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Geographic profiling and animal foraging.

Geographic profiling was originally developed as a statistical tool for use in criminal cases, particularly those involving serial killers and rapists. It is designed to help police forces prioritize lists of suspects by using the location of crime scenes to identify the areas in which the criminal is most likely to live. Two important concepts are the buffer zone (criminals are less likely to commit crimes in the immediate vicinity of their home) and distance decay (criminals commit fewer crimes as the distance from their home increases). In this study, we show how the techniques of geographic profiling may be applied to animal data, using as an example foraging patterns in two sympatric colonies of pipistrelle bats, Pipistrellus pipistrellus and P. pygmaeus, in the northeast of Scotland. We show that if model variables are fitted to known roost locations, these variables may be used as numerical descriptors of foraging patterns. We go on to show that these variables can be used to differentiate patterns of foraging in these two species.

Animals↗

Acoustic characteristics of air puff-induced 22-kHz alarm calls in direct recordings.

Alarm calls were induced in adult Wistar rats by an air puff. Emitted calls were digitized and directly recorded on a computer hard drive. The long-duration 22-kHz calls were emitted almost exclusively in series. Initial calls in the series tended to have the longest durations, higher frequency range, and the highest degree of frequency modulation, as compared to other calls. The frequency modulation always appeared as a downward sweep and seemed to represent a tuning of individual calls to a 3 kHz communicatory band. Regardless of the maximum frequency, rats always reached approximately the same minimum frequency, common to all calls. Thus, the broader was the frequency range of a given call, the longer the call duration. It is postulated, therefore, that rats emit 22-kHz calls at the minimum possible ultrasonic frequency they are able to produce, which is synonymous with peak frequency. It is further postulated that production of alarm calls in series, with long call duration and the invariably low ultrasonic frequency, maximizes successful communication in dangerous situations. Exceptions to this rule were observed immediately following air puffs, suggesting that acoustic parameters of the initial calls may differ from the alarming properties of the remaining 22-kHz calls.

Acoustics↗

GABAergic inhibition contributes to pulse repetition rate-dependent frequency selectivity in the inferior colliculus of the big brown bat, Eptesicus fuscus.

This study examined the effect of bicuculline application on sharpness of frequency tuning curves (FTCs) of bat inferior collicular neurons plotted under three different pulse repetition rates (PRRs) of 10, 30 and 90 pulses per second. The sharpness of FTCs of collicular neurons, which was expressed in Q(n) (Q(10), Q(20), Q(30)) and bandwidths (90, 75 and 50% of the maximal response at the best frequency), improved with increasing PRR. However, this PRR-dependent frequency selectivity of collicular neurons was abolished during bicuculline application. This observation suggests that GABAergic inhibition contributes more effectively to sharpening of FTCs at higher than at lower PRRs.

Action Potentials↗

Multiple combination-sensitive neurons in the auditory cortex of the mustached bat.

The mustached bat, Pteronotus parnellii, emits biosonar pulses consisting of four constant-frequency (CF(1-4)) and four frequency-modulated (FM(1-4)) components. The FM-FM area of its auditory cortex consists of three subdivisions, containing either FM(1)-FM(2), FM(1)-FM(3) or FM(1)-FM(4) combination-sensitive neurons. The FM-FM area also contains 'multiple combination-sensitive' neurons: FM(1)-FM(2,3), FM(1)-FM(3,4), FM(1)-FM(2,4), and FM(1)-FM(2,3,4) neurons. All FM-FM neurons are tuned to a time delay (echo delay) of FM(n) (n=2-4) from FM(1). In the present study, we made the following four major findings. (1) Multiple combination-sensitive neurons show the strongest response to a combination of more than two signal elements. (2) Multiple combination-sensitive neurons are located in about 100 microm wide bands at the boundaries between two adjacent subdivisions of the FM-FM area. (3) Iso-best-delay contour lines across the three single combination-sensitive subdivisions are not interrupted by multiple combination-sensitive bands. (4) Each subdivision of the FM-FM area has frequency-vs.-frequency coordinates in terms of best FM(1) and best FM(n) frequencies for facilitation, although such coordinates were not obtained with single tone bursts.

Acoustic Stimulation↗

Paradoxical lateral suppression in the dolphin's auditory system: weak sounds suppress response to strong sounds.

A paradoxical phenomenon was found in the auditory system of dolphins: weak sounds suppressed the brain responses to much stronger sounds. This occurred when the brain evoked potentials to rhythmic sound amplitude modulations were recorded. The response was markedly suppressed by addition of another sound of higher frequency and down to 40 dB lower intensity than the amplitude-modulated signal. Only the sustained rhythmic response was suppressed while transient on-response was not, thus indicating that the suppression influenced the ability of evoked potentials to follow rapid amplitude modulations. This prevents weak sounds from being masked by stronger ones. It may help a dolphin to perceive weaker echo-signals in the background of stronger emitted pulses.

Acoustic Stimulation↗

Topographical distribution of delay-tuned responses in the mustached bat inferior colliculus.

In the mustached bat, delay-tuned neurons respond best to specific delays between the first harmonic frequency modulated (FM) component (FM1; 24-29 kHz) of the emitted biosonar pulse and a higher harmonic FM component in returning echoes (e.g. FM3, 72-89 kHz). These delay-tuned, combinatorial responses predominate in the inferior colliculus (IC) of the mustached bat. This study examined the topographical distribution of delay-tuned neurons in the 72-89 kHz frequency representation of the IC. We recorded and histologically localized 163 single units. Ninety units were facilitated and 41 were inhibited by the combination of two frequencies in the 24-29 kHz and 72-89 kHz ranges. The facilitatory responses were selective for delays up to 20 ms between the two signals. To determine if delay-tuned neurons were topographically organized, we plotted the dorsomedio-ventrolateral and caudo-rostral positions of each unit versus its best delay. Best delay was not correlated with either location. Response latency to best frequency tones was topographically organized, but was not correlated with best delay. This indicates that the latency axis in the IC is unrelated to the delay tuning of these combinatorial neurons. Because delay-tuned neurons are not topographically organized in the IC but are in the auditory cortex, our findings suggest that the creation and organization of delay-tuned neurons occur at different stages in the ascending auditory system.

Acoustic Stimulation↗

Excitatory and facilitatory frequency response areas in the inferior colliculus of the mustached bat.

In the mustached bat's central nucleus of the inferior colliculus (ICC), many neurons display facilitatory or inhibitory responses when presented with two tones of distinctly different frequencies. Our previous studies have focused on spectral interactions between specific frequency bands contained in the bat's sonar vocalization. In this study, we describe excitatory and facilitatory frequency response areas across all frequencies in the mustached bat's audible range. We show that many neurons in the ICC have more extensive frequency interactions than previously documented. We recorded responses of 96 single units to single tones and combinations of two tones. Best frequencies of the units ranged from 59-15 kHz. Forty-one units had a single, excitatory frequency response area. The rest of the units had more complex frequency tuning that included multiple excitatory frequency response areas and facilitatory frequency response areas. Some of the facilitatory frequency interactions were between one sound with energy in a sonar frequency band and a second sound with energy in a non-sonar frequency band. We also found that neurons could be facilitated by more than one additional frequency band. Our findings of extensive frequency interactions in the ICC of the mustached bat suggest that some neurons may be well suited for the analysis of complex sounds, possibly including social communication sounds.

Acoustic Stimulation↗

NMDA-mediated facilitation in the echo-delay tuned areas of the auditory cortex of the mustached bat.

We recorded the responses of single delay-tuned neurons in the dorsal fringe (DF) area and the FM-FM area of the auditory cortex of the mustached bat using multi-barreled carbon-fiber electrodes. An iontophoretic application of N-methyl-D-aspartate (NMDA) or kainate (KA) to a DF neuron evoked a burst of discharges from the neuron. The burst of discharges evoked by NMDA was always smaller than that evoked by KA. Simultaneous application of D-2-Amino-5-phosphonovalerate (APV) with NMDA and KA abolished the NMDA-evoked but not the KA-evoked discharges. APV did not evoke any significant changes in the auditory responses of 43 out of the 47 delay-tuned neurons studied in the DF area, and in all 20 neurons studied in the FM-FM area. In the remaining four DF neurons, however, APV either increased the initial discharges of their auditory response or decreased the late discharges of their response. These results indicate that in the majority of neurons in the DF and FM-FM areas NMDA receptors do not play a significant role in the processing of target-distance information, and that their facilitative auditory responses are basically created by synaptic interactions occurring in the subcortical auditory nuclei.

2-Amino-5-phosphonovalerate↗

Perception of two-tone complexes by the goldfish (Carassius auratus).

Previous experiments on the sense of hearing in goldfish have used a stimulus generalization paradigm to investigate the perceptual dimensions evoked by spectrally and temporally complex sounds. The present experiments investigated the effects on perception of the frequency separation between two tones. In the first set of experiments, six groups of goldfish were classically conditioned to a single tone and then tested for generalization to two-tone complexes having one frequency component equal to the conditioning tone, and the other differing by 2-256 Hz. Generalization declined with increasing frequency differences up to about 32 Hz, and then increased for wider frequency separations. These functions indicate that a restricted range of beat rates produces a perceptual quality that is quite unlike that of a single tone. The generalization function of frequency separation resembles the inverse of the 'fluctuation strength' and 'roughness' functions for human listeners. The second experiment investigated the effects of spectral location on the perception of a 32 Hz beat rate. Goldfish were conditioned to a two-tone complex (500 and 532 Hz) and then tested for generalization to single tones at various frequencies between 200 and 1200 Hz, and to two-tone complexes having a 32 Hz beat rate but with the lower tone component at various frequencies. For single-tone stimuli, generalization was relatively weak but showed a peak at 500 Hz. For the two-tone stimuli, generalization was more robust, but showed a similarly shaped gradient centered on 500 Hz. Thus, goldfish behaved as if they had acquired information about both temporal modulation and the frequency location of the tone components. These perceptual behaviors appear to be shared with humans and other vertebrates.

Acoustic Stimulation↗

Processing of sinusoidally frequency modulated signals in the nuclei of the lateral lemniscus of the big brown bat, Eptesicus fuscus.

Neurons in the nuclei of the lateral lemniscus (NLL) of the big brown bat, Eptesicus fuscus, show several distinctive patterns of response to unmodulated tones. Previous work suggests that sustained responders are specialized to transmit information about sound level and duration while onset responders transmit precise timing information. The biosonar signals of E. fuscus consist of multiple, downward frequency modulated sweeps that change in slope and repetition rate as the bat approaches a target. An obvious hypothesis would be that NLL neurons with sustained responses should discharge during the time when the frequency of a signal is within their response area, but that onset responders should discharge each time the frequency enters the excitatory portion of their response area. In this study we examined the responses of NLL neurons to sinusoidally frequency modulated (SFM) signals presented monaurally to awake, restrained bats. Extracellular recordings were obtained from single neurons in the multipolar and columnar divisions of the ventral nucleus (VNLLm and VNLLc), the intermediate nucleus (INLL) and the dorsal nucleus of the lateral lemniscus (DNLL). All NLL neurons responded synchronously to SFM signals under some conditions. The temporal precision of synchronization was quantified using a coefficient of synchronization (CS), where a value of I equals perfect synchrony. Maximum CS values ranged from 0.70 to >0.99, were generally highest at low modulation rates ( <200 Hz), and showed lowpass characteristics for modulation rate. The maximal modulation rates that elicited synchronous discharge ranged from 50 to 500 Hz. The highest maximal rates were found in the VNLLm and VNLLc, the lowest in DNLL. The ability of NLL neurons to synchronize their discharge to the pattern of an SFM signal is intermediate between that of neurons in the cochlear nucleus and in the inferior colliculus. For the majority of neurons in VNLLm, INLL and DNLL, the precision of synchronization was approximately equal for the downward and upward components of the SFM signal; in contrast, 69% of VNLLc neurons responded selectively to the downward component of the SFM signal. All VNLLc neurons and a subset of those in VNLLm, INLL, and DNLL responded synchronously to SFM signals only if the frequency excursions included a border of the excitatory frequency bandwidth, suggesting that the synchronous discharge was due primarily to the repeated passage of the stimulus frequency into and out of the excitatory portion of the response area. In the case of VNLLc neurons, only the high frequency border was effective; Other neurons, especially those in DNLL, responded synchronously to SFM signals with frequency excursions that were confined entirely within the excitatory response area.

Acoustic Stimulation↗

Tonotopic representation and space map in the non-primary auditory cortex of the mustached bat.

As auditory system has no sensory epithelium into which auditory space are projected, we studied the physiological map of the auditory space in the non-primary auditory cortex of the mustached bat by using the echo of their orientation sound. Ten bats were used as experimental subjects. Tungsten wire electrodes were inserted obliquely in the dorsomedial (DM) and ventroposterior (VP) areas of the non-primary auditory cortex. When single neuron was isolated, best frequency (BF), best azymuth (BAZ) and best elevation (BEL) were measured and were plotted on a schematic figure. To mimic its biosonar, one loudspeaker, delivering synthesized orientation sounds, was placed in front of the animal, and another loudspeaker delivering synthesized echo was mounted on a movable hoop. Tonotopic representation was observed but complicated in both areas, and those areas could be divided into several subdivisions consisting of the neuron groups characterized by three frequency bands. The neurons were thought to be related to the processing of biosonar informations from the facts that their BFs agreed with the scope of the FM sweep of each echo harmonics. The magnitude of the response showed rapid increase at their BAZ or BEL, so that the neurons seemed to tune to a certain direction in the auditory space. Especially in the DM area, neurons assumed a systematic arrangement of their BAZs on the cerebral surface and showed some tendency of a systematic arrangement of their BELs. The DM area was thought to have a kind of neural map of the auditory space.

Acoustic Stimulation↗

Learning from dolphins.

Dolphins have been shown to have a powerful impact on the wellbeing of humans, how do they do it? This article reflects the thoughts of one person after spending time with these wonderful creatures.

Adult↗

Anatomical and functional imaging of the auditory cortex in awake mustached bats using magnetic resonance technology.

The auditory cortex of mustached bats, Pteronotus parnellii, has been studied extensively using neuroanatomical tract-tracing and electrophysiological techniques to elucidate the functional organization and neural mechanisms important for auditory processing. While these techniques have identified several cortical maps involved in processing auditory information, there has been no direct observation of the dynamics of simultaneous activation of several discrete areas. We applied magnetic resonance (MR) imaging techniques for visualizing brain structures in awake bats using a 7-Tesla magnet system; we also investigated functional MR imaging by measuring changes in stimulus-correlated blood oxygenation levels to detect cortical areas exhibiting evoked neural activity. High resolution (100 microm) anatomical images were successfully acquired without any motion artifacts. It was possible to reconstruct the whole brain image and analyze brain surface structures with three dimensional (3D) MR imaging data. These data provide detailed morphometric measurements that will allow localization of stimulus specific neural activity patterns using modified functional magnetic-resonance-imaging (fMRI) protocols. Motion artifacts is the primary disadvantage of using awake bats; our study shows that fMRI of a bat's brain is feasible and may prove to be an important advancement for a further understanding of auditory processing in this species.Themes: Sensory systems, Neural basis of behavior.

Acoustic Stimulation↗