines and bat mortality: Doppler shift profiles and ultrasonic bat-like pulse reflection from moving turbine blades.

PubMed

Long, Chloe V; Flint, James A; Lepper, Paul A

2010-10-01

Bat mortality resulting from actual or near-collision with operational wind turbine rotors is a phenomenon that is widespread but not well understood. Because bats rely on information contained in high-frequency echoes to determine the nature and movement of a target, it is important to consider how ultrasonic pulses similar to those used by bats for echolocation may be interacting with operational turbine rotor blades. By assessing the characteristics of reflected ultrasonic echoes, moving turbine blades operating under low wind speed conditions (<6 m s(-1)) were found to produce distinct Doppler shift profiles at different angles to the rotor. Frequency shifts of up to ±700-800 Hz were produced, which may not be perceptible by some bat species. Monte Carlo simulation of bat-like sampling by echolocation revealed that over 50 rotor echoes could be required by species such as Pipistrellus pipistrellus for accurate interpretation of blade movement, which may not be achieved in the bat's approach time-window. In summary, it was found that echoes returned from moving blades had features which could render them attractive to bats or which might make it difficult for the bat to accurately detect and locate blades in sufficient time to avoid a collision.

Comparison of echolocation clicks from geographically sympatric killer whales and long-finned pilot whales (L).

PubMed

Eskesen, Ida G; Wahlberg, Magnus; Simon, Malene; Larsen, Ole Næsbye

2011-07-01

The source characteristics of biosonar signals from sympatric killer whales and long-finned pilot whales in a Norwegian fjord were compared. A total of 137 pilot whale and more than 2000 killer whale echolocation clicks were recorded using a linear four-hydrophone array. Of these, 20 pilot whale clicks and 28 killer whale clicks were categorized as being recorded on-axis. The clicks of pilot whales had a mean apparent source level of 196 dB re 1 μPa pp and those of killer whales 203 dB re 1 μPa pp. The duration of pilot whale clicks was significantly shorter (23 μs, S.E.=1.3) and the centroid frequency significantly higher (55 kHz, S.E.=2.1) than killer whale clicks (duration: 41 μs, S.E.=2.6; centroid frequency: 32 kHz, S.E.=1.5). The rate of increase in the accumulated energy as a function of time also differed between clicks from the two species. The differences in duration, frequency, and energy distribution may have a potential to allow for the distinction between pilot and killer whale clicks when using automated detection routines for acoustic monitoring. © 2011 Acoustical Society of America

Active Listening in a Bat Cocktail Party: Adaptive Echolocation and Flight Behaviors of Big Brown Bats, Eptesicus fuscus, Foraging in a Cluttered Acoustic Environment.

PubMed

Warnecke, Michaela; Chiu, Chen; Engelberg, Jonathan; Moss, Cynthia F

2015-09-01

In their natural environment, big brown bats forage for small insects in open spaces, as well as in vegetation and in the presence of acoustic clutter. While searching and hunting for prey, bats experience sonar interference, not only from densely cluttered environments, but also from calls of conspecifics foraging in close proximity. Previous work has shown that when two bats compete for a single prey item in a relatively open environment, one of the bats may go silent for extended periods of time, which can serve to minimize sonar interference between conspecifics. Additionally, pairs of big brown bats have been shown to adjust frequency characteristics of their vocalizations to avoid acoustic interference in echo processing. In this study, we extended previous work by examining how the presence of conspecifics and environmental clutter influence the bat's echolocation behavior. By recording multichannel audio and video data of bats engaged in insect capture in open and cluttered spaces, we quantified the bats' vocal and flight behaviors. Big brown bats flew individually and in pairs in an open and cluttered room, and the results of this study shed light on the different strategies that this species employs to negotiate a complex and dynamic environment. © 2015 S. Karger AG, Basel.

The Origin of High-Frequency Hearing in Whales.

PubMed

Churchill, Morgan; Martinez-Caceres, Manuel; de Muizon, Christian; Mnieckowski, Jessica; Geisler, Jonathan H

2016-08-22

Odontocetes (toothed whales) rely upon echoes of their own vocalizations to navigate and find prey underwater [1]. This sensory adaptation, known as echolocation, operates most effectively when using high frequencies, and odontocetes are rivaled only by bats in their ability to perceive ultrasonic sound greater than 100 kHz [2]. Although features indicative of ultrasonic hearing are present in the oldest known odontocetes [3], the significance of this finding is limited by the methods employed and taxa sampled. In this report, we describe a new xenorophid whale (Echovenator sandersi, gen. et sp. nov.) from the Oligocene of South Carolina that, as a member of the most basal clade of odontocetes, sheds considerable light on the evolution of ultrasonic hearing. By placing high-resolution CT data from Echovenator sandersi, 2 hippos, and 23 fossil and extant whales in a phylogenetic context, we conclude that ultrasonic hearing, albeit in a less specialized form, evolved at the base of the odontocete radiation. Contrary to the hypothesis that odontocetes evolved from low-frequency specialists [4], we find evidence that stem cetaceans, the archaeocetes, were more sensitive to high-frequency sound than their terrestrial ancestors. This indicates that selection for high-frequency hearing predates the emergence of Odontoceti and the evolution of echolocation. Copyright © 2016 Elsevier Ltd. All rights reserved.

Insight on how fishing bats discern prey and adjust their mechanic and sensorial features during the attack sequence

PubMed Central

Aizpurua, Ostaizka; Alberdi, Antton; Aihartza, Joxerra; Garin, Inazio

2015-01-01

Several insectivorous bats have included fish in their diet, yet little is known about the processes underlying this trophic shift. We performed three field experiments with wild fishing bats to address how they manage to discern fish from insects and adapt their hunting technique to capture fish. We show that bats react only to targets protruding above the water and discern fish from insects based on prey disappearance patterns. Stationary fish trigger short and shallow dips and a terminal echolocation pattern with an important component of the narrowband and low frequency calls. When the fish disappears during the attack process, bats regulate their attack increasing the number of broadband and high frequency calls in the last phase of the echolocation as well as by lengthening and deepening their dips. These adjustments may allow bats to obtain more valuable sensorial information and to perform dips adjusted to the level of uncertainty on the location of the submerged prey. The observed ultrafast regulation may be essential for enabling fishing to become cost-effective in bats, and demonstrates the ability of bats to rapidly modify and synchronise their sensorial and motor features as a response to last minute stimulus variations. PMID:26196094

Marine Mammals: Hearing and Echolocation at Coconut Island

DTIC Science & Technology

2009-09-30

113(5), 2408-2411 5 Supin, A. Ya., Nachtigall, P.E., and Breese, M. (2009) Forward masking based gain control in odontocete biosonar : an evoked...odontocete biosonar : an evoked-potential study. J Acous Soc Am (published, refereed) Mooney T.A., Nachtigall, P.E. Breese, M. Vlachos, S. and...Nachtigall, P.E. and Breese, M. (2008) Forward masking as a mechanism of automatic gain control in whale biosonar : a psychophysical study, Journal of the

Echolocation-Based Foraging by Harbor Porpoises and Sperm Whales, Including Effects on Noise and Acoustic Propagation

DTIC Science & Technology

2008-09-01

Behavioural Point Process Data 234 Appendix B: Matlab Code 258 Matlab Code Used in Chapter 2 (Porpoise Prey Capture Analysis) 258 Click Extraction and...Measurement of Click Properties 258 Envelope-based Click Detector 262 Matlab Code Used in Chapter 3 (Transmission Loss in Porpoise Habitats) ..267...Click Extraction from Data Wavefiles 267 Click Level Determination (Grand Manan Datasets) 270 Click Level Determination (Danish Datasets) 287 Matlab

Effect of Broadband Nature of Marine Mammal Echolocation Clicks on Click-Based Population Density Estimates

DTIC Science & Technology

2014-09-30

initially on beaked whales (Blainville’s or Cuvier’s), for which high quality click recordings of clicks are available from DTAG data, as these are known...will be applied also to other species such as sperm whale (Physeter macrocephalus) (whose high source level assures long range detection and amplifies...REFERENCES Gillespie, D. and Leaper, R. (1996). Detection of sperm whale Physeter macrocephalus clicks and discrimination of individual vocalizations

Effect of Broadband Nature of Marine Mammal Echolocation Clicks on Click-Based Population Density Estimates

DTIC Science & Technology

2014-09-30

research will focus initially on beaked whales (Blainville’s or Cuvier’s), for which high quality click recordings of clicks are available from DTAG...The same methodology will be applied also to other species such as sperm whale (Physeter macrocephalus) (whose high source level assures long range...Thomas, University of St. Andrews). REFERENCES Gillespie, D. and Leaper, R. (1996). Detection of sperm whale Physeter macrocephalus clicks and

Ambient Noise in the Sea

DTIC Science & Technology

1984-01-01

the 14 to 222 Hz band. In a tank, the echolocation signals made by dolphins were found (167) to be directional, with peak energies above 100 kHz...17. Daniels, F. B., Mechanisms of Generation of Infrasound by Ocean Waves, JASA 24, 83, 1952. 18. Daniels, F. B., Generation of Infrasound by Ocean...in the Ocean by Surface Waves, J. Sound and Vibration 37, 185, 1974. 58. Hughes, B., Estimates of Underwater Sound (and Infrasound ) Produced by Non

Ocean, Platform, and Signal Processing Effects on Synthetic Aperture Sonar Performance

DTIC Science & Technology

1991-02-01

Canadian Reginald A. Fessenden , of the Submarine Signal Company, Boston, MA during the period from 1912 to 1914 [Fay, c. 19441. Fessenden’s invention...usually referred to as the Fessenden oscillator. It was conceived by Fessenden as an instrument for underwater communication from ship to ship, and more...accident it was observed that the ocean depth could also be measured. In these landmark experiments, Fessenden showed telemetry, echolocation and

A Fully Integrated Materials Framework for Enabling the Wireless Detection of Micro-defects in Aging and Battle-worn Structures

DTIC Science & Technology

2012-05-01

field-programmable gate array (FPGA) uses digital signal processing (DSP) algorithms to decode echo-location information from the backscattered signal ...characterizing and understanding of the physical properties of the BST and PZT thin films. Using microwave reflection spectroscopy, the complex...acoustic data, , would be encoded in the reflected MW signal by means of phase modulation (PM). By using high-Q resonators as the reactive

Advanced Methods for Passive Acoustic Detection, Classification, and Localization of Marine Mammals

DTIC Science & Technology

2015-09-30

floor 1176 Howell St. Newport, RI 02842 phone: (401) 832-5749 fax: (401) 832-4441 email: David.Moretti@navy.mil Steve W. Martin... Jarvis et al. 2008). This classifier both detects and classifies echolocation clicks from five species of odontocetes, including Blainville’s and...Cuvier’s beaked whales, Risso’s dolphins, short-finned pilot whales, and sperm whales. Here Moretti’s group, particularly S. Jarvis , is improving the

Advanced Methods for Passive Acoustic Detection, Classification, and Localization of Marine Mammals

DTIC Science & Technology

2013-09-30

N0001411WX21394 Steve W. Martin SPAWAR Systems Center Pacific 53366 Front St. San Diego, CA 92152-6551 phone: (619) 553-9882 email: Steve.W.Martin...multiclass support vector machine (SVM) classifier was previously developed ( Jarvis et al. 2008). This classifier both detects and classifies echolocation...whales. Here Moretti’s group, particularly S. Jarvis , will improve the SVM classifier by resolving confusion between species whose clicks overlap in

Multiecho Processing by an Echolocating Dolphin

DTIC Science & Technology

2003-08-01

with the number of stimulus information, is less clear (Dankiewicz et al., 2002; Moore presentations. etal., 1991; Roitblat etal., 1991). Dependence...W. B., Roitblat , H. L., Penner, R. H., and Nachtigall, P. E. (1991). ting yields a large false-alarm rate that cannot be changed "Recognizing... Roitblat , H. L., Moore, P. W. B., Nachtigall, P. E., and Penner, R. H. (1991). Au, W. W. L., and Banks, K. (1998). "The acoustics of the snapping shrimp

Development of Automated Whistle and Click Classifiers for Odontocete Species in the Western Atlantic Ocean and the Waters Surrounding the Hawaiian Islands

DTIC Science & Technology

2015-09-30

for Odontocete Species in the Western Atlantic Ocean and the Waters Surrounding the Hawaiian Islands Julie N. Oswald & Tina M. Yack Bio-Waves... Atlantic Ocean, the temperate Pacific Ocean and the waters surrounding the Hawaiian Islands. These classifiers will also incorporate ancillary...and echolocation click classifiers for odontocete species in the northwest Atlantic Ocean, the waters surrounding the Hawaiian Islands and the

Effect of Broadband Nature of Marine Mammal Echolocation Clicks on Click-Based Population Density Estimates

DTIC Science & Technology

2015-09-30

sonar sound and therefore of high relevance to the US Navy, and sperm whale (Physeter macrocephalus) clicks, whose high source level assures long...beaked whale, and Zimmer (2011) for the sperm whale. 5 For background noise we used measurements obtained with the Delphinus towed array at the US...Atlantic Undersea Test and Evaluation Center (AUTEC), where both Blainville’s beaked whales and sperm whales are commonly present. Analysis of the

Single-locus species delimitation: a test of the mixed Yule-coalescent model, with an empirical application to Philippine round-leaf bats.

PubMed

Esselstyn, Jacob A; Evans, Ben J; Sedlock, Jodi L; Anwarali Khan, Faisal Ali; Heaney, Lawrence R

2012-09-22

Prospects for a comprehensive inventory of global biodiversity would be greatly improved by automating methods of species delimitation. The general mixed Yule-coalescent (GMYC) was recently proposed as a potential means of increasing the rate of biodiversity exploration. We tested this method with simulated data and applied it to a group of poorly known bats (Hipposideros) from the Philippines. We then used echolocation call characteristics to evaluate the plausibility of species boundaries suggested by GMYC. In our simulations, GMYC performed relatively well (errors in estimated species diversity less than 25%) when the product of the haploid effective population size (N(e)) and speciation rate (SR; per lineage per million years) was less than or equal to 10(5), while interspecific variation in N(e) was twofold or less. However, at higher but also biologically relevant values of N(e) × SR and when N(e) varied tenfold among species, performance was very poor. GMYC analyses of mitochondrial DNA sequences from Philippine Hipposideros suggest actual diversity may be approximately twice the current estimate, and available echolocation call data are mostly consistent with GMYC delimitations. In conclusion, we consider the GMYC model useful under some conditions, but additional information on N(e), SR and/or corroboration from independent character data are needed to allow meaningful interpretation of results.

Timing matters: sonar call groups facilitate target localization in bats.

PubMed

Kothari, Ninad B; Wohlgemuth, Melville J; Hulgard, Katrine; Surlykke, Annemarie; Moss, Cynthia F

2014-01-01

To successfully negotiate a cluttered environment, an echolocating bat must control the timing of motor behaviors in response to dynamic sensory information. Here we detail the big brown bat's adaptive temporal control over sonar call production for tracking prey, moving predictably or unpredictably, under different experimental conditions. We studied the adaptive control of vocal-motor behaviors in free-flying big brown bats, Eptesicus fuscus, as they captured tethered and free-flying insects, in open and cluttered environments. We also studied adaptive sonar behavior in bats trained to track moving targets from a resting position. In each of these experiments, bats adjusted the features of their calls to separate target and clutter. Under many task conditions, flying bats produced prominent sonar sound groups identified as clusters of echolocation pulses with relatively stable intervals, surrounded by longer pulse intervals. In experiments where bats tracked approaching targets from a resting position, bats also produced sonar sound groups, and the prevalence of these sonar sound groups increased when motion of the target was unpredictable. We hypothesize that sonar sound groups produced during flight, and the sonar call doublets produced by a bat tracking a target from a resting position, help the animal resolve dynamic target location and represent the echo scene in greater detail. Collectively, our data reveal adaptive temporal control over sonar call production that allows the bat to negotiate a complex and dynamic environment.

Timing matters: sonar call groups facilitate target localization in bats

PubMed Central

Kothari, Ninad B.; Wohlgemuth, Melville J.; Hulgard, Katrine; Surlykke, Annemarie; Moss, Cynthia F.

2014-01-01

To successfully negotiate a cluttered environment, an echolocating bat must control the timing of motor behaviors in response to dynamic sensory information. Here we detail the big brown bat's adaptive temporal control over sonar call production for tracking prey, moving predictably or unpredictably, under different experimental conditions. We studied the adaptive control of vocal-motor behaviors in free-flying big brown bats, Eptesicus fuscus, as they captured tethered and free-flying insects, in open and cluttered environments. We also studied adaptive sonar behavior in bats trained to track moving targets from a resting position. In each of these experiments, bats adjusted the features of their calls to separate target and clutter. Under many task conditions, flying bats produced prominent sonar sound groups identified as clusters of echolocation pulses with relatively stable intervals, surrounded by longer pulse intervals. In experiments where bats tracked approaching targets from a resting position, bats also produced sonar sound groups, and the prevalence of these sonar sound groups increased when motion of the target was unpredictable. We hypothesize that sonar sound groups produced during flight, and the sonar call doublets produced by a bat tracking a target from a resting position, help the animal resolve dynamic target location and represent the echo scene in greater detail. Collectively, our data reveal adaptive temporal control over sonar call production that allows the bat to negotiate a complex and dynamic environment. PMID:24860509

Effects of competitive prey capture on flight behavior and sonar beam pattern in paired big brown bats, Eptesicus fuscus.

PubMed

Chiu, Chen; Reddy, Puduru Viswanadha; Xian, Wei; Krishnaprasad, Perinkulam S; Moss, Cynthia F

2010-10-01

Foraging and flight behavior of echolocating bats were quantitatively analyzed in this study. Paired big brown bats, Eptesicus fuscus, competed for a single food item in a large laboratory flight room. Their sonar beam patterns and flight paths were recorded by a microphone array and two high-speed cameras, respectively. Bats often remained in nearly classical pursuit (CP) states when one bat is following another bat. A follower can detect and anticipate the movement of the leader, while the leader has the advantage of gaining access to the prey first. Bats in the trailing position throughout the trial were more successful in accessing the prey. In this study, bats also used their sonar beam to monitor the conspecific's movement and to track the prey. Each bat tended to use its sonar beam to track the prey when it was closer to the worm than to another bat. The trailing bat often directed its sonar beam toward the leading bat in following flight. When two bats flew towards each other, they tended to direct their sonar beam axes away from each other, presumably to avoid signal jamming. This study provides a new perspective on how echolocating bats use their biosonar system to coordinate their flight with conspecifics in a group and how they compete for the same food source with conspecifics.

Acoustic and foraging behavior of a Baird's beaked whale, Berardius bairdii, exposed to simulated sonar

PubMed Central

Stimpert, A. K.; DeRuiter, S. L.; Southall, B. L.; Moretti, D. J.; Falcone, E. A.; Goldbogen, J. A.; Friedlaender, A.; Schorr, G. S.; Calambokidis, J.

2014-01-01

Beaked whales are hypothesized to be particularly sensitive to anthropogenic noise, based on previous strandings and limited experimental and observational data. However, few species have been studied in detail. We describe the underwater behavior of a Baird's beaked whale (Berardius bairdii) from the first deployment of a multi-sensor acoustic tag on this species. The animal exhibited shallow (23 ± 15 m max depth), intermediate (324 ± 49 m), and deep (1138 ± 243 m) dives. Echolocation clicks were produced with a mean inter-click interval of approximately 300 ms and peak frequency of 25 kHz. Two deep dives included presumed foraging behavior, with echolocation pulsed sounds (presumed prey capture attempts) associated with increased maneuvering, and sustained inverted swimming during the bottom phase of the dive. A controlled exposure to simulated mid-frequency active sonar (3.5–4 kHz) was conducted 4 hours after tag deployment, and within 3 minutes of exposure onset, the tagged whale increased swim speed and body movement, and continued to show unusual dive behavior for each of its next three dives, one of each type. These are the first data on the acoustic foraging behavior in this largest beaked whale species, and the first experimental demonstration of a response to simulated sonar. PMID:25391309

The Source Parameters of Echolocation Clicks from Captive and Free-Ranging Yangtze Finless Porpoises (Neophocaena asiaeorientalis asiaeorientalis)

PubMed Central

Fang, Liang; Wang, Ding; Li, Yongtao; Cheng, Zhaolong; Pine, Matthew K.; Wang, Kexiong; Li, Songhai

2015-01-01

The clicks of Yangtze finless porpoises (Neophocaena asiaeorientalis asiaeorientalis) from 7 individuals in the tank of Baiji aquarium, 2 individuals in a netted pen at Shishou Tian-e-zhou Reserve and 4 free-ranging individuals at Tianxingzhou were recorded using a broadband digital recording system with four element hydrophones. The peak-to-peak apparent source level (ASL_pp) of clicks from individuals at the Baiji aquarium was 167 dB re 1 μPa with mean center frequency of 133 kHz, -3dB bandwidth of 18 kHz and -10 dB duration of 58 μs. The ASL_pp of clicks from individuals at the Shishou Tian-e-zhou Reserve was 180 dB re 1 μPa with mean center frequency of 128 kHz, -3dB bandwidth of 20 kHz and -10 dB duration of 39 μs. The ASL_pp of clicks from individuals at Tianxingzhou was 176 dB re 1 μPa with mean center frequency of 129 kHz, -3dB bandwidth of 15 kHz and -10 dB duration of 48 μs. Differences between the source parameters of clicks among the three groups of finless porpoises suggest these animals adapt to their echolocation signals depending on their surroundings. PMID:26053758

Echolocation clicks of free-ranging Cuvier's beaked whales (Ziphius cavirostris)

NASA Astrophysics Data System (ADS)

Zimmer, Walter M. X.; Johnson, Mark P.; Madsen, Peter T.; Tyack, Peter L.

2005-06-01

Strandings of beaked whales of the genera Ziphius and Mesoplodon have been reported to occur in conjunction with naval sonar use. Detection of the sounds from these elusive whales could reduce the risk of exposure, but descriptions of their vocalizations are at best incomplete. This paper reports quantitative characteristics of clicks from deep-diving Cuvier's beaked whales (Ziphius cavirostris) using a unique data set. Two whales in the Ligurian Sea were simultaneously tagged with sound and orientation recording tags, and the dive tracks were reconstructed allowing for derivation of the range and relative aspect between the clicking whales. At depth, the whales produced trains of regular echolocation clicks with mean interclick intervals of 0.43 s (+/-0.09) and 0.40 s (+/-0.07). The clicks are frequency modulated pulses with durations of ~200 μs and center frequencies around 42 kHz, -10 dB bandwidths of 22 kHz, and Q3 dB of 4. The sound beam is narrow with an estimated directionality index of more than 25 dB, source levels up to 214 dBpp re: 1 μPa at 1 m, and energy flux density of 164 dB re: 1 μPa2 s. As the spectral and temporal properties are different from those of nonziphiid odontocetes the potential for passive detection is enhanced. .

Echolocating bats use future-target information for optimal foraging.

PubMed

Fujioka, Emyo; Aihara, Ikkyu; Sumiya, Miwa; Aihara, Kazuyuki; Hiryu, Shizuko

2016-04-26

When seeing or listening to an object, we aim our attention toward it. While capturing prey, many animal species focus their visual or acoustic attention toward the prey. However, for multiple prey items, the direction and timing of attention for effective foraging remain unknown. In this study, we adopted both experimental and mathematical methodology with microphone-array measurements and mathematical modeling analysis to quantify the attention of echolocating bats that were repeatedly capturing airborne insects in the field. Here we show that bats select rational flight paths to consecutively capture multiple prey items. Microphone-array measurements showed that bats direct their sonar attention not only to the immediate prey but also to the next prey. In addition, we found that a bat's attention in terms of its flight also aims toward the next prey even when approaching the immediate prey. Numerical simulations revealed a possibility that bats shift their flight attention to control suitable flight paths for consecutive capture. When a bat only aims its flight attention toward its immediate prey, it rarely succeeds in capturing the next prey. These findings indicate that bats gain increased benefit by distributing their attention among multiple targets and planning the future flight path based on additional information of the next prey. These experimental and mathematical studies allowed us to observe the process of decision making by bats during their natural flight dynamics.

Mechanisms underlying intensity-dependent changes in cortical selectivity for frequency-modulated sweeps.

PubMed

Razak, K A

2012-04-01

Frequency-modulated (FM) sweeps are common components of species-specific vocalizations. The intensity of FM sweeps can cover a wide range in the natural environment, but whether intensity affects neural selectivity for FM sweeps is unclear. Bats, such as the pallid bat, which use FM sweeps for echolocation, are suited to address this issue, because the intensity of echoes will vary with target distance. In this study, FM sweep rate selectivity of pallid bat auditory cortex neurons was measured using downward sweeps at different intensities. Neurons became more selective for FM sweep rates present in the bat's echolocation calls as intensity increased. Increased selectivity resulted from stronger inhibition of responses to slower sweep rates. The timing and bandwidth of inhibition generated by frequencies on the high side of the excitatory tuning curve [sideband high-frequency inhibition (HFI)] shape rate selectivity in cortical neurons in the pallid bat. To determine whether intensity-dependent changes in FM rate selectivity were due to altered inhibition, the timing and bandwidth of HFI were quantified at multiple intensities using the two-tone inhibition paradigm. HFI arrived faster relative to excitation as sound intensity increased. The bandwidth of HFI also increased with intensity. The changes in HFI predicted intensity-dependent changes in FM rate selectivity. These data suggest that neural selectivity for a sweep parameter is not static but shifts with intensity due to changes in properties of sideband inhibition.

High Frequency Acoustic Recording Package, Data Summary Report, PS09, February 26, 2010 - November 03, 2010

DTIC Science & Technology

2012-10-01

blue whale vocalizations consisted of A and B calls, either as songs or individual calls, as well as several D calls associated with foraging. See...humpback whale songs had bi- modal annual structure: continuous presence in September–October and secondary peak in March–April (Fig. 6). Sperm whale ...Blue whale B call 1429 24% 19 8% Blue whale D call 40 ə% 10 4% Humpback whale Song 2417 40% 119 47% Sperm whale echolocation 302 25% 55 22

Cetacean Bioacoustics with Emphasis on Recording and Monitoring

NASA Astrophysics Data System (ADS)

Akamatsu, Tomonari

More than 80 cetacean species live in oceans, lakes, and rivers. For underwater navigation and recognition, whales and dolphins have developed unique sensory systems using acoustic signals. Toothed whales, such as dolphins and porpoises, have sonar using ultrasonic pulse trains called echolocations (Au, 1993). As top predators in the water, dolphins and porpoises rely on accurate and long-range sensory systems for catching prey. Dolphins have another type of vocalization called a whistle that is narrowband with a long duration.

Effect of Broadband Nature of Marine Mammal Echolocation Clicks on Click-Based Population Density Estimates

DTIC Science & Technology

2015-09-30

Blainville’s beaked whales (Mesoplodon densirostris), known to be sensitive to sonar sound and therefore of high relevance to the US Navy, and sperm whale...this study were obtained from von Benda-Beckmann et al. (2010) for the beaked whale, and Zimmer (2011) for the sperm whale. 4 For background...Blainville’s beaked whales and sperm whales are commonly present. Analysis of the noise levels on the system indicated that the background noise was system

Navigation: bat orientation using Earth's magnetic field.

PubMed

Holland, Richard A; Thorup, Kasper; Vonhof, Maarten J; Cochran, William W; Wikelski, Martin

2006-12-07

Bats famously orientate at night by echolocation, but this works over only a short range, and little is known about how they navigate over longer distances. Here we show that the homing behaviour of Eptesicus fuscus, known as the big brown bat, can be altered by artificially shifting the Earth's magnetic field, indicating that these bats rely on a magnetic compass to return to their home roost. This finding adds to the impressive array of sensory abilities possessed by this animal for navigation in the dark.

Discrimination of Amplitude-Modulated Synthetic Echo Trains by an Echolocating Bottlenose Dolphin

DTIC Science & Technology

2002-07-09

resulted in improved echo categorization (Moore FIG. 1. (A) Enlargement of a single stimulus pulse (left) and its spectrum eta!., 1991; Roitblat eta!., 1991...Floyd, 1980; Moore et aL., 1991; Roitblat J. Acoust. Soc. Am., Vol. 112, No. 4, October 2002 Dankiewicz et al.: Discrimination of AM by an...Those studies were designed to elu- Harley, H. E., Roitblat , H. L., and Nachtigall, P. E. (1996). "Object repre- cidate the possible mechanisms by which

Portable Multi Hydrophone Array for Field and Laboratory Measurements of Odontocete Acoustic Signals

DTIC Science & Technology

2015-09-30

bottlenose dolphin and a false killer whale as well as other species such as the Risso’s dolphin . We are currently comparing various file types and methods...echolocation abilities of a Risso’s dolphin are currently being reviewed for publication. 3 The hook detection study recordings are being analyzed for basic...element array in order to obtain horizontal and vertical beam patterns of acoustic signals of a false killer whale and a bottlenose dolphin . The data

The sonar aperture and its neural representation in bats.

PubMed

Heinrich, Melina; Warmbold, Alexander; Hoffmann, Susanne; Firzlaff, Uwe; Wiegrebe, Lutz

2011-10-26

As opposed to visual imaging, biosonar imaging of spatial object properties represents a challenge for the auditory system because its sensory epithelium is not arranged along space axes. For echolocating bats, object width is encoded by the amplitude of its echo (echo intensity) but also by the naturally covarying spread of angles of incidence from which the echoes impinge on the bat's ears (sonar aperture). It is unclear whether bats use the echo intensity and/or the sonar aperture to estimate an object's width. We addressed this question in a combined psychophysical and electrophysiological approach. In three virtual-object playback experiments, bats of the species Phyllostomus discolor had to discriminate simple reflections of their own echolocation calls differing in echo intensity, sonar aperture, or both. Discrimination performance for objects with physically correct covariation of sonar aperture and echo intensity ("object width") did not differ from discrimination performances when only the sonar aperture was varied. Thus, the bats were able to detect changes in object width in the absence of intensity cues. The psychophysical results are reflected in the responses of a population of units in the auditory midbrain and cortex that responded strongest to echoes from objects with a specific sonar aperture, regardless of variations in echo intensity. Neurometric functions obtained from cortical units encoding the sonar aperture are sufficient to explain the behavioral performance of the bats. These current data show that the sonar aperture is a behaviorally relevant and reliably encoded cue for object size in bat sonar.

Acoustic surveys of Hawaiian Hoary Bats in Kahikinui Forest Reserve and Nakula Natural Area Reserve on the Island of Maui

USGS Publications Warehouse

Todd, Christopher M.; Pinzari, Corinna A.; Bonaccorso, Frank

2016-01-01

The Kahikinui Forest Reserve and the adjoining Nakula Natural Area Reserve (KFR-NNAR) was established in 2011 as a conservation area on the leeward slope of Haleakalā Volcano on the island of Maui to protect unique natural features and endangered species including the Hawaiian hoary bat, Lasiurus cinereus semotus. We recorded bat vocalizations from July 2012 to November 2014 using automated echolocation detectors at 14 point locations in the KFRNNAR. Our study area included remnants of recovering mesic montane forest with interspersed grasses (1,250‒1,850 m elevation, hereafter called “forest”) and xeric subalpine shrubland plant communities (1,860‒2,800 m, hereafter called “shrubland”). Monthly detections of Hawaiian hoary bats, Lasiurus cinereus semotus, within the KFR-NNAR identified areas of high and low detection probability as well as foraging activity. Sixty per cent of all detector-nights had confirmed bat vocalizations and included detections in every month of the study. Monthly detection probability values were highest from July to November 2012; these values were significantly greater than values measured in any month thereafter. Pooled values of detection probabilities, mean pulses/night, percentage of nights with feeding activity, and acoustic detections all were greater in the recovering forest zone than corresponding values from the shrublands. Our data provide baseline levels of hoary bat echolocation activity that may be compared with future studies in the KFR-NNAR relative to success criteria for Hawaiian hoary bat habitat restoration.

Echolocation signals and transmission beam pattern of a false killer whale (Pseudorca crassidens).

PubMed

Au, W W; Pawloski, J L; Nachtigall, P E; Blonz, M; Gisner, R C

1995-07-01

The echolocation transmission beam pattern of a false killer whale (Pseudorca crassidens) was measured in the vertical and horizontal planes. A vertical array of seven broadband miniature hydrophones was used to measure the beam pattern in the vertical plane and a horizontal array of the same hydrophones was used in the horizontal plane. The measurements were performed in the open waters of Kaneohe Bay, Oahu, Hawaii, while the whale performed a target discrimination task. Four types of signals, characterized by their frequency spectra, were measured. Type-1 signals had a single low-frequency peak at 40 +/- 9 kHz and a low-amplitude shoulder at high frequencies. Type-2 signals had a bimodal frequency characteristic with a primary peak at 46 +/- 7 kHz and a secondary peak at 88 +/- 13 kHz. Type-3 signals were also bimodal but with a primary peak at 100 +/- 7 kHz and a secondary peak at 49 +/- 9 kHz. Type-4 signals had a single high-frequency peak at 104 +/- 7 kHz. The center frequency of the signals were found to be linearly correlated to the peak-to-peak source level, increasing with increasing source level. The major axis of the vertical beam was directed slightly downward between 0 and -5 degrees, in contrast to the +5 to 10 degrees for Tursiops and Delphinapterus. The beam in the horizontal plane was directed forward between 0 degrees and -5 degrees.(ABSTRACT TRUNCATED AT 250 WORDS)

Spike Train Similarity Space (SSIMS) Method Detects Effects of Obstacle Proximity and Experience on Temporal Patterning of Bat Biosonar

PubMed Central

Accomando, Alyssa W.; Vargas-Irwin, Carlos E.; Simmons, James A.

2018-01-01

Bats emit biosonar pulses in complex temporal patterns that change to accommodate dynamic surroundings. Efforts to quantify these patterns have included analyses of inter-pulse intervals, sonar sound groups, and changes in individual signal parameters such as duration or frequency. Here, the similarity in temporal structure between trains of biosonar pulses is assessed. The spike train similarity space (SSIMS) algorithm, originally designed for neural activity pattern analysis, was applied to determine which features of the environment influence temporal patterning of pulses emitted by flying big brown bats, Eptesicus fuscus. In these laboratory experiments, bats flew down a flight corridor through an obstacle array. The corridor varied in width (100, 70, or 40 cm) and shape (straight or curved). Using a relational point-process framework, SSIMS was able to discriminate between echolocation call sequences recorded from flights in each of the corridor widths. SSIMS was also able to tell the difference between pulse trains recorded during flights where corridor shape through the obstacle array matched the previous trials (fixed, or expected) as opposed to those recorded from flights with randomized corridor shape (variable, or unexpected), but only for the flight path shape in which the bats had previous training. The results show that experience influences the temporal patterns with which bats emit their echolocation calls. It is demonstrated that obstacle proximity to the bat affects call patterns more dramatically than flight path shape. PMID:29472848

Echolocation parameters of Australian humpback dolphins (Sousa sahulensis) and Indo-Pacific bottlenose dolphins (Tursiops aduncus) in the wild.

PubMed

de Freitas, Mafalda; Jensen, Frants H; Tyne, Julian; Bejder, Lars; Madsen, Peter T

2015-06-01

Echolocation is a key sensory modality for toothed whale orientation, navigation, and foraging. However, a more comparative understanding of the biosonar properties of toothed whales is necessary to understand behavioral and evolutionary adaptions. To address this, two free-ranging sympatric delphinid species, Australian humpback dolphins (Sousa sahulensis) and Indo-Pacific bottlenose dolphins (Tursiops aduncus), were studied. Biosonar clicks from both species were recorded within the same stretch of coastal habitat in Exmouth Gulf, Western Australia, using a vertical seven element hydrophone array. S. sahulensis used biosonar clicks with a mean source level of 199 ± 3 dB re 1 μPa peak-peak (pp), mean centroid frequency of 106 ± 11 kHz, and emitted at interclick intervals (ICIs) of 79 ± 33 ms. These parameters were similar to click parameters of sympatric T. aduncus, characterized by mean source levels of 204 ± 4 dB re 1 μPa pp, centroid frequency of 112 ± 9 kHz, and ICIs of 73 ± 29 ms. These properties are comparable to those of other similar sized delphinids and suggest that biosonar parameters are independent of sympatric delphinids and possibly driven by body size. The dynamic biosonar behavior of these delphinids may have, consequently, allowed for adaptations to local environments through high levels of control over sonar beam properties.

Biosonar resolving power: echo-acoustic perception of surface structures in the submillimeter range.

PubMed

Simon, Ralph; Knörnschild, Mirjam; Tschapka, Marco; Schneider, Annkathrin; Passauer, Nadine; Kalko, Elisabeth K V; von Helversen, Otto

2014-01-01

The minimum distance for which two points still can be separated from each other defines the resolving power of a visual system. In an echo-acoustic context, the resolving power is usually measured as the smallest perceivable distance of two reflecting surfaces on the range axis and is found to be around half a millimeter for bats employing frequency modulated (FM) echolocation calls. Only few studies measured such thresholds with physical objects, most often bats were trained on virtual echoes i.e., echoes generated and played back by a computer; moreover, bats were sitting while they received the stimuli. In these studies differences in structure depth between 200 and 340 μm were found. However, these low thresholds were never verified for free-flying bats and real physical objects. Here, we show behavioral evidence that the echo-acoustic resolving power for surface structures in fact can be as low as measured for computer generated echoes and even lower, sometimes below 100 μm. We found this exceptional fine discrimination ability only when one of the targets showed spectral interferences in the frequency range of the bats' echolocation call while the other target did not. This result indicates that surface structure is likely to be perceived as a spectral quality rather than being perceived strictly in the time domain. Further, it points out that sonar resolving power directly depends on the highest frequency/shortest wavelength of the signal employed.

Biosonar navigation above water I: estimating flight height.

PubMed

Hoffmann, Susanne; Genzel, Daria; Prosch, Selina; Baier, Leonie; Weser, Sabrina; Wiegrebe, Lutz; Firzlaff, Uwe

2015-02-15

Locomotion and foraging on the wing require precise navigation in more than just the horizontal plane. Navigation in three dimensions and, specifically, precise adjustment of flight height are essential for flying animals. Echolocating bats drink from water surfaces in flight, which requires an exceptionally precise vertical navigation. Here, we exploit this behavior in the bat, Phyllostomus discolor, to understand the biophysical and neural mechanisms that allow for sonar-guided navigation in the vertical plane. In a set of behavioral experiments, we show that for echolocating bats, adjustment of flight height depends on the tragus in their outer ears. Specifically, the tragus imposes elevation-specific spectral interference patterns on the echoes of the bats' sonar emissions. Head-related transfer functions of our bats show that these interference patterns are most conspicuous in the frequency range ∼55 kHz. This conspicuousness is faithfully preserved in the frequency tuning and spatial receptive fields of cortical single and multiunits recorded from anesthetized animals. In addition, we recorded vertical spatiotemporal response maps that describe neural tuning in elevation over time. One class of units that were very sharply tuned to frequencies ∼55 kHz showed unusual spatiotemporal response characteristics with a preference for paired echoes where especially the first echo originates from very low elevations. These behavioral and neural data provide the first insight into biosonar-based processing and perception of acoustic elevation cues that are essential for bats to navigate in three-dimensional space. Copyright © 2015 the American Physiological Society.

Spike Train Similarity Space (SSIMS) Method Detects Effects of Obstacle Proximity and Experience on Temporal Patterning of Bat Biosonar.

PubMed

Accomando, Alyssa W; Vargas-Irwin, Carlos E; Simmons, James A

2018-01-01

Bats emit biosonar pulses in complex temporal patterns that change to accommodate dynamic surroundings. Efforts to quantify these patterns have included analyses of inter-pulse intervals, sonar sound groups, and changes in individual signal parameters such as duration or frequency. Here, the similarity in temporal structure between trains of biosonar pulses is assessed. The spike train similarity space (SSIMS) algorithm, originally designed for neural activity pattern analysis, was applied to determine which features of the environment influence temporal patterning of pulses emitted by flying big brown bats, Eptesicus fuscus . In these laboratory experiments, bats flew down a flight corridor through an obstacle array. The corridor varied in width (100, 70, or 40 cm) and shape (straight or curved). Using a relational point-process framework, SSIMS was able to discriminate between echolocation call sequences recorded from flights in each of the corridor widths. SSIMS was also able to tell the difference between pulse trains recorded during flights where corridor shape through the obstacle array matched the previous trials (fixed, or expected) as opposed to those recorded from flights with randomized corridor shape (variable, or unexpected), but only for the flight path shape in which the bats had previous training. The results show that experience influences the temporal patterns with which bats emit their echolocation calls. It is demonstrated that obstacle proximity to the bat affects call patterns more dramatically than flight path shape.

Acoustic Sensors for Air and Surface Navigation Applications

PubMed Central

Kapoor, Rohan; Ramasamy, Subramanian; Schyndel, Ron Van

2018-01-01

This paper presents the state-of-the-art and reviews the state-of-research of acoustic sensors used for a variety of navigation and guidance applications on air and surface vehicles. In particular, this paper focuses on echolocation, which is widely utilized in nature by certain mammals (e.g., cetaceans and bats). Although acoustic sensors have been extensively adopted in various engineering applications, their use in navigation and guidance systems is yet to be fully exploited. This technology has clear potential for applications in air and surface navigation/guidance for intelligent transport systems (ITS), especially considering air and surface operations indoors and in other environments where satellite positioning is not available. Propagation of sound in the atmosphere is discussed in detail, with all potential attenuation sources taken into account. The errors introduced in echolocation measurements due to Doppler, multipath and atmospheric effects are discussed, and an uncertainty analysis method is presented for ranging error budget prediction in acoustic navigation applications. Considering the design challenges associated with monostatic and multi-static sensor implementations and looking at the performance predictions for different possible configurations, acoustic sensors show clear promises in navigation, proximity sensing, as well as obstacle detection and tracking. The integration of acoustic sensors in multi-sensor navigation systems is also considered towards the end of the paper and a low Size, Weight and Power, and Cost (SWaP-C) sensor integration architecture is presented for possible introduction in air and surface navigation systems. PMID:29414894

Biosonar resolving power: echo-acoustic perception of surface structures in the submillimeter range

PubMed Central

Simon, Ralph; Knörnschild, Mirjam; Tschapka, Marco; Schneider, Annkathrin; Passauer, Nadine; Kalko, Elisabeth K. V.; von Helversen, Otto

2014-01-01

The minimum distance for which two points still can be separated from each other defines the resolving power of a visual system. In an echo-acoustic context, the resolving power is usually measured as the smallest perceivable distance of two reflecting surfaces on the range axis and is found to be around half a millimeter for bats employing frequency modulated (FM) echolocation calls. Only few studies measured such thresholds with physical objects, most often bats were trained on virtual echoes i.e., echoes generated and played back by a computer; moreover, bats were sitting while they received the stimuli. In these studies differences in structure depth between 200 and 340 μm were found. However, these low thresholds were never verified for free-flying bats and real physical objects. Here, we show behavioral evidence that the echo-acoustic resolving power for surface structures in fact can be as low as measured for computer generated echoes and even lower, sometimes below 100 μm. We found this exceptional fine discrimination ability only when one of the targets showed spectral interferences in the frequency range of the bats′ echolocation call while the other target did not. This result indicates that surface structure is likely to be perceived as a spectral quality rather than being perceived strictly in the time domain. Further, it points out that sonar resolving power directly depends on the highest frequency/shortest wavelength of the signal employed. PMID:24616703

Bats coordinate sonar and flight behavior as they forage in open and cluttered environments.

PubMed

Falk, Benjamin; Jakobsen, Lasse; Surlykke, Annemarie; Moss, Cynthia F

2014-12-15

Echolocating bats use active sensing as they emit sounds and listen to the returning echoes to probe their environment for navigation, obstacle avoidance and pursuit of prey. The sensing behavior of bats includes the planning of 3D spatial trajectory paths, which are guided by echo information. In this study, we examined the relationship between active sonar sampling and flight motor output as bats changed environments from open space to an artificial forest in a laboratory flight room. Using high-speed video and audio recordings, we reconstructed and analyzed 3D flight trajectories, sonar beam aim and acoustic sonar emission patterns as the bats captured prey. We found that big brown bats adjusted their sonar call structure, temporal patterning and flight speed in response to environmental change. The sonar beam aim of the bats predicted the flight turn rate in both the open room and the forest. However, the relationship between sonar beam aim and turn rate changed in the forest during the final stage of prey pursuit, during which the bat made shallower turns. We found flight stereotypy developed over multiple days in the forest, but did not find evidence for a reduction in active sonar sampling with experience. The temporal patterning of sonar sound groups was related to path planning around obstacles in the forest. Together, these results contribute to our understanding of how bats coordinate echolocation and flight behavior to represent and navigate their environment. © 2014. Published by The Company of Biologists Ltd.

Bats coordinate sonar and flight behavior as they forage in open and cluttered environments

PubMed Central

Falk, Benjamin; Jakobsen, Lasse; Surlykke, Annemarie; Moss, Cynthia F.

2014-01-01

Echolocating bats use active sensing as they emit sounds and listen to the returning echoes to probe their environment for navigation, obstacle avoidance and pursuit of prey. The sensing behavior of bats includes the planning of 3D spatial trajectory paths, which are guided by echo information. In this study, we examined the relationship between active sonar sampling and flight motor output as bats changed environments from open space to an artificial forest in a laboratory flight room. Using high-speed video and audio recordings, we reconstructed and analyzed 3D flight trajectories, sonar beam aim and acoustic sonar emission patterns as the bats captured prey. We found that big brown bats adjusted their sonar call structure, temporal patterning and flight speed in response to environmental change. The sonar beam aim of the bats predicted the flight turn rate in both the open room and the forest. However, the relationship between sonar beam aim and turn rate changed in the forest during the final stage of prey pursuit, during which the bat made shallower turns. We found flight stereotypy developed over multiple days in the forest, but did not find evidence for a reduction in active sonar sampling with experience. The temporal patterning of sonar sound groups was related to path planning around obstacles in the forest. Together, these results contribute to our understanding of how bats coordinate echolocation and flight behavior to represent and navigate their environment. PMID:25394632

Distribution and morphology of cholinergic, putative catecholaminergic and serotonergic neurons in the brain of the Egyptian rousette flying fox, Rousettus aegyptiacus.

PubMed

Maseko, Busisiwe C; Bourne, James A; Manger, Paul R

2007-11-01

Over the past decade much controversy has surrounded the hypothesis that the megachiroptera, or megabats, share unique neural characteristics with the primates. These observations, which include similarities in visual pathways, have suggested that the megabats are more closely related to the primates than to the other group of the Chiropteran order, the microbats, and suggests a diphyletic origin of the Chiroptera. To contribute data relevant to this debate, we used immunohistochemical techniques to reveal the architecture of the neuromodulatory systems of the Egyptian rousette (Rousettus aegypticus), an echolocating megabat. Our findings revealed many similarities in the nuclear parcellation of the cholinergic, putative catecholaminergic and serotonergic systems with that seen in other mammals including the microbat. However, there were 11 discrete nuclei forming part of these systems in the brain of the megabat studied that were not evident in an earlier study of a microbat. The occurrence of these nuclei align the megabat studied more closely with primates than any other mammalian group and clearly distinguishes them from the microbat, which aligns with the insectivores. The neural systems investigated are not related to such Chiropteran specializations as echolocation, flight, vision or olfaction. If neural characteristics are considered strong indicators of phylogenetic relationships, then the data of the current study strongly supports the diphyletic origin of Chiroptera and aligns the megabat most closely with primates in agreement with studies of other neural characters.

Duration-sensitive neurons in the inferior colliculus of horseshoe bats: adaptations for using CF-FM echolocation pulses.

PubMed

Luo, Feng; Metzner, Walter; Wu, Feijian; Wu, Feijian J; Zhang, Shuyi; Zhang, Shuyi Y; Chen, Qicai; Chen, Qicai C

2008-01-01

The present study examines duration-sensitive neurons in the inferior colliculus (IC) of the least horseshoe bat, Rhinolophus pusillus, from China. In contrast to other bat species tested for duration selectivity so far, echolocation pulses emitted by horseshoe bats are generally longer and composed of a long constant-frequency (CF) component followed by a short downward frequency-modulated (FM) sweep (CF-FM pulse). We used combined CF-FM pulses to analyze the differential effects that these two pulse components had on the duration tuning in neurons of the horseshoe bat's IC. Consistent with results from other mammals, duration-sensitive neurons found in the least horseshoe bat fall into three main classes: short-pass, band-pass, and long-pass. Using a CF stimulus alone, 54% (51/95) of all IC neurons showed at least one form of duration selectivity at one or more stimulus intensities. In 65 of the 95 IC neurons tested with CF pulses, we were also able to test their duration selectivity for a combined CF-FM pulse, which increased the ratio of duration-sensitive neurons to 66% (43/65). Seven to 15 neurons that failed to show duration tuning for CF bursts became duration sensitive for CF-FM pulses, with most of them exhibiting short-pass (depending on stimulus intensity, between 4 and 8 neurons) or band-pass tuning (1-3 neurons). Increasing stimulus intensities did not affect the duration tuning in 53% (23/43) of duration-sensitive neurons for CF bursts and in about 26% (7/27) for CF-FM stimuli. In the remaining neurons, increasing sound levels generally reduced the ratio of duration-sensitive neurons to 33% for CF and 37% for CF-FM stimulation. In those that remained duration sensitive, louder CF bursts shortened best durations in band-pass neurons and cutoff durations in short- and long-pass neurons, whereas louder CF-FM stimuli reduced the cutoff durations only in short-pass neurons. Bandwidths of band-pass neurons were not significantly affected by any stimulus configuration, with only a slight trend for increasing bandwidths for louder CF bursts (but not CF-FM stimuli). Best durations and cutoff durations reached higher values than those in the other bat species examined so far and roughly match the longer durations of echolocation pulses emitted by horseshoe bats. Therefore presentation of a CF-FM stimulus improved the duration sensitivity in IC neurons by increasing the ratio of duration-tuned neurons and making them less susceptible to changes in signal intensity.

Spatio-temporal variation and seasonality of Odontocetes' foraging activity in the leeward side of the island of Hawaii

NASA Astrophysics Data System (ADS)

Giorli, Giacomo; Au, Whitlow W. L.

2017-03-01

The Kona coast of the island of Hawaii hosts many species of odontocetes. These marine mammals are top predators and their foraging activity plays an important role in the ecosystem dynamics. Three passive acoustics recorders were used to study the temporal and spatial occurrence of the foraging activity of odontocetes (excluding beaked and sperm whales) at three locations along the Kona coast of Hawaii between 2012 and 2013. Echolocation clicks were detected using the M3R1

Tempo and mode of antibat ultrasound production and sonar jamming in the diverse hawkmoth radiation.

PubMed

Kawahara, Akito Y; Barber, Jesse R

2015-05-19

The bat-moth arms race has existed for over 60 million y, with moths evolving ultrasonically sensitive ears and ultrasound-producing organs to combat bat predation. The evolution of these defenses has never been thoroughly examined because of limitations in simultaneously conducting behavioral and phylogenetic analyses across an entire group. Hawkmoths include >1,500 species worldwide, some of which produce ultrasound using genital stridulatory structures. However, the function and evolution of this behavior remain largely unknown. We built a comprehensive behavioral dataset of hawkmoth hearing and ultrasonic reply to sonar attack using high-throughput field assays. Nearly half of the species tested (57 of 124 species) produced ultrasound to tactile stimulation or playback of bat echolocation attack. To test the function of ultrasound, we pitted big brown bats (Eptesicus fuscus) against hawkmoths over multiple nights and show that hawkmoths jam bat sonar. Ultrasound production was immediately and consistently effective at thwarting attack and bats regularly performed catching behavior without capturing moths. We also constructed a fossil-calibrated, multigene phylogeny to study the evolutionary history and divergence times of these antibat strategies across the entire family. We show that ultrasound production arose in multiple groups, starting in the late Oligocene (∼ 26 Ma) after the emergence of insectivorous bats. Sonar jamming and bat-detecting ears arose twice, independently, in the Miocene (18-14 Ma) either from earless hawkmoths that produced ultrasound in response to physical contact only, or from species that did not respond to touch or bat echolocation attack.

Echolocation behaviour of the big brown bat (Eptesicus fuscus) in an obstacle avoidance task of increasing difficulty.

PubMed

Sändig, Sonja; Schnitzler, Hans-Ulrich; Denzinger, Annette

2014-08-15

Four big brown bats (Eptesicus fuscus) were challenged in an obstacle avoidance experiment to localize vertically stretched wires requiring progressively greater accuracy by diminishing the wire-to-wire distance from 50 to 10 cm. The performance of the bats decreased with decreasing gap size. The avoidance task became very difficult below a wire separation of 30 cm, which corresponds to the average wingspan of E. fuscus. Two of the bats were able to pass without collisions down to a gap size of 10 cm in some of the flights. The other two bats only managed to master gap sizes down to 20 and 30 cm, respectively. They also performed distinctly worse at all other gap sizes. With increasing difficulty of the task, the bats changed their flight and echolocation behaviour. Especially at gap sizes of 30 cm and below, flight paths increased in height and flight speed was reduced. In addition, the bats emitted approach signals that were arranged in groups. At all gap sizes, the largest numbers of pulses per group were observed in the last group before passing the obstacle. The more difficult the obstacle avoidance task, the more pulses there were in the groups and the shorter the within-group pulse intervals. In comparable situations, the better-performing bats always emitted groups with more pulses than the less well-performing individuals. We hypothesize that the accuracy of target localization increases with the number of pulses per group and that each group is processed as a package. © 2014. Published by The Company of Biologists Ltd.

Spatio-temporal patterns of beaked whale echolocation signals in the North Pacific.

PubMed

Baumann-Pickering, Simone; Roch, Marie A; Brownell, Robert L; Simonis, Anne E; McDonald, Mark A; Solsona-Berga, Alba; Oleson, Erin M; Wiggins, Sean M; Hildebrand, John A

2014-01-01

At least ten species of beaked whales inhabit the North Pacific, but little is known about their abundance, ecology, and behavior, as they are elusive and difficult to distinguish visually at sea. Six of these species produce known species-specific frequency modulated (FM) echolocation pulses: Baird's, Blainville's, Cuvier's, Deraniyagala's, Longman's, and Stejneger's beaked whales. Additionally, one described FM pulse (BWC) from Cross Seamount, Hawai'i, and three unknown FM pulse types (BW40, BW43, BW70) have been identified from almost 11 cumulative years of autonomous recordings at 24 sites throughout the North Pacific. Most sites had a dominant FM pulse type with other types being either absent or limited. There was not a strong seasonal influence on the occurrence of these signals at any site, but longer time series may reveal smaller, consistent fluctuations. Only the species producing BWC signals, detected throughout the Pacific Islands region, consistently showed a diel cycle with nocturnal foraging. By comparing stranding and sighting information with acoustic findings, we hypothesize that BWC signals are produced by ginkgo-toothed beaked whales. BW43 signal encounters were restricted to Southern California and may be produced by Perrin's beaked whale, known only from Californian waters. BW70 signals were detected in the southern Gulf of California, which is prime habitat for Pygmy beaked whales. Hubb's beaked whale may have produced the BW40 signals encountered off central and southern California; however, these signals were also recorded off Pearl and Hermes Reef and Wake Atoll, which are well south of their known range.

Situational and Age-Dependent Decision Making during Life Threatening Distress in Myotis macrodactylus.

PubMed

Huang, Xiaobin; Kanwal, Jagmeet S; Jiang, Tinglei; Long, Zhenyu; Luo, Bo; Yue, Xinke; Gu, Yongbo; Feng, Jiang

2015-01-01

Echolocation and audiovocal communication have been studied extensively in bats. The manner in which these abilities are incorporated within escape behaviors during life-threatening distress is largely unknown. Here we tested the hypothesis that behavioral response profiles expressed during distress are relatively stereotypic given their evolutionary adaptations to avoid predators. We subjected juvenile and adult big-footed myotis (Myotis macrodactylus) to a sequence of three types of life threatening distress: 1) trapping them in a mist-net (environmental threat), 2) approaching them when trapped (predator threat), and 3) partially restraining their freedom to move (arrest), and recorded their escape behavior in each of the three conditions. Response profiles differed across individuals and with the context in which they were expressed. During environmental and predator threat, bats displayed significantly more biting and wing-flapping behaviors and emitted more echolocation pulses than during arrest. Response profiles also varied with age. During arrest, juveniles were more likely than adults to emit distress calls and vice-versa for biting and wing flapping during environmental and predator threat. Overall, individualized response profiles were classified into ten clusters that were aligned along two divergent response trajectories when viewed within two-dimensional, multifactorial decision space. Juvenile behaviors tended to follow a predominantly "social-dependence" trajectory, whereas adult behaviors were mostly aligned along a "self-reliance" trajectory. We conclude that bats modify their vocal behavior and make age-appropriate and contextually adaptive decisions when distressed. This decision-making ability is consistent with observations in other social species, including humans.

Situational and Age-Dependent Decision Making during Life Threatening Distress in Myotis macrodactylus

PubMed Central

Huang, Xiaobin; Kanwal, Jagmeet S.; Jiang, Tinglei; Long, Zhenyu; Luo, Bo; Yue, Xinke; Gu, Yongbo; Feng, Jiang

2015-01-01

Echolocation and audiovocal communication have been studied extensively in bats. The manner in which these abilities are incorporated within escape behaviors during life-threatening distress is largely unknown. Here we tested the hypothesis that behavioral response profiles expressed during distress are relatively stereotypic given their evolutionary adaptations to avoid predators. We subjected juvenile and adult big-footed myotis (Myotis macrodactylus) to a sequence of three types of life threatening distress: 1) trapping them in a mist-net (environmental threat), 2) approaching them when trapped (predator threat), and 3) partially restraining their freedom to move (arrest), and recorded their escape behavior in each of the three conditions. Response profiles differed across individuals and with the context in which they were expressed. During environmental and predator threat, bats displayed significantly more biting and wing-flapping behaviors and emitted more echolocation pulses than during arrest. Response profiles also varied with age. During arrest, juveniles were more likely than adults to emit distress calls and vice-versa for biting and wing flapping during environmental and predator threat. Overall, individualized response profiles were classified into ten clusters that were aligned along two divergent response trajectories when viewed within two-dimensional, multifactorial decision space. Juvenile behaviors tended to follow a predominantly “social-dependence” trajectory, whereas adult behaviors were mostly aligned along a “self-reliance” trajectory. We conclude that bats modify their vocal behavior and make age-appropriate and contextually adaptive decisions when distressed. This decision-making ability is consistent with observations in other social species, including humans. PMID:26181328

Amazon river dolphins (Inia geoffrensis) modify biosonar output level and directivity during prey interception in the wild.

PubMed

Ladegaard, Michael; Jensen, Frants Havmand; Beedholm, Kristian; da Silva, Vera Maria Ferreira; Madsen, Peter Teglberg

2017-07-15

Toothed whales have evolved to live in extremely different habitats and yet they all rely strongly on echolocation for finding and catching prey. Such biosonar-based foraging involves distinct phases of searching for, approaching and capturing prey, where echolocating animals gradually adjust sonar output to actively shape the flow of sensory information. Measuring those outputs in absolute levels requires hydrophone arrays centred on the biosonar beam axis, but this has never been done for wild toothed whales approaching and capturing prey. Rather, field studies make the assumption that toothed whales will adjust their biosonar in the same manner to arrays as they will when approaching prey. To test this assumption, we recorded wild botos ( Inia geoffrensis ) as they approached and captured dead fish tethered to a hydrophone in front of a star-shaped seven-hydrophone array. We demonstrate that botos gradually decrease interclick intervals and output levels during prey approaches, using stronger adjustment magnitudes than predicted from previous boto array data. Prey interceptions are characterised by high click rates, but although botos buzz during prey capture, they do so at lower click rates than marine toothed whales, resulting in a much more gradual transition from approach phase to buzzing. We also demonstrate for the first time that wild toothed whales broaden biosonar beamwidth when closing in on prey, as is also seen in captive toothed whales and bats, thus resulting in a larger ensonified volume around the prey, probably aiding prey tracking by decreasing the risk of prey evading ensonification. © 2017. Published by The Company of Biologists Ltd.

Biosonar navigation above water II: exploiting mirror images.

PubMed

Genzel, Daria; Hoffmann, Susanne; Prosch, Selina; Firzlaff, Uwe; Wiegrebe, Lutz

2015-02-15

As in vision, acoustic signals can be reflected by a smooth surface creating an acoustic mirror image. Water bodies represent the only naturally occurring horizontal and acoustically smooth surfaces. Echolocating bats flying over smooth water bodies encounter echo-acoustic mirror images of objects above the surface. Here, we combined an electrophysiological approach with a behavioral experimental paradigm to investigate whether bats can exploit echo-acoustic mirror images for navigation and how these mirrorlike echo-acoustic cues are encoded in their auditory cortex. In an obstacle-avoidance task where the obstacles could only be detected via their echo-acoustic mirror images, most bats spontaneously exploited these cues for navigation. Sonar ensonifications along the bats' flight path revealed conspicuous changes of the reflection patterns with slightly increased target strengths at relatively long echo delays corresponding to the longer acoustic paths from the mirrored obstacles. Recordings of cortical spatiotemporal response maps (STRMs) describe the tuning of a unit across the dimensions of elevation and time. The majority of cortical single and multiunits showed a special spatiotemporal pattern of excitatory areas in their STRM indicating a preference for echoes with (relative to the setup dimensions) long delays and, interestingly, from low elevations. This neural preference could effectively encode a reflection pattern as it would be perceived by an echolocating bat detecting an object mirrored from below. The current study provides both behavioral and neurophysiological evidence that echo-acoustic mirror images can be exploited by bats for obstacle avoidance. This capability effectively supports echo-acoustic navigation in highly cluttered natural habitats. Copyright © 2015 the American Physiological Society.

Dynamic behavioral strategies during sonar signal emission in roundleaf bats.

PubMed

Feng, Lin; Li, Yitan; Lu, Hongwang

2013-10-02

For echolocating bats which emit biosonar pulses nasally, their nostrils are surrounded by fleshy appendages that diffract the outgoing ultrasonic waves. The posterior leaf, as a prominent part of the noseleaf, was mentioned in previous preliminary observations to move during flight in some species of bats, yet the detailed motion patterns and thus the possible functional role of the posterior leaf movement in biosonar systems remain unclear. In the current work, the motion of the posterior leaf of living pratt's roundleaf bats has been investigated quantitatively. Temporal characterizations of the noseleaf movement and the ultrasonic pulse emission were performed by virtue of synchronized laser vibrometry and sound recording. The results showed that the posterior leaf tilted forwards and restored to original position within tens of milliseconds. Noseleaf motions were temporally correlated with the emitted ultrasonic pulses. The surfaces of the posterior leaf were moving in the anterior direction in most of the pulse duration. The bats were able to switch the motions on or off. From the comparison with the previously reported noseleaf dynamics in horseshoe bat, we find similar ratio sizes and displacements of the noseleaves compared to the used wavelengths, implying that similar behavioral strategies are utilized by species of bats and it may be applied to different components of the signal emitting apparatus. It suggests that the dynamic sensing principles may widely play a role in the biosonar systems and the investigation on time-variant mechanisms is of capital importance to understand the biosonar sensing strategies used by echolocating bats. © 2013.

Forest structure affects trophic linkages: How silvicultural disturbance impacts bats and their insect prey

USGS Publications Warehouse

Dodd, L.E.; Lacki, M.J.; Britzke, E.R.; Buehler, D.A.; Keyser, P.D.; Larkin, J.L.; Rodewald, A.D.; Wigley, T.B.; Wood, P.B.; Rieske, L.K.

2012-01-01

Vertebrate insectivores such as bats are a pervasive top-down force on prey populations in forest ecosystems. Conservation focusing on forest-dwelling bats requires understanding of community-level interactions between these predators and their insect prey. Our study assessed bat activity and insect occurrence (abundance and diversity) across a gradient of forest disturbance and structure (silvicultural treatments) in the Central Appalachian region of North America. We conducted acoustic surveys of bat echolocation concurrent with insect surveys using blacklight and malaise traps over 2 years. Predator activity, prey occurrence and prey biomass varied seasonally and across the region. The number of bat echolocation pulses was positively related with forest disturbance, whereas prey demonstrated varied trends. Lepidopteran abundance was negatively related with disturbance, while dipteran abundance and diversity was positively related with disturbance. Coleoptera were unaffected. Neither bat nor insect response variables differed between plot interiors and edges. Correlations between bat activity and vegetative structure reflected differences in foraging behavior among ensembles. Activity of myotine bats was correlated with variables describing sub-canopy vegetation, whereas activity of lasiurine bats was more closely correlated with canopy-level vegetation. Lepidopteran abundance was correlated with variables describing canopy and sub-canopy vegetation, whereas coleopteran and dipteran occurrence were more closely correlated with canopy-level vegetative structure. Our study demonstrates regional variation in bat activity and prey occurrence across a forested disturbance gradient. Land management and conservation efforts should consider the importance of vegetation structure and plant species richness to sustain forest-dwelling bats and their insect prey.

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

PubMed

Ghose, Kaushik; Moss, Cynthia F

2006-02-08

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.

Sequential assessment of prey through the use of multiple sensory cues by an eavesdropping bat

NASA Astrophysics Data System (ADS)

Page, Rachel A.; Schnelle, Tanja; Kalko, Elisabeth K. V.; Bunge, Thomas; Bernal, Ximena E.

2012-06-01

Predators are often confronted with a broad diversity of potential prey. They rely on cues associated with prey quality and palatability to optimize their hunting success and to avoid consuming toxic prey. Here, we investigate a predator's ability to assess prey cues during capture, handling, and consumption when confronted with conflicting information about prey quality. We used advertisement calls of a preferred prey item (the túngara frog) to attract fringe-lipped bats, Trachops cirrhosus, then offered palatable, poisonous, and chemically manipulated anurans as prey. Advertisement calls elicited an attack response, but as bats approached, they used additional sensory cues in a sequential manner to update their information about prey size and palatability. While both palatable and poisonous small anurans were readily captured, large poisonous toads were approached but not contacted suggesting the use of echolocation for assessment of prey size at close range. Once prey was captured, bats used chemical cues to make final, post-capture decisions about whether to consume the prey. Bats dropped small, poisonous toads as well as palatable frogs coated in toad toxins either immediately or shortly after capture. Our study suggests that echolocation and chemical cues obtained at close range supplement information obtained from acoustic cues at long range. Updating information about prey quality minimizes the occurrence of costly errors and may be advantageous in tracking temporal and spatial fluctuations of prey and exploiting novel food sources. These findings emphasize the sequential, complex nature of prey assessment that may allow exploratory and flexible hunting behaviors.

Migratory bats respond to artificial green light with positive phototaxis.

PubMed

Voigt, Christian C; Roeleke, Manuel; Marggraf, Lara; Pētersons, Gunārs; Voigt-Heucke, Silke L

2017-01-01

Artificial light at night is spreading worldwide at unprecedented rates, exposing strictly nocturnal animals such as bats to a novel anthropogenic stressor. Previous studies about the effect of artificial light on bats focused almost exclusively on non-migratory species, yet migratory animals such as birds are known to be largely affected by light pollution. Thus, we conducted a field experiment to evaluate if bat migration is affected by artificial light at night. In late summer, we presented artificial green light of 520 nm wavelength to bats that were migrating south along the shoreline of the Baltic Sea. Using a light on-off treatment, we observed that the activity of Pipistrellus nathusii and P. pygmaeus, the two most abundant migratory species at our site, increased by more than 50% in the light-on compared to the light-off treatment. We observed an increased number of feeding buzzes during the light-on compared to the light-off treatment for P. nathusii. However, feeding activity was low in general and did not increase disproportionately during the light-on treatment in relation to the overall echolocation call activity of bats. Further, P. nathusii were attracted towards the green light at a distance of about 23 m, which is way beyond the echolocation detection range for insects of Nathusius' bats. We therefore infer that migratory bats were not attracted to artificial green light because of high insect densities, but instead by positive phototaxis. We conclude that artificial light at night may potentially impact bat migration in a yet unrecognized way.

Spatial and Temporal Variations in the Occurrence and Foraging Activity of Coastal Dolphins in Menai Bay, Zanzibar, Tanzania.

PubMed

Temple, Andrew J; Tregenza, Nick; Amir, Omar A; Jiddawi, Narriman; Berggren, Per

2016-01-01

Understanding temporal patterns in distribution, occurrence and behaviour is vital for the effective conservation of cetaceans. This study used cetacean click detectors (C-PODs) to investigate spatial and temporal variation in occurrence and foraging activity of the Indo-Pacific bottlenose (Tursiops aduncus) and Indian Ocean humpback (Sousa plumbea) dolphins resident in the Menai Bay Conservation Area (MBCA), Zanzibar, Tanzania. Occurrence was measured using detection positive minutes. Inter-click intervals were used to identify terminal buzz vocalisations, allowing for analysis of foraging activity. Data were analysed in relation to spatial (location) and temporal (monsoon season, diel phase and tidal phase) variables. Results showed significantly increased occurrence and foraging activity of dolphins in southern areas and during hours of darkness. Higher occurrence at night was not explained by diel variation in echolocation rate and so were considered representative of occurrence patterns. Both tidal phase and monsoon season influenced occurrence but results varied among sites, with no general patterns found. Foraging activity was greatest during hours of darkness, High water and Flood tidal phases. Comparisons of echolocation data among sites suggested differences in the broadband click spectra of MBCA dolphins, possibly indicative of species differences. These dolphin populations are threatened by unsustainable fisheries bycatch and tourism activities. The spatial and temporal patterns identified in this study have implications for future conservation and management actions with regards to these two threats. Further, the results indicate future potential for using passive acoustics to identify and monitor the occurrence of these two species in areas where they co-exist.

Spatio-Temporal Patterns of Beaked Whale Echolocation Signals in the North Pacific

PubMed Central

Baumann-Pickering, Simone; Roch, Marie A.; Brownell Jr, Robert L.; Simonis, Anne E.; McDonald, Mark A.; Solsona-Berga, Alba; Oleson, Erin M.; Wiggins, Sean M.; Hildebrand, John A.

2014-01-01

At least ten species of beaked whales inhabit the North Pacific, but little is known about their abundance, ecology, and behavior, as they are elusive and difficult to distinguish visually at sea. Six of these species produce known species-specific frequency modulated (FM) echolocation pulses: Baird’s, Blainville’s, Cuvier’s, Deraniyagala’s, Longman’s, and Stejneger’s beaked whales. Additionally, one described FM pulse (BWC) from Cross Seamount, Hawai’i, and three unknown FM pulse types (BW40, BW43, BW70) have been identified from almost 11 cumulative years of autonomous recordings at 24 sites throughout the North Pacific. Most sites had a dominant FM pulse type with other types being either absent or limited. There was not a strong seasonal influence on the occurrence of these signals at any site, but longer time series may reveal smaller, consistent fluctuations. Only the species producing BWC signals, detected throughout the Pacific Islands region, consistently showed a diel cycle with nocturnal foraging. By comparing stranding and sighting information with acoustic findings, we hypothesize that BWC signals are produced by ginkgo-toothed beaked whales. BW43 signal encounters were restricted to Southern California and may be produced by Perrin’s beaked whale, known only from Californian waters. BW70 signals were detected in the southern Gulf of California, which is prime habitat for Pygmy beaked whales. Hubb’s beaked whale may have produced the BW40 signals encountered off central and southern California; however, these signals were also recorded off Pearl and Hermes Reef and Wake Atoll, which are well south of their known range. PMID:24465877

Kinesonde observations of ionosphere modification by intense electromagnetic fields from Platteville, Colorado.

NASA Technical Reports Server (NTRS)

Wright, J. W.

1973-01-01

Observations by the Kinesonde (a multifrequency, spaced antenna, digitized complex-amplitude radio sounding system) of ionospheric responses to excitation by the high-power transmitter at Platteville, Colo., are described. Increases of echo scintillation rate and intensity at frequencies reflected near and far from the excitation level are shown. Significant onset delays of these responses suggest disturbance propagation velocities of a few kilometers per second. Calculated echolocations show a time-dependent development toward the excitation region, again with a delayed response. Comments are offered regarding the relative utility of ionogram and Kinesonde observations for study of these phenomena.

An acoustic survey of beaked whales at Cross Seamount near Hawaii.

PubMed

McDonald, Mark A; Hildebrand, John A; Wiggins, Sean M; Johnston, David W; Polovina, Jeffrey J

2009-02-01

An acoustic record from Cross Seamount, southwest of Hawaii, revealed sounds characteristic of beaked whale echolocation at the same relative abundance year-around (270 of 356 days), occurring almost entirely at night. The most common sound had a linear frequency upsweep from 35 to 100 kHz (the bandwidth of recording), an interpulse interval of 0.11 s, and duration of at least 932 mus. A less common upsweep sound with shorter interpulse interval and slower sweep rate was also present. Sounds matching Cuvier's beaked whale were not detected, and Blainville's beaked whale sounds were detected on only one occasion.

The Genomes of Two Bat Species with Long Constant Frequency Echolocation Calls.

PubMed

Dong, Dong; Lei, Ming; Hua, Panyu; Pan, Yi-Hsuan; Mu, Shuo; Zheng, Guantao; Pang, Erli; Lin, Kui; Zhang, Shuyi

2017-01-01

Bats can perceive the world by using a wide range of sensory systems, and some of the systems have become highly specialized, such as auditory sensory perception. Among bat species, the Old World leaf-nosed bats and horseshoe bats (rhinolophoid bats) possess the most sophisticated echolocation systems. Here, we reported the whole-genome sequencing and de novo assembles of two rhinolophoid bats-the great leaf-nosed bat (Hipposideros armiger) and the Chinese rufous horseshoe bat (Rhinolophus sinicus). Comparative genomic analyses revealed the adaptation of auditory sensory perception in the rhinolophoid bat lineages, probably resulting from the extreme selectivity used in the auditory processing by these bats. Pseudogenization of some vision-related genes in rhinolophoid bats was observed, suggesting that these genes have undergone relaxed natural selection. An extensive contraction of olfactory receptor gene repertoires was observed in the lineage leading to the common ancestor of bats. Further extensive gene contractions can be observed in the branch leading to the rhinolophoid bats. Such concordance suggested that molecular changes at one sensory gene might have direct consequences for genes controlling for other sensory modalities. To characterize the population genetic structure and patterns of evolution, we re-sequenced the genome of 20 great leaf-nosed bats from four different geographical locations of China. The result showed similar sequence diversity values and little differentiation among populations. Moreover, evidence of genetic adaptations to high altitudes in the great leaf-nosed bats was observed. Taken together, our work provided a useful resource for future research on the evolution of bats. © The Author 2016. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.

A comparison of survey methods for documenting presence of Myotis leibii (Eastern Small-Footed Bats) at roosting areas in Western Virginia

USGS Publications Warehouse

Huth, John K.; Silvis, Alexander; Moosman, Paul R.; Ford, W. Mark; Sweeten, Sara E.

2015-01-01

Many aspects of foraging and roosting habitat of Myotis leibii (Eastern Small-Footed Bat), an emergent rock roosting-obligate, are poorly described. Previous comparisons of effectiveness of acoustic sampling and mist-net captures have not included Eastern Small-Footed Bat. Habitat requirements of this species differ from congeners in the region, and it is unclear whether survey protocols developed for other species are applicable. Using data from three overlapping studies at two sampling sites in western Virginia’s central Appalachian Mountains, detection probabilities were examined for three survey methods (acoustic surveys with automated identification of calls, visual searches of rock crevices, and mist-netting) for use in the development of “best practices” for future surveys and monitoring. Observer effects were investigated using an expanded version of visual search data. Results suggested that acoustic surveys with automated call identification are not effective for documenting presence of Eastern Small-Footed Bats on talus slopes (basal detection rate of 0%) even when the species is known to be present. The broadband, high frequency echolocation calls emitted by Eastern Small-Footed Bat may be prone to attenuation by virtue of their high frequencies, and these factors, along with signal reflection, lower echolocation rates or possible misidentification to other bat species over talus slopes may all have contributed to poor acoustic survey success. Visual searches and mist-netting of emergent rock had basal detection probabilities of 91% and 75%, respectively. Success of visual searches varied among observers, but detection probability improved with practice. Additionally, visual searches were considerably more economical than mist-netting.

Optimal Predator Risk Assessment by the Sonar-Jamming Arctiine Moth Bertholdia trigona

PubMed Central

Corcoran, Aaron J.; Wagner, Ryan D.; Conner, William E.

2013-01-01

Nearly all animals face a tradeoff between seeking food and mates and avoiding predation. Optimal escape theory holds that an animal confronted with a predator should only flee when benefits of flight (increased survival) outweigh the costs (energetic costs, lost foraging time, etc.). We propose a model for prey risk assessment based on the predator's stage of attack. Risk level should increase rapidly from when the predator detects the prey to when it commits to the attack. We tested this hypothesis using a predator – the echolocating bat – whose active biosonar reveals its stage of attack. We used a prey defense – clicking used for sonar jamming by the tiger moth Bertholdia trigona– that can be readily studied in the field and laboratory and is enacted simultaneously with evasive flight. We predicted that prey employ defenses soon after being detected and targeted, and that prey defensive thresholds discriminate between legitimate predatory threats and false threats where a nearby prey is attacked. Laboratory and field experiments using playbacks of ultrasound signals and naturally behaving bats, respectively, confirmed our predictions. Moths clicked soon after bats detected and targeted them. Also, B. trigona clicking thresholds closely matched predicted optimal thresholds for discriminating legitimate and false predator threats for bats using search and approach phase echolocation – the period when bats are searching for and assessing prey. To our knowledge, this is the first quantitative study to correlate the sensory stimuli that trigger defensive behaviors with measurements of signals provided by predators during natural attacks in the field. We propose theoretical models for explaining prey risk assessment depending on the availability of cues that reveal a predator's stage of attack. PMID:23671686

The diel rhythms of biosonar behavior in the Yangtze finless porpoise (Neophocaena asiaeorientalis asiaeorientalis) in the port of the Yangtze River: The correlation between prey availability and boat traffic.

PubMed

Wang, Zhitao; Akamatsu, Tomonari; Wang, Kexiong; Wang, Ding

2014-01-01

Information on the habitat use of the critically endangered Yangtze finless porpoise (Neophocaena asiaeorientalis asiaeorientalis) is critical for its conservation. The diel biosonar behavior of the porpoise in the port areas of the Yangtze River was examined along with simultaneous observations of fish density and boat traffic. Biosonar pulses from the porpoises were detected for 1233 min (5.77%) over a 21,380 min duration of effective observations. In total, 190 (5.63%) buzzes (an indication of prey capture attempts) were recorded among the 3372 identified click trains. Of the 168 echolocation encounters (bouts of click trains less than eight min apart), 150 (89.3%) involved single animals, indicating that solitary porpoises were frequently present and feeding in the port areas. Significant diel patterns were evident involving the biosonar behavior of the porpoises (including click trains and buzzes), fish density and boat traffic. The frequencies of the click trains and buzzes were significantly lower during the day than in the evening and at night, which suggests that porpoises in this region are primarily engaged in crepuscular and nocturnal foraging. The lack of a significant diel pattern in the echolocation encounters indicates the importance of the port in porpoise conservation. A forced feeding schedule may be associated with the lack of a significant correlation between porpoise acoustics and boat traffic. Overall, prey availability appears to be the primary factor that attracts porpoises. Additionally, porpoises tend to migrate or remain downstream in the morning and migrate or remain upstream in the evening, most likely to follow their prey. The findings of this study can be used to improve the conservation of the Yangtze finless porpoise.

The Diel Rhythms of Biosonar Behavior in the Yangtze Finless Porpoise (Neophocaena asiaeorientalis asiaeorientalis) in the Port of the Yangtze River: The Correlation between Prey Availability and Boat Traffic

PubMed Central

Wang, Zhitao; Akamatsu, Tomonari; Wang, Kexiong; Wang, Ding

2014-01-01

Information on the habitat use of the critically endangered Yangtze finless porpoise (Neophocaena asiaeorientalis asiaeorientalis) is critical for its conservation. The diel biosonar behavior of the porpoise in the port areas of the Yangtze River was examined along with simultaneous observations of fish density and boat traffic. Biosonar pulses from the porpoises were detected for 1233 min (5.77%) over a 21,380 min duration of effective observations. In total, 190 (5.63%) buzzes (an indication of prey capture attempts) were recorded among the 3372 identified click trains. Of the 168 echolocation encounters (bouts of click trains less than eight min apart), 150 (89.3%) involved single animals, indicating that solitary porpoises were frequently present and feeding in the port areas. Significant diel patterns were evident involving the biosonar behavior of the porpoises (including click trains and buzzes), fish density and boat traffic. The frequencies of the click trains and buzzes were significantly lower during the day than in the evening and at night, which suggests that porpoises in this region are primarily engaged in crepuscular and nocturnal foraging. The lack of a significant diel pattern in the echolocation encounters indicates the importance of the port in porpoise conservation. A forced feeding schedule may be associated with the lack of a significant correlation between porpoise acoustics and boat traffic. Overall, prey availability appears to be the primary factor that attracts porpoises. Additionally, porpoises tend to migrate or remain downstream in the morning and migrate or remain upstream in the evening, most likely to follow their prey. The findings of this study can be used to improve the conservation of the Yangtze finless porpoise. PMID:24823945

Trawling bats exploit an echo-acoustic ground effect

PubMed Central

Zsebok, Sandor; Kroll, Ferdinand; Heinrich, Melina; Genzel, Daria; Siemers, Björn M.; Wiegrebe, Lutz

2013-01-01

A water surface acts not only as an optic mirror but also as an acoustic mirror. Echolocation calls emitted by bats at low heights above water are reflected away from the bat, and hence the background clutter is reduced. Moreover, targets on the surface create an enhanced echo. Here, we formally quantified the effect of the surface and target height on both target detection and -discrimination in a combined laboratory and field approach with Myotis daubentonii. In a two-alternative, forced-choice paradigm, the bats had to detect a mealworm and discriminate it from an inedible dummy (20 mm PVC disc). Psychophysical performance was measured as a function of height above either smooth surfaces (water or PVC) or above a clutter surface (artificial grass). At low heights above the clutter surface (10, 20, or 35 cm), the bats' detection performance was worse than above a smooth surface. At a height of 50 cm, the surface structure had no influence on target detection. Above the clutter surface, also target discrimination was significantly impaired with decreasing target height. A detailed analysis of the bats' echolocation calls during target approach shows that above the clutter surface, the bats produce calls with significantly higher peak frequency. Flight-path reconstruction revealed that the bats attacked an target from below over water but from above over a clutter surface. These results are consistent with the hypothesis that trawling bats exploit an echo-acoustic ground effect, in terms of a spatio-temporal integration of direct reflections with indirect reflections from the water surface, to optimize prey detection and -discrimination not only for prey on the water but also for some range above. PMID:23576990

Ultrasonic predator-prey interactions in water-convergent evolution with insects and bats in air?

PubMed

Wilson, Maria; Wahlberg, Magnus; Surlykke, Annemarie; Madsen, Peter Teglberg

2013-01-01

Toothed whales and bats have independently evolved biosonar systems to navigate and locate and catch prey. Such active sensing allows them to operate in darkness, but with the potential cost of warning prey by the emission of intense ultrasonic signals. At least six orders of nocturnal insects have independently evolved ears sensitive to ultrasound and exhibit evasive maneuvers when exposed to bat calls. Among aquatic prey on the other hand, the ability to detect and avoid ultrasound emitting predators seems to be limited to only one subfamily of Clupeidae: the Alosinae (shad and menhaden). These differences are likely rooted in the different physical properties of air and water where cuticular mechanoreceptors have been adapted to serve as ultrasound sensitive ears, whereas ultrasound detection in water have called for sensory cells mechanically connected to highly specialized gas volumes that can oscillate at high frequencies. In addition, there are most likely differences in the risk of predation between insects and fish from echolocating predators. The selection pressure among insects for evolving ultrasound sensitive ears is high, because essentially all nocturnal predation on flying insects stems from echolocating bats. In the interaction between toothed whales and their prey the selection pressure seems weaker, because toothed whales are by no means the only marine predators placing a selection pressure on their prey to evolve specific means to detect and avoid them. Toothed whales can generate extremely intense sound pressure levels, and it has been suggested that they may use these to debilitate prey. Recent experiments, however, show that neither fish with swim bladders, nor squid are debilitated by such signals. This strongly suggests that the production of high amplitude ultrasonic clicks serve the function of improving the detection range of the toothed whale biosonar system rather than debilitation of prey.

Sensory Ecology of Water Detection by Bats: A Field Experiment

PubMed Central

Russo, Danilo; Cistrone, Luca; Jones, Gareth

2012-01-01

Bats face a great risk of dehydration, so sensory mechanisms for water recognition are crucial for their survival. In the laboratory, bats recognized any smooth horizontal surface as water because these provide analogous reflections of echolocation calls. We tested whether bats also approach smooth horizontal surfaces other than water to drink in nature by partly covering watering troughs used by hundreds of bats with a Perspex layer mimicking water. We aimed 1) to confirm that under natural conditions too bats mistake any horizontal smooth surface for water by testing this on large numbers of individuals from a range of species and 2) to assess the occurrence of learning effects. Eleven bat species mistook Perspex for water relying chiefly on echoacoustic information. Using black instead of transparent Perspex did not deter bats from attempting to drink. In Barbastella barbastellus no echolocation differences occurred between bats approaching the water and the Perspex surfaces respectively, confirming that bats perceive water and Perspex to be acoustically similar. The drinking attempt rates at the fake surface were often lower than those recorded in the laboratory: bats then either left the site or moved to the control water surface. This suggests that bats modified their behaviour as soon as the lack of drinking reward had overridden the influence of echoacoustic information. Regardless of which of two adjoining surfaces was covered, bats preferentially approached and attempted to drink from the first surface encountered, probably because they followed a common route, involving spatial memory and perhaps social coordination. Overall, although acoustic recognition itself is stereotyped and its importance in the drinking process overwhelming, our findings point at the role of experience in increasing behavioural flexibility under natural conditions. PMID:23133558

Effects of sound intensity on temporal properties of inhibition in the pallid bat auditory cortex.

PubMed

Razak, Khaleel A

2013-01-01

Auditory neurons in bats that use frequency modulated (FM) sweeps for echolocation are selective for the behaviorally-relevant rates and direction of frequency change. Such selectivity arises through spectrotemporal interactions between excitatory and inhibitory components of the receptive field. In the pallid bat auditory system, the relationship between FM sweep direction/rate selectivity and spectral and temporal properties of sideband inhibition have been characterized. Of note is the temporal asymmetry in sideband inhibition, with low-frequency inhibition (LFI) exhibiting faster arrival times compared to high-frequency inhibition (HFI). Using the two-tone inhibition over time (TTI) stimulus paradigm, this study investigated the interactions between two sound parameters in shaping sideband inhibition: intensity and time. Specifically, the impact of changing relative intensities of the excitatory and inhibitory tones on arrival time of inhibition was studied. Using this stimulation paradigm, single unit data from the auditory cortex of pentobarbital-anesthetized cortex show that the threshold for LFI is on average ~8 dB lower than HFI. For equal intensity tones near threshold, LFI is stronger than HFI. When the inhibitory tone intensity is increased further from threshold, the strength asymmetry decreased. The temporal asymmetry in LFI vs. HFI arrival time is strongest when the excitatory and inhibitory tones are of equal intensities or if excitatory tone is louder. As inhibitory tone intensity is increased, temporal asymmetry decreased suggesting that the relative magnitude of excitatory and inhibitory inputs shape arrival time of inhibition and FM sweep rate and direction selectivity. Given that most FM bats use downward sweeps as echolocation calls, a similar asymmetry in threshold and strength of LFI vs. HFI may be a general adaptation to enhance direction selectivity while maintaining sweep-rate selective responses to downward sweeps.

Acoustic communication in plant-animal interactions.

PubMed

Schöner, Michael G; Simon, Ralph; Schöner, Caroline R

2016-08-01

Acoustic communication is widespread and well-studied in animals but has been neglected in other organisms such as plants. However, there is growing evidence for acoustic communication in plant-animal interactions. While knowledge about active acoustic signalling in plants (i.e. active sound production) is still in its infancy, research on passive acoustic signalling (i.e. reflection of animal sounds) revealed that bat-dependent plants have adapted to the bats' echolocation systems by providing acoustic reflectors to attract their animal partners. Understanding the proximate mechanisms and ultimate causes of acoustic communication will shed light on an underestimated dimension of information transfer between plants and animals. Copyright © 2016 Elsevier Ltd. All rights reserved.

Hearing Sensation Changes When a Warning Predicts a Loud Sound in the False Killer Whale (Pseudorca crassidens).

PubMed

Nachtigall, Paul E; Supin, Alexander Y

2016-01-01

Stranded whales and dolphins have sometimes been associated with loud anthropogenic sounds. Echolocating whales produce very loud sounds themselves and have developed the ability to protect their hearing from their own signals. A false killer whale's hearing sensitivity was measured when a faint warning sound was given just before the presentation of an increase in intensity to 170 dB. If the warning occurred within 1-9 s, as opposed to 20-40 s, the whale showed a 13-dB reduction in hearing sensitivity. Warning sounds before loud pulses may help mitigate the effects of loud anthropogenic sounds on wild animals.