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  • 1
    Electronic Resource
    Electronic Resource
    Springer
    Journal of comparative physiology 186 (2000), S. 413-423 
    ISSN: 1432-1351
    Keywords: Key words Medial superior olive ; Interaural time ; difference ; Receptive field ; Precedence effect ; Temporal processing
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Medicine
    Notes: Abstract Traditionally, the medial superior olive, a mammalian auditory brainstem structure, is considered to encode interaural time differences, the main cue for localizing low-frequency sounds. Detection of binaural excitatory and inhibitory inputs are considered as an underlying mechanism. Most small mammals, however, hear high frequencies well beyond 50 kHz and have small interaural distances. Therefore, they can not use interaural time differences for sound localization and yet possess a medial superior olive. Physiological studies in bats revealed that medial superior olive cells show similar interaural time difference coding as in larger mammals tuned to low-frequency hearing. Their interaural time difference sensitivity, however, is far too coarse to serve in sound localization. Thus, interaural time difference sensitivity in medial superior olive of small mammals is an epiphenomenon. We propose that the original function of the medial superior olive is a binaural cooperation causing facilitation due to binaural excitation. Lagging inhibitory inputs, however, suppress reverberations and echoes from the acoustic background. Thereby, generation of antagonistically organized temporal fields is the basic and original function of the mammalian medial superior olive. Only later in evolution with the advent of larger mammals did interaural distances, and hence interaural time differences, became large enough to be used as cues for sound localization of low-frequency stimuli.
    Type of Medium: Electronic Resource
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  • 2
    Electronic Resource
    Electronic Resource
    Springer
    Journal of comparative physiology 74 (1971), S. 153-155 
    ISSN: 1432-1351
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Medicine
    Notes: Summary Collicular evoked potentials in Rhinolophus ferrum equinum show very prominent responses to the final frequency modulated part of a acoustic stimulus, simulating the natural echolocation sound.
    Type of Medium: Electronic Resource
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  • 3
    Electronic Resource
    Electronic Resource
    Springer
    Journal of comparative physiology 125 (1978), S. 217-225 
    ISSN: 1432-1351
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Medicine
    Notes: Summary The responses of 230 single neurons in the inferior colliculus of the horseshoebat to single tones have been studied, emphasizing systematic analysis of the effective frequency bands, dynamic properties and the time course of responses. Distribution of the units' best excitatory frequencies (BEF) is: low frequency neurons 23% (BEF 3–65 kHz); FM-frequency neurons 25% (BEF 65–81 kHz, i.e., frequencies occurring in the FM-part of the bat's echo signal); filter neurons 45% (BEF 81–88 kHz, i.e., frequencies occurring in the stabilized CF-part of the bat's echo=reference frequency (RF)); high frequency neurons about 7% (BEF 〉 88 kHz). Tuning curves show conventional shapes (Fig. 1), apart from those of filter neurons, which are extremely narrow. Accordingly, Q10dB-values (BEF divided by the bandwidth of the tuning curve at 10 dB above threshold) are 80–450 in filter neurons (Fig. 2). Response patterns (Fig. 3) are similar to those of Nucleus cochlearis units (transient, sustained, negative and complex responders) with an increased percentage of complex responders up to 38% and a decreased number of transient responders. All types of spike-count functions are found (Fig. 4); nonmonotonic ones dominating. Maximal spike counts are not at the BEF but a few kHz below. Distinct upper thresholds, especially at the BEF of filter neurons (Fig. 5) lead to abrupt changes in activity by slightly shifting stimulus frequency or intensity. The hallmark of inferior colliculus neurons is inhibition, disclosed by distinct inhibitory areas enfolding and overlapping excitatory ones (Figs. 3 and 5). Duration of inhibition varies with stimulus frequency, but is largely independent of stimulus duration (Fig. 6), whereas rebound of inhibition depends on stimulus duration building up periodic rebound activities, if stimulus duration is lengthened. In addition, there are neurons responding only periodically, regardless of stimulus frequency and intensity (Fig. 7). Inhibition is discussed in terms of improving the neuronal signal/spontaneous noise ratio and altering responsiveness of neurons after stimulation, so that these neurons may be suited to time processing in the acoustic pathway.
    Type of Medium: Electronic Resource
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  • 4
    Electronic Resource
    Electronic Resource
    Springer
    Journal of comparative physiology 163 (1988), S. 271-285 
    ISSN: 1432-1351
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Medicine
    Notes: Summary The tonotopic organization of the inferior colliculus (IC) in two echolocating bats,Hipposideros speoris andMegaderma lyra, was studied by multiunit recordings. InHipposideros speoris frequencies below the range of the echolocation signals (i.e. below 120 kHz) are compressed into a dorsolateral cap about 400–600 Μm thick. Within this region, neuronal sheets of about 4–5 Μm thickness represent a 1 kHz-band. In contrast, the frequencies of the echolocation signals (120–140 kHz) are overrepresented and occupy the central and ventral parts of the IC (Fig. 3). In this region, neuronal sheets of about 80 Μm thickness represent a 1 kHz-band. The largest 1 kHz-slabs (400–600 Μm) represent frequencies of the pure tone components of the echolocation signals (130–140 kHz). The frequency of the pure tone echolocation component is specific for any given individual and always part of the overrepresented frequency range but did not necessarily coincide with the BF of the thickest isofrequency slab. Thus hipposiderid bats have an auditory fovea (Fig. 10). In the IC ofMegaderma lyra the complete range of audible frequencies, from a few kHz to 110 kHz, is represented in fairly equal proportions (Fig. 7). On the average, a neuronal sheet of 30 Μm thickness is dedicated to a 1 kHz-band, however, frequencies below 20 kHz, i.e. below the range of the echolocation signals, are overrepresented. Audiograms based on thresholds determined from multiunit recordings demonstrate the specific sensitivities of the two bat species. InHipposideros speoris the audiogram shows two sensitivity peaks, one in the nonecholocating frequency range (10–60 kHz) and one within the auditory fovea for echolocation (130–140 kHz).Megaderma lyra has extreme sensitivity between 15–20 kHz, with thresholds as low as −24 dB SPL, and a second sensitivity peak at 50 kHz (Fig. 8). InMegaderma lyra, as in common laboratory mammals, Q10dB-values of single units do not exceed 30, whereas inHipposideros speoris units with BFs within the auditory fovea reach Q10dB-values of up to 130. InMegaderma lyra, many single units and multiunit clusters with BFs below 30 kHz show upper thresholds of 40–50 dB SPL and respond most vigorously to sound intensities below 30 dB SPL (Fig. 9). Many of these units respond preferentially or exclusively to noise. These features are interpreted as adaptations to detection of prey-generated noises. The two different tonotopic arrangements (compare Figs. 3 and 7) in the ICs of the two species are correlated with their different foraging behaviours. It is suggested that pure tone echolocation and auditory foveae are primarily adaptations to echo clutter rejection for species foraging on the wing close to vegetation.
    Type of Medium: Electronic Resource
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  • 5
    Electronic Resource
    Electronic Resource
    Springer
    Journal of comparative physiology 152 (1983), S. 421-432 
    ISSN: 1432-1351
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Medicine
    Notes: Summary 1. When catching flying prey under laboratory conditionsRhinolophus ferrumequinum typically emit FM-CF-FM signals (Fig. 2). Except for the last two sounds during approach and final buzz the FM-parts are fainter than the CF-component by a factor of 0.76+-14 (Fig. 1). The final FM-part was undetectable in some signals emitted during approach (Fig. 3). 2. In obstacle avoidance flights both, preceding and final FM-parts are prominent and louder than the CF-part by a factor of 1.14 to 1.63 (Fig. 1 and 3). Bandwidths of the FM-components increased from ca. 3.5 to 12 kHz for the starting and from 12 to 20 kHz on the average for the final FM-sweep (Table 1). 3. In the open field during cruising flightsNyctalus noctula emits pure tones of 22.5 to 25.0 kHz without any frequency modulated components and a duration of 10 to 50 ms (Fig. 4). Brief frequency modulated signals sweeping from ca. 50 to 20 kHz in about 1–2 ms are emitted during pursuit of prey (Fig. 4). 4. Under laboratory conditionsNyctalus noctula does not emit pure tones and is not able to catch flying prey in a flight chamber 10.5×3.5×2.15 m in size. During flights towards a landing platformNyctalus noctula invariably emits brief frequency modulated pulses. During an individual flight the structure is not changed (Fig. 5). 5. InNyctalus noctula specific features of the echolocation pulses, e.g. frequency range swept through, presence of harmonics and double pulses (Fig. 6) are maintained during an individual flight. These specific characteristics of the signal may be used to identify echoes belonging to its own emitted echolocation pulse.
    Type of Medium: Electronic Resource
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  • 6
    Electronic Resource
    Electronic Resource
    Springer
    Journal of comparative physiology 159 (1986), S. 403-413 
    ISSN: 1432-1351
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Medicine
    Notes: Summary 1. The echolocating ‘long CF/FM-bat’Rhinolophus rouxi and the ‘short CF/FM-bats’Hipposideros bicolor andHipposideros speoris were tested for catching responses to moving and non-moving targets. 2. Under our experimental conditions (freshly caught caged bats in a natural environment)Rhinolophus rouxi andHipposideros speoris only responded to insects of any sort that were beating their wings. The bats showed no reactions whatsoever to nonmoving insects or those walking on the floor or the sides of the cage. 3. Hipposideros bicolor responded in the same way as the above species to wingbeating insects but in addition also attacked walking insects. In 27 presentations 15 walking insects were caught (Fig. 2). 4. Rhinolophus rouxi, Hipposideros speoris andHipposideros bicolor also detected, approached and seized tethered cockroaches hanging from the ceiling when these were vibrating up and down (Fig. 3). This indicates that any oscillating movement and not specific aspects of wing beating were the key releasers for catching behaviour in all three species. However, a wing beating insect is strongly preferred over a vibrating one in all three species (Fig. 4). 5. Rhinolophus rouxi, Hipposideros speoris andHipposideros bicolor attacked and seized a dead bait when it was associated with a wing beating device (Fig. 1). All three species responded effectively to beat frequencies as low as 10 beats/s (peak-to-peak amplitude of the wing excursion 20 mm). For lower frequencies the response rates rapidly deteriorated (Fig. 5). 6. Horseshoe bats no longer responded to wing beats of 5 beats/s when the wing beat amplitude was 2 to 1 mm or to wing beats of 2 to 1 beats/s when the amplitude was 3 mm or lower (Fig. 6). This suggests that the speed of the wing is a critical parameter. From these data we infer that the threshold for the catching responses is at a wing speed of about 2 to 1 cm/s. 7. In horseshoe bats (experimental tests) and the two hipposiderid species (behavioural observations) one single wing beat was enough to elicit a catching response (Fig. 8). 8. It is concluded that ‘long’ and ‘short’ CF/ FM-bats feature a similar responsiveness to fluttering targets. The sensitivity to oscillating movements is considered as an effective detection mechanism for any sort of potential prey.
    Type of Medium: Electronic Resource
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  • 7
    Electronic Resource
    Electronic Resource
    Springer
    Journal of comparative physiology 106 (1976), S. 111-125 
    ISSN: 1432-1351
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Medicine
    Notes: Summary For echolocation,Rhinolophus ferrumequinum emits orientation sounds, each of which consists of a long constant-frequency (CF) component and short frequency-modulated (FM) components. The CF component is about 83 kHz and is used for Doppler-shift compensation. In this bat, single auditory nerve fibers and cochlear nuclear neurons tuned at about 83 kHz show low threshold and very sharp filter characteristics. The slopes of their tuning curves ranged between 1,000 and 3,500 dB/octave and their Q-10 dB values were between 20 and 400, 140 on the average (Figs. 3–5). The peripheral auditory system is apparently specialized for the reception and fine frequency analysis of the CF component in orientation sounds and Doppler-shift compensated echoes. This specialization is not due to suppression or inhibition comparable to lateral inhibition, but due to the mechanical specialization of the cochlea. Peripheral auditory neurons with the best frequency between 77 and 87 kHz showed not only on-responses, but also off-responses to tonal stimuli (Figs. 1, 2, and 6). The off-responses with a latency comparable to that of N1-off were not due to a rebound from either suppression or inhibition, but probably due to a mechanical transient occurring in the cochlea at the cessation of a tone burst.
    Type of Medium: Electronic Resource
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  • 8
    Electronic Resource
    Electronic Resource
    Springer
    Journal of comparative physiology 160 (1987), S. 509-515 
    ISSN: 1432-1351
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Medicine
    Notes: Summary 1. The response of the echolocating bat,Megaderma lyra, was tested to different kinds of prey in an outdoor cage. The bats caught larger flying insects (moths, beetles, grasshoppers, and cockroaches) on the wing and also picked up arthropods (solifugid spiders, beetles and cockroaches) and small vertebrates (mice, fishes, frogs and geckoes) from the ground. After touching the prey with the muzzle, the bats were able to differentiate between species. Scorpions and toads were not taken byM. lyra. 2. In lighted and in dark conditions,M. lyra detected and caught prey only when it moved. Dead frogs briskly pulled over the floor were also detected and caught, whereas stationary dead frogs were disregarded by the bats (Table 1). 3. When dead frogs were pulled over the watered surface of a glass plate to eliminate noises by motion, the motion no longer alarmed the bats. From the results of these experiments it was concluded thatM. lyra detects prey on the ground by listening to the noise of the moving target only, and not by echolocation (Table 1 C, Fig. 1). Furthermore,M. lyra were not attracted by frog calls. 4. M. lyra differentiated between palatable frogs and non-palatable toads only after touching the prey with the muzzle. 5. Experiments with freshly killed frogs coated with toad secretions or covered with toad skins indicate thatM. lyra differentiates between frogs and toads by chemical means. There was no evidence that these prey were differentiated by means of echolocation.
    Type of Medium: Electronic Resource
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  • 9
    Electronic Resource
    Electronic Resource
    Springer
    Naturwissenschaften 58 (1971), S. 627-627 
    ISSN: 1432-1904
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Chemistry and Pharmacology , Natural Sciences in General
    Type of Medium: Electronic Resource
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  • 10
    Electronic Resource
    Electronic Resource
    Springer
    Journal of comparative physiology 154 (1984), S. 133-142 
    ISSN: 1432-1351
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Medicine
    Notes: Summary 1. Audiograms are recorded from one non-echolocating and nine echolocating sympatrically living bat species of South India. These species areCynopterus sphinx (non-echolocating),Tadarida aegyptiaca, Taphozous melanopogon, T. kachhensis, Rhinopoma hardwickei, Pipistrellus dormeri, P. mimus, Hipposideros speoris, H. bicolor andMegaderma lyra. 2. InRhinopoma hardwickei a highly sensitive frequency range was found which is narrowly tuned to the frequency band of the bat's CF-echolocation call (32–35 kHz, Fig. 3). In hipposiderids a ‘filter’ narrowly tuned to the frequency of the CF-part of the CF-FM echolocation sounds (137.5 kHz inH. speoris and 151.5 kHz inH. bicolor, Fig. 5) could be recorded from deeper parts of IC. 3. In the echolocating species the best frequency of the audiograms closely matched with that frequency range in the echolocation calls containing most energy. 4. In bat species foraging flying prey best frequencies of audiograms and height of preferred foraging areas are inversely related, i.e. bat species hunting high above canopy have lower best frequencies than those foraging close to or within canopy (Fig. 6). 5. A hypothesis is forwarded explaining how fluttering target detection by constant frequency echolocation might have evolved from long distance echolocation by pure tone signals.
    Type of Medium: Electronic Resource
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