Differential phylogenetic development of the acoustic nuclei among chiroptera.
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The calls of five syntopic species of Neoconocephalus varied significantly in their spectral composition. The center-frequency of the narrow-band low-frequency component varied from 7 kHz to 15 kHz among the five species. Hearing thresholds, as determined from whole nerve recordings, did not vary accordingly among the five species but were lowest in the range from 16 kHz to 18 kHz in all five species. Iso-intensity response functions were flat for stimulus intensities up to 27 dB above threshold, indicating an even distribution of the best frequencies of individual receptor cells. At higher stimulus intensities, the intensity/response functions were steeper at frequencies above 35 kHz than at lower frequencies. This suggests the presence of a second receptor cell population for such high frequencies, with 25-30 dB higher thresholds. This receptor cell population is interpreted as an adaptation for bat avoidance. The transmission properties of the Neoconocephalus habitat (grassland) had low-pass characteristics for pure tones. Frequencies below 10 kHz passed almost unaffected, while attenuation in excess of spherical attenuation increased at higher frequencies. Considering these transmission properties and the tuning of female hearing sensitivity, call frequencies of approximately 9-10 kHz should be most effective as communication signals in this group of insects. It is discussed that the frequency of male calls is strongly influenced by bat predation and by the transmission properties of the habitat but is not strongly influenced by the tuning of the female hearing system.
For auditory imaging, a bat emits orientation sounds (pulses) and listens to echoes. The parameters characterizing a pulse-echo pair each convey particular types of biosonar information. For example, a Doppler shift (a difference in frequency between an emitted pulse and its echo) carries velocity information. For a 61-kHz sound, a 1.0-kHz Doppler shift corresponds to 2.8 ms-1 velocity. The delay of the echo from the pulse conveys distance (range) information. A 1.0-ms echo delay corresponds to a target distance of 17 cm. The auditory system of the mustached bat, Pteronotus parnelli, from Central America solves the computational problems in analyzing these parameters by creating maps in the cerebral cortex. The pulse of the mustached bat is complex. It consists of four harmonics, each of which contains a long constant-frequency (CF) component and a short frequency-modulated (FM) component. Therefore, there are eight components in the emitted pulse (CF1-4 and FM1-4). The CF signal is particularly suited for target velocity measurement, whereas the FM signal is suited for target distance measurement. Since the eight components differ from each other in frequency, they are analyzed in parallel at different regions of the basilar membrane in the inner ear. Then, they are separately coded by primary auditory neurons and are sent up to the auditory cortex through several auditory nuclei. During the ascent of the signals through these auditory nuclei, neurons responding to the FM components process range information, while other neurons responding to the CF components process velocity information. A comparison of the data obtained from the mustached bat with those obtained from other species illustrates both the specialized neural mechanisms specific to the bat's auditory system, and the general neural mechanisms which are probably shared with many different types of animals.
Quantitative data for Doppler-shift compensation by Pteronotus parnellii parnellii were obtained with a device which propelled the bats at constant velocities over a distance of 12 m. The bats compensated for Doppler shifts at all velocities tested (0.1-5.0 ms-1). The main findings were (1) that compensation was usually accomplished by a progressive lowering of the approximately 61 kHz second harmonic constant-frequency component of emitted sounds in small frequency steps (93 +/- 72 Hz); (2) that the time needed to reach a steady compensation level averaged 514 +/- 230 ms and the number of pulses required to reach full compensation averaged 10.78 +/- 5.16; (3) that the animals compensated to hold the echo (reference) frequency at a value that was slightly higher than the resting frequency and slightly lower than the cochlear resonance frequency; (4) that reference frequency varied as a function of velocity, the higher the velocity of the animal, the higher was the reference frequency (slope 55 Hz m-1s-2); and (5) that the mean reference frequency was always an undercompensation. The average amount of undercompensation was 15.8%. There was a significant difference (P < or = 0.005) in Doppler-shift compensation data collected at velocities that differed by 0.1 ms-1. A velocity difference of 0.1 ms-1 corresponds to a Doppler-shift difference of about 35 Hz in the approximately 61 kHz signals reaching the ear.
Atmospheric conditions conducive to long-range transmission of low-frequency sound as used by elephants are found to exist in the Etosha National Park in Namibia during the late dry season. Meteorological measurements show that strong temperature inversions form at the surface before sunset and decay with sunrise, often accompanied by calm wind conditions during the early evening. These observations are used in an acoustic model to determine the sensitivity of infrasound to the effects of (a) the strength, thickness and elevation of temperature inversions, and (b) the growth and decay of an inversion typical of dry, elevated African savannas. The results suggest that the range over which elephants communicate more than doubles at night. Optimum conditions occur 1-2 h after sunset on clear, relatively cold, calm nights. At these times, ranges of over 10 km are likely, with the greatest amplification occurring at the lowest frequency tested. This strong diurnal cycle in communication range may be reflected in longer-lasting changes in weather and may exert a significant influence on elephant behaviour on time scales from days to many years.
We studied the acoustic behaviour of bats (Eptesicus nilssonii) hunting for large (wing span 5 cm) non-hearing hepialid moths (Hepialus humuli). Groups of silvery-white male H. humuli perform a short (30 min) hovering display flight over grassland at dusk. They typically hover at, or below, the tops of grass panicles and are therefore situated in a highly acoustically cluttered habitat. Occasionally, they move to a new position by making short (1-5 s) flights at higher levels. E. nilssonii is not a clutter specialist, and yet we found that they attacked H. humuli within the 'clutter overlap zone'. The bats did not change their signal design in any marked manner for this specific task. Measurements of echoes from a moth 10 cm above or below the grass tops showed that information for detecting the moths was available to the bats. Nevertheless, the bats did not attack moths in stationary hovering display flight, only when they moved above the grass panicles. The duration of the up/down flights (movements) were almost always longer than an entire capture sequence by the bats. Apparently, the bats rely on the movement of the moth in space, monitored over successive echoes, to discriminate moth echoes from overlapping clutter echoes.
Using an implanted electrode, we recorded the responses from the ultrasound-sensitive mantis interneuron 501-T3 during flying bat attacks in a large flight room where the mantis served as the target. 501-T3 responds to each vocalization emitted with multi-spike bursts when pulse repetition rates (PRRs) are below 55 pulses x s(-1). As PRR increases and pulse durations fall below 3 ms, 501-T3 ceases burst activity. On average, spike bursts cease 272 ms before contact (when the bat is 73 cm away from the preparation). The timing of cessation of activity in 501-T3 is similar to the latency for the diving portion of the response of the mantid (242 ms). Experiments using vocalizing stationary bats confirm that 501-T3 responds more reliably to longer pulse durations (> or =3 ms) when intensities are below 90 dB pe SPL. The cessation of 501-T3 activity is probably due both to the increasing PRR and to the decreasing pulse duration that occur in the terminal buzz phase of a bat attack. 501-T3 may be actively shut off at high PRRs and/or intensities to protect the interneuron from habituation while the mantis performs an escape response. The cessation of 501-T3 activity is consistent with the lack of a very late ultrasound-mediated evasive response by the mantis. However, cessation of 501-T3 activity may allow a true 'last-chance' response to be mediated by other neural systems.
Pteronotus parnellii uses the second harmonic (61-62 kHz) of the CF component in its orientation sounds for Doppler-shift compensation. The bat's inner ear is mechanically specialized for fine analysis of sounds at about 61-62 kHz. Because of this specialization, cochlear microphonics (CM) evoked by 61-62 kHz tone bursts exhibit prominent transients, slow increase and decrease in amplitude at the onset and cessation of these stimuli. CM-responses to 60-61 kHz tone bursts show a prominent input-output non-linearity and transients. Accordingly, a summated response of primary auditory neurones (N1) appears not only at the onset of the stimuli, but also at the cessation. N1-off is sharply tuned at 60-61 kHz, while N1-on is tuned at 63-64 kHz, which is 2 kHz higher than the best frequency of the auditory system because of the envelope-distortion originating from sharp mechanical tuning. Single peripheral neurones sensitive to 61-62 kHz sounds have an unusually sharp tuning curve and show phase-locked responses to beats of up to 3 kHz. Information about the frequencies of Doppler-shifted echoes is thus coded by a set of sharply tuned neurones and also discharges phase-locked to beats. Neurones with a best frequency between 55 and 64 kHz show not only tonic on-responses but also off-responses which are apparently related to the mechanical off-transient occuring in the inner ear and not to a rebound from neural inhibition.
The marine catfish (Arius felis) uses directional hearing for the acoustical detection of obstacles, and is primarily dependent on the vector components of near-field acoustics. Directional sound, detected as lateral line action potentials, produces the strongest response on the side toward the stimulus. Responses are maximal in the 50-150 Hz range. The swim bladder is asymmetrical as a sound generator (7 dB front-to-back ratio). As a sound receiver, the swim bladder is even more directional (24 dB front-to-back ratio at 100 Hz). It is concluded that directional response to sound in fishes can involve the interaction of at least two mechanisms, in this case the lateral line and the swim bladder.
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Mammalian vocalizations require the precise coordination of separate laryngeal and respiratory motor pathways. Precisely how and where in the brain vocal motor patterns interact with respiratory rhythm control is unknown. The parabrachial nucleus (PB) is known to mediate key respiratory reflexes and is also considered a principle component of the mammalian vocal motor pathway, making it a likely site for vocal-respiratory interactions, yet a specific role for the PB in vocalizing has yet to be demonstrated. To investigate the role of the PB in vocal-respiratory coordination, we pharmacologically manipulated synaptic activity in the PB while spontaneously vocalizing horseshoe bats were provoked to emit either short, single syllable or long, multisyllabic vocal motor patterns. Iontophoresis of the GABAA agonist muscimol (MUS) into the lateral PB extended expiratory durations surrounding all vocalizations and increased mean call durations. Alternatively, application of the GABAA antagonist bicuculline methiodide (BIC) shortened expirations and call durations. In addition, BIC eliminated the occurrence of multisyllabic vocalizations. BIC caused a mild increase in quiet breathing rates, whereas MUS tended to slow quiet breathing. The results indicate that GABAA receptor-mediated inhibition in the lateral PB modulates the time course of respiratory phase switching during vocalizing, and is needed for proper coordination of calling and breathing in mammals. We hypothesize that vocal-respiratory rhythm entrainment is achieved at least in part via mechanisms similar to other forms of locomotor-respiratory coupling, namely somatosensory feedback influences on respiratory phase-switching in the lateral PB.
Vocal amplitude, one of the crucial factors for the exchange of acoustic signals, has been neglected in studies of animal communication, but recent studies on song variation in Common Nightingales Luscinia megarhynchos have revealed new insights into its importance in the singing behavior of territorial birds. In nightingales song amplitude is not maximized per se, but is individually regulated according to the level of masking background noise. Also, birds adjust their vocal intensity according to social variables, as in male-male interactions. Moreover, during such interactions, males exploited the directionality of their songs to broadcast them in the direction of the intended receivers ensuring the most effective signal transmission. Studies of the development of this typical long-range signaling suggest that sound level is highly interrelated with overall developmental progression and learning, and thus should be viewed as an integral part of song ontogeny. I conclude that song amplitude is a dynamic feature of the avian signal system, which is individually regulated according to the ecological demands of signal transmission and the social context of communication.
We recorded vocalizations of megachiropteran bats, Cynopterus brachyotis, C. horsfieldi, Megaerops ecaudatus and Macroglossus sobrinus as they were held in our hands, in a hanging position or moved up and down, simulating flight. All four species produced regular audible tone pulses with a frequency spectrum of less than 9 kHz (peak at 4-6 kHz). The duration of a single pulse varied from 14.9 to 23.6 msec on average among these species. The interpulse interval was between 108.3 and 127.7 msec. In M. ecaudatus and M. sobrinus, double pulses were occasionally emitted, with interpulse intervals averaging 39 msec and 31 msec, respectively. The regular pulse emission may indicate that the bats studied use sounds to probe space.
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One of the fundamental organizational principles of the mammalian sensory cortex is the topographic representation of the sensory epithelium. A cochleotopic topographic organization has been demonstrated for a number of auditory cortical fields, including the primary auditory field (AI). In the spatial cortical dimension orthogonal to the cochleotopic frequency gradient, systematic arrangements of other functional parameters have been observed. The first evidence of a spatial segregation of functional parameters along the isofrequency domain of cat AI was the discovery of binaural interaction bands. Recent studies show topographic representations of additional functional parameters in the central auditory system of the cat. Among the functional parameters that are evidence of systematic and topographic representations in cat auditory cortical fields are spectral resolution (sharpness of tuning), temporal resolution (preferred repetition rate), and inhibitory properties related to the coding of broadband signals.
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In a series of publications, the author used the correlation hypothesis to explain the main mechanisms of functioning of bat and dolphin echo locators. A good fit of calculated parameters to experimental data was shown. In this work, by the example of generalization of the recognition problem, the use by the brain of correlation methods for solving more general problems independently on the modality of sensor signals was analyzed. In favor of the hypothesis that correlation methods are widely used in brain functioning are the data presented in the paper, which prove that an associative neuron is a suitable analog computer. It is suited for rapid "computation" of the intercorrelation function of discrete input and reference signals. The set of weighting coefficients of neuron synaptic inputs serve as such signals. The pool of associative neurons determines the values on the correlation function in the required range by changing the "numbers" of inputs from neuron to neuron at which discrete signals arrive.