Vocalization Influences Auditory Processing in Collicular Neurons of the CF-FM-Bat, Rhinolophus ferrumequinum

1. In awake Greater Horseshoe bats (Rhinolophus ferrumequinum) the responses of 64 inferior colliculus neurons to electrically elicited vocalizations (VOC) and combinations of these with simulated echoes (AS: pure tones and AS(FM): sinusoidally frequency-modulated tones mimicking echoes from wing beating insects) were recorded. 2. The neurons responding to the species-specific echolocation sound elicited by electrical stimulation of the central grey matter had best frequencies between 76 and 86 kHz. The response patterns to the invariable echolocation sound varied from unit to unit (Fig. 1). 3. In 26 neurons the responses to vocalized echolocation sounds markedly differed from those to identical artificial ones copying the CF-portion of the vocalized sound (AS). These neurons reacted with a different response to the same pure tone whether it was presented artificially or vocalized by the bat (Fig. 2). In these neurons vocalization activities qualitatively alter the responsiveness to the stimulus parameters of the echoes. 4. A few neurons neither responded to vocalization nor to an identical pure tone but discharged when vocalization and pure tone were presented simultaneously. 5. In 2 neurons synchronized encoding of small frequency-modulations of the pure tone (mimicking an echo returning from a wing beating prey) occurred only during vocalization. Without vocalization the neurons did not respond to the identical stimulus set (Fig. 3). In these neurons vocalization activities enhanced FM-encoding capabilities otherwise not present in these neurons. 6. FM-encoding depended on the timing between vocalization and frequency-modulated signal (echo). As soon as vocalization and FM-signal no more overlapped or at least 60–80 ms after onset of vocalization synchronized firing to the FM was lost (4 neurons) (Fig. 4). 7. 4 neurons weakly responded to playbacks of the bat's own vocalization 1 ms after onset of vocalization. But when the playback frequency was shifted to higher frequencies by more than 400 Hz the neurons changed firing patterns and the latency of the first response peak (Fig. 5). These neurons sensitive to frequency shifts in the echoes returning during vocalization may be relevant to the Doppler-shift compensation mechanism in Greater Horseshoe bats

Coding of Small Sinusoidal Frequency and Amplitude Modulations in the Inferior Colliculus of 'CF-FM' Bat, Rhinolophus Ferrumequinum

Single neurons in the inferior colliculus of the Greater Horseshoe bat, Rhinolophus ferrumequinum, showed two broad categories of response patterns to sinusoidally frequency (SFM) or amplitude (SAM) modulated stimuli. Tonic responding cells (best excitatory frequency (BEF) between 10 and 90 kHz) showed a rough sinusoidal modulation of the discharge pattern to SFM. Transient responding neurons, generally showing on- or off-responses to pure tones, (BEF between 65 and 88 kHz), displayed highly synchronized discharge patterns to SFM-cycles (Fig. 1). Modulation rates between 20 and 100 Hz were most effective and some neurons encoded modulation rates up to 350 Hz (Figs. 2 and 3). The SFM responses were best synchronized to the modulation envelope for center frequencies in the upper portion of the tuning curve (Figs. 4 and 5). Sharply tuned neurons with BEF around 80 kHz had the lowest threshold for modulation depth (± 10 Hz or 0.025%) (Fig. 6). In general, SAMs evoked the same type of response patterns and were encoded down to modulation index of 3% (Fig. 7). The fine frequency and amplitude discriminations for periodical modulations by collicular neurons is discussed as related to the detection and discrimination performance of bats, when preying on flying insects in clustered surroundings

The functional role of GABA and glycine in monaural and binaural processing in the inferior colliculus of horseshoe bats

The functional role of GABA and glycine in monaural and binaural signal analysis was studied in single unit recordings from the central nucleus of the inferior colliculus (IC) of horseshoe bats (Rhinolophus rouxi) employing microiontophoresis of the putative neurotransmitters and their antagonists bicuculline and strychnine. Most neurons were inhibited by GABA (98%; N= 107) and glycine (92%; N = 118). Both neurotransmitters appear involved in several functional contexts, but to different degrees. Bicuculline-induced increases of discharge activity (99% of cells; N= 191) were accompanied by changes of temporal response patterns in 35 % of neurons distributed throughout the IC. Strychnine enhanced activity in only 53% of neurons (N= 147); cells exhibiting response pattern changes were rare (9%) and confined to greater recording depths. In individual cells, the effects of both antagonists could markedly differ, suggesting a differential supply by GABAergic and glycinergic networks. Bicuculline changed the shape of the excitatory tuning curve by antagonizing lateral inhibition at neighboring frequencies and/or inhibition at high stimulation levels. Such effects were rarely observed with strychnine. Binaural response properties of single units were influenced either by antagonization of inhibition mediated by ipsilateral stimulation (bicuculline) or by changing the strength of the main excitatory input (bicuculline and strychnine)

The auditory cortex of the bat Phyllostomus discolor: Localization and organization of basic response properties

<p>Abstract</p> <p>Background</p> <p>The mammalian auditory cortex can be subdivided into various fields characterized by neurophysiological and neuroarchitectural properties and by connections with different nuclei of the thalamus. Besides the primary auditory cortex, echolocating bats have cortical fields for the processing of temporal and spectral features of the echolocation pulses. This paper reports on location, neuroarchitecture and basic functional organization of the auditory cortex of the microchiropteran bat <it>Phyllostomus discolor </it>(family: Phyllostomidae).</p> <p>Results</p> <p>The auditory cortical area of <it>P. discolor </it>is located at parieto-temporal portions of the neocortex. It covers a rostro-caudal range of about 4800 μm and a medio-lateral distance of about 7000 μm on the flattened cortical surface.</p> <p>The auditory cortices of ten adult <it>P. discolor </it>were electrophysiologically mapped in detail. Responses of 849 units (single neurons and neuronal clusters up to three neurons) to pure tone stimulation were recorded extracellularly. Cortical units were characterized and classified depending on their response properties such as best frequency, auditory threshold, first spike latency, response duration, width and shape of the frequency response area and binaural interactions.</p> <p>Based on neurophysiological and neuroanatomical criteria, the auditory cortex of <it>P. discolor </it>could be subdivided into anterior and posterior ventral fields and anterior and posterior dorsal fields. The representation of response properties within the different auditory cortical fields was analyzed in detail. The two ventral fields were distinguished by their tonotopic organization with opposing frequency gradients. The dorsal cortical fields were not tonotopically organized but contained neurons that were responsive to high frequencies only.</p> <p>Conclusion</p> <p>The auditory cortex of <it>P. discolor </it>resembles the auditory cortex of other phyllostomid bats in size and basic functional organization. The tonotopically organized posterior ventral field might represent the primary auditory cortex and the tonotopically organized anterior ventral field seems to be similar to the anterior auditory field of other mammals. As most energy of the echolocation pulse of <it>P. discolor </it>is contained in the high-frequency range, the non-tonotopically organized high-frequency dorsal region seems to be particularly important for echolocation.</p

Bat azimuthal echolocation using interaural level differences: modeling and implementation by a VLSI-based hardware system

Bats have long fascinated both scientists and engineers due to their superb ability to use echolocation to fly with speed and agility through complex natural environments in complete darkness. This dissertation presents a neuromorphic VLSI circuit model of bat azimuthal echolocation. Interaural level differences (ILDs) are the cues for bat azimuthal echolocation and are also the primary cues used by other mammals to localize high frequency sounds. The fact that neurons in bats respond to short echoes by one or two spikes strongly suggests that the conventionally used firing rate is an unlikely code. The operation of first spike latency in ILD computation and transformation is investigated in a network of spiking neurons linking the lateral superior olive (LSO), dorsal nucleus of the lateral lemniscus (DNLL), and inferior colliculus (IC). The results of the investigation suggest that spatially distributed first spike latencies can serve as a fast code for azimuth that can be ``read-out'' by ascending stages. With the hardware echolocation model that uses spike timing representation, we study how multiple echoes can affect bat echolocation and demonstrate that the response to multiple sounds is not a simple linear addition of the response to single sounds. By developing functional models of the bat echolocation system, we can study the efficient implementation demonstrated by nature. For example, variations among analog VLSI circuit units due to the unavoidable transistor mismatch - traditionally thought of as a hurdle to overcome - have been found beneficial in generating the desired diversity of response that is similar to their neural counterparts. This work advocates the use and design of summating and exponentially decaying synapses. A compact and easily controllable synapse circuit has found an application in achieving a linear temporal spike summation by operating with a very short time constants. It has also been applied in modeling a nonlinear intensity-latency trading by working with a long synaptic time constant. We propose a new synapse circuit model that is compatible with those used in computational models and implementable by CMOS transistors operating in the subthreshold region

Biologically inspired processing of radar and sonar target echoes

Modern radar and sonar systems rely on active sensing to accomplish a variety of tasks, including detection and classification of targets, accurate localization and tracking, autonomous navigation and collision avoidance. Bats have relied on active sensing for over 50 million years and their echolocation system provides remarkable perceptual and navigational performance that are of envy to synthetic systems. The aim of this study is to investigate the mechanisms bats use to process echo acoustic signals and investigate if there are lessons that can be learned and ultimately applied to radar systems. The basic principles of the bat auditory system processing are studied and applied to radio frequencies. A baseband derivative of the Spectrogram Correlation and Transformation (SCAT) model of the bat auditory system, called Baseband SCAT (BSCT), has been developed. The BSCT receiver is designed for processing radio-frequency signals and to allow an analytical treatment of the expected performance. Simulations and experiments have been carried out to confirm that the outputs of interest of both models are “equivalent”. The response of the BSCT to two closely spaced targets is studied and it is shown that the problem of measuring the relative distance between two targets is converted to a problem of measuring the range to a single target. Nearly double improvement in the resolution between two close scatterers is achieved with respect to the matched filter. The robustness of the algorithm has been demonstrated through laboratory measurements using ultrasound and radio frequencies (RF). Pairs of spheres, flat plates and vertical rods were used as targets to represent two main reflectors