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

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.     

Network Models of Frequency Modulated Sweep Detection

Frequency modulated (FM) sweeps are common in species-specific vocalizations, including human speech. Auditory neurons selective for the direction and rate of frequency change in FM sweeps are present across species, but the synaptic mechanisms underlying such selectivity are only beginning to be understood. Even less is known about mechanisms of experience-dependent changes in FM sweep selectivity. We present three network models of synaptic mechanisms of FM sweep direction and rate selectivity that explains experimental data: (1) The ‘facilitation’ model contains frequency selective cells operating as coincidence detectors, summing up multiple excitatory inputs with different time delays. (2) The ‘duration tuned’ model depends on interactions between delayed excitation and early inhibition. The strength of delayed excitation determines the preferred duration. Inhibitory rebound can reinforce the delayed excitation. (3) The ‘inhibitory sideband’ model uses frequency selective inputs to a network of excitatory and inhibitory cells. The strength and asymmetry of these connections results in neurons responsive to sweeps in a single direction of sufficient sweep rate. Variations of these properties, can explain the diversity of rate-dependent direction selectivity seen across species. We show that the inhibitory sideband model can be trained using spike timing dependent plasticity (STDP) to develop direction selectivity from a non-selective network. These models provide a means to compare the proposed synaptic and spectrotemporal mechanisms of FM sweep processing and can be utilized to explore cellular mechanisms underlying experience- or training-dependent changes in spectrotemporal processing across animal models. Given the analogy between FM sweeps and visual motion, these models can serve a broader function in studying stimulus movement across sensory epithelia.     

Recovery cycle of neurons in the inferior colliculus of the FM bat determined with varied pulse-echo duration and amplitude

The recovery cycle of auditory neurons is an important neuronal property which underlies a bat's ability in analyzing returning echoes and to determine target distance (i.e., echo ranging). In the same token, duration selectivity of auditory neurons plays an important role in pulse recognition in bat echolocation. Because insectivorous bats progressively vary the pulse parameters (repetition rate, duration, and amplitude) during hunting, the recovery cycle of auditory neurons is inevitably affected by their selectivity to other co-varying echo parameters. This study examines the effect of pulse duration and amplitude on recovery cycle of neurons in the central nucleus of the inferior colliculus (IC) of the FM bat, Pipistrellus abramus, using biologically relevant pulse-echo (P-E) pairs with varied duration and amplitude difference. We specifically examine how duration selectivity may affect a neuron's recovery cycle. IC neurons have wide range of recovery cycle and best duration (BD) covering P-E intervals and duration occurring different phases of hunting. The recovery cycle of most IC neurons increases with P-E duration and amplitude difference. Most duration-selective IC neurons recover rapidly when stimulated with biologically relevant P-E pairs. As such, neurons with short BD recover rapidly when stimulated with P-E pairs of short duration and small P-E amplitude difference. Conversely, neurons with long BD recover rapidly when stimulated with P-E pairs of long duration and large P-E amplitude difference. These data suggest that bats may potentially utilize the response of IC neurons with different BD and recovery cycle to effectively perform echo detection, recognition of echo duration and echo ranging throughout a target approaching sequence.     

The generation of direction selectivity in the auditory system

Both human speech and animal vocal signals contain frequency-modulated (FM) sounds. Although central auditory neurons that selectively respond to the direction of frequency modulation are known, the synaptic mechanisms underlying the generation of direction selectivity (DS) remain elusive. Here we show the emergence of DS neurons in the inferior colliculus by mapping the three major subcortical auditory nuclei. Cell-attached recordings reveal a highly reliable and precise firing of DS neurons to FM sweeps in a preferred direction. By using in vivo whole-cell current-clamp and voltage-clamp recordings, we found that the synaptic inputs to DS neurons are not direction selective, but temporally reversed excitatory and inhibitory synaptic inputs are evoked in response to opposing directions of FM sweeps. The construction of such temporal asymmetry, resulting DS, and its topography can be attributed to the spectral disparity of the excitatory and the inhibitory synaptic tonal receptive fields.     

Responses of midbrain auditory neurons in rats to FM and AM tones presented simultaneously

Speech and other communication signals contain components of frequency and amplitude modulations (FM, AM) that often occur together. Auditory midbrain (or inferior colliculus, IC) is an important center for coding time-varying features of sounds. It remains unclear how IC neurons respond when FM and AM stimuli are both presented. Here we studied IC neurons in the urethane-anesthetized rats when animals were simultaneously stimulated with FM and AM tones. Of 122 units that were sensitive to the dual stimuli, the responses could be grossly divided into two types: one that resembled the respective responses to FM or AM stimuli presented separately ("simple" sensitivity, 45% of units), and another that appeared markedly different from their respective responses to FM or AM tones ("complex" sensitivity, 55%). These types of combinational sensitivities were further correlated with individual cell's frequency tuning pattern (response area) and with their common response pattern to FM and AM sounds. Results suggested that such combinational sensitivity could reflect local synaptic interactions on IC neurons and that the neural mechanisms could underlie more developed sensitivities to acoustic combinations found at the auditory cortex.     

Duration tuning in the auditory midbrain of echolocating and non-echolocating vertebrates

Neurons tuned for stimulus duration were first discovered in the auditory midbrain of frogs. Duration-tuned neurons (DTNs) have since been reported from the central auditory system of both echolocating and non-echolocating mammals, and from the central visual system of cats. We hypothesize that the functional significance of auditory duration tuning likely varies between species with different evolutionary histories, sensory ecologies, and bioacoustic constraints. For example, in non-echolocating animals such as frogs and mice the temporal filtering properties of auditory DTNs may function to discriminate species-specific communication sounds. In echolocating bats duration tuning may also be used to create cells with highly selective responses for specific rates of frequency modulation and/or pulse-echo delays. The ability to echolocate appears to have selected for high temporal acuity in the duration tuning curves of inferior colliculus neurons in bats. Our understanding of the neural mechanisms underlying sound duration selectivity has improved substantially since DTNs were first discovered almost 50 years ago, but additional research is required for a comprehensive understanding of the functional role and the behavioral significance that duration tuning plays in sensory systems.     

The adaptive value of increasing pulse repetition rate during hunting by echolocating bats

During hunting, bats of suborder Microchiropetra emit intense ultrasonic pulses and analyze the weak returning echoes with their highly developed auditory system to extract the information about insects or obstacles. These bats progressively shorten the duration, lower the frequency, decrease the intensity and increase the repetition rate of emitted pulses as they search, approach, and finally intercept insects or negotiate obstacles. This dynamic variation in multiple parameters of emitted pulses predicts that analysis of an echo parameter by the bat would be inevitably affected by other co-varying echo parameters. The progressive increase in the pulse repetition rate throughout the entire course of hunting would presumably enable the bat to extract maximal information from the increasing number of echoes about the rapid changes in the target or obstacle position for successful hunting. However, the increase in pulse repetition rate may make it difficult to produce intense short pulse at high repetition rate at the end of long-held breath. The increase in pulse repetition rate may also make it difficult to produce high frequency pulse due to the inability of the bat laryngeal muscles to reach its full extent of each contraction and relaxation cycle at a high repetition rate. In addition, the increase in pulse repetition rate increases the minimum threshold (i.e. decrease auditory sensitivity) and the response latency of auditory neurons. In spite of these seemingly physiological disadvantages in pulse emission and auditory sensitivity, these bats do progressively increase pulse repetition rate throughout a target approaching sequence. Then, what is the adaptive value of increasing pulse repetition rate during echolocation? What are the underlying mechanisms for obtaining maximal information about the target features during increasing pulse repetition rate? This article reviews the electrophysiological studies of the effect of pulse repetition rate on multiple-parametric selectivity of neurons in the central nucleus of the inferior colliculus of the big brown bat, Eptesicus fuscus using single repetitive sound pulses and temporally patterned trains of sound pulses. These studies show that increasing pulse repetition rate improves multiple-parametric selectivity of inferior collicular neurons. Conceivably, this improvement of multiple-parametric selectivity of collicular neurons with increasing pulse repetition rate may serve as the underlying mechanisms for obtaining maximal information about the prey features for successful hunting by bats.     

The role of GABAergic inhibition in shaping directional selectivity of bat inferior collicular neurons determined with temporally patterned pulse trains

This study examined the role of GABAergic inhibition in shaping directional selectivity of neurons in the inferior colliculus of the big brown bat, Eptesicus fuscus. When determined with temporally patterned pulse trains at different pulse repetition rates, 93 inferior colliculus neurons displayed three types of directional selectivity curves. A directionally selective curve always showed a maximum to a certain azimuthal angle (the best angle). A hemifield curve showed a maximum to a range of contralateral azimuthal angles. A non-directional curve did not show a maximum to any particular azimuthal angles. Directional selectivity curves of 42% neurons changed from hemifield or non-directional to directionally selective and the best angles of 16-21% neurons shifted toward the midline with increasing pulse repetition rate of pulse trains. Directional selectivity curves of most (74%) neurons that discharged impulses to each pulse of a pulse train also became sharper with increasing pulse repetition rate of pulse trains. Bicuculline application produced more pronounced broadening of directional selective curves of inferior colliculus neurons at higher than at lower pulse repetition rates. As a result, pulse repetition rate-dependent directional selectivity of inferior colliculus neurons was abolished. Possible mechanisms and biological significance of these findings are discussed.     

Bicuculline application affects discharge pattern and pulse-duration tuning characteristics of bat inferior collicular neurons

This study examines the contribution of GABAergic inhibition to the discharge pattern and pulse duration tuning characteristics of 101 bat inferior collicular neurons by means of bicuculline application to their recording sites. When stimulated with single pulses, 56 (55%) neurons discharged 1 or 2 impulses (phasic responders), 42 (42%) discharged 3–10 impulses (phasic bursters) and 3 (3%) discharged impulses throughout the stimulus duration (tonic responders). Bicuculline application increased the number of impulses and changed the discharge patterns of 66 neurons. Using 50% difference between maximal and minimal responses as a criterion, the duration tuning characteristics of these neurons can be described as band-pass (20, 20%), long-pass (17, 17%), short-pass (33, 32%), and all-pass (31, 31%). Each band-pass neuron discharged maximally to a specific duration (the best duration) which was at least 50% larger than the neuron's responses to a long-duration pulse and a short-duration pulse. In contrast, each long- or short-pass neuron discharged maximally to a range of long or short duration pulses. Bicuculline application changed the duration tuning characteristics of 65 neurons. Possible mechanisms underlying duration tuning characteristics and the behavioral relevance to bat echolocation are discussed. Accepted: 4 November 1998     

The influence of stimulus duration on the delay tuning of cortical neurons in the FM bat,Myotis lucifugus

1. Echolocating bats use echo delay as the primary cue to determine target distance. During target-directed flight, the emitted pulses increase in repetition rate and shorten in duration as distance decreases. To determine how these parameters affect the delay tuning of neurons in the auditory cortex of the awake bat, Myotis lucifugus, we examined the responses of 104 delay-sensitive neurons as the pulse repetition rate (PRR) and duration were independently varied. Stimulus duration of 4, 2 and 1 ms and PRR of 5-100/s were used for both the pulse and echo to determine delay sensitivity. These parameter ranges span those used during the search, approach, and the initial terminal phases of echolocation. 2. As the stimulus duration was shortened, the range of PRRs for delay sensitivity was extended to higher rates in 41% of the neurons, narrowed or disappeared in 40%, and remained unchanged in 4%. The remaining 15% were not categorized since it was not possible to determine a trend in which the range of delay-sensitive PRRs changed with stimulus duration. 3. Three types of tracking neurons (i.e., neurons that change their best delay during target-directed flight) were found. For the first type, the best delay (BD) shortened with shorter stimulus duration, for the second type, BD shortened with both shorter stimulus durations and higher PRRs, and for the third type, BD shortened with higher PRRs. 4. These results suggest that the stimulus parameters of sonar emission influence delay tuning and hence processing by cortical neurons in FM bats.     

Echolocation sound features processed to provide distance information in the CF/FM bat,Noctilio albiventris: evidence for a gated time window utilizing both CF and FM components

Summary Bats of the speciesNoctilio albiventris, trained to discriminate differences in target distance, emitted pairs of pulses at a rate of 7–10/s, the first a constant frequency (CF) pulse of about 8 ms duration and 75 kHz frequency, followed after about 28 ms by a CF/FM pulse having a 6 ms, 75 kHz CF component that terminates in a 2 ms FM sweep to about 57 kHz.Loud free-running artificial pulses, simulating the bat's natural CF/FM echolocation sound, interfered with distance discrimination at repetition rates exceeding 5/s. Systematic modifications in the temporal and frequency structure of the artificial pulses resulted in orderly changes in the degree of interference. Artificial pulses simulating the natural CF or FM components alone had no effect, nor did 10/s white noise pulses, although constant white noise of the same intensity masked the behavior.Interference occurred when the CF of the artificial pulses was between 52 and 77 kHz, ending with a downward FM sweep of 25 kHz from the CF. For interference to occur there was a much more critical requirement that the FM sweep begin at approximately the frequency of the CF component. The FM sweep needed to be 11 kHz or greater bandwidth. Interference occurred when the duration of the CF component of the CF/FM artificial pulse was between 2 and 30 ms, with maximal effect between 10 and 20 ms. However, a brief (2.0 ms) CF signal 2–27 ms before an isolated FM signal was as effective as a continuous CF component of the same duration.When coupled with the bat's own emissions, artificial CF/FM pulses interfered if they occurred after the bat's CF/FM pulse and before the next natural emission. A 2 ms FM sweep alone was effective in interfering with distance discrimination when it came 8–27 ms after the onset of the bat's own CF/FM pulse. Neither CF/FM nor FM artificial pulses interfered when they began during the bat's own emission. A 10 ms CF pulse alone had no effect at any time.These findings indicate thatN. albiventris uses both the CF and FM components of its short-CF/FM echolocation sound for distance discrimination. The CF onset activates a gating mechanism that, during a narrowly defined subsequent time window, enables the nervous system to process FM pulse-echo pairs for distance information, within a fairly broad frequency range, as long as the frequencies of the CF and the beginning of the FM sweep are nearly identical.Abbreviations CF constant frequency - FM frequency modulation     

Sound-evoked oscillation and paradoxical latency shift in the inferior colliculus neurons of the big fruit-eating bat, <Emphasis Type="Italic">Artibeus jamaicensis</Emphasis>

Frequency tuning, temporal response pattern and latency properties of inferior colliculus neurons were investigated in the big fruit-eating bat, Artibeus jamaicensis. Neurons having best frequencies between 48–72 kHz and between 24–32 kHz are overrepresented. The inferior colliculus neurons had either phasic (consisting in only one response cycle at all stimulus intensities) or long-lasting oscillatory responses (consisting of multiple response cycles). Seventeen percent of neurons displayed paradoxical latency shift, i.e. their response latency increased with increasing sound level. Three types of paradoxical latency shift were found: (1) stable, that does not depend on sound duration, (2) duration-dependent, that grows with increasing sound duration, and (3) progressive, whose magnitude increases with increasing sound level. The temporal properties of paradoxical latency shift neurons compare well with those of neurons having long-lasting oscillatory responses, i.e. median inter-spike intervals and paradoxical latency shift below 6 ms are overrepresented. In addition, oscillatory and paradoxical latency shift neurons behave similarly when tested with tones of different durations. Temporal properties of oscillation and PLS found in the IC of fruit-eating bats are similar to those found in the IC of insectivorous bats using downward frequency-modulated echolocation calls.     

Serotonin modulates responses to species-specific vocalizations in the inferior colliculus

Neuromodulators such as serotonin are capable of altering the neural processing of stimuli across many sensory modalities. In the inferior colliculus, a major midbrain auditory gateway, serotonin alters the way that individual neurons respond to simple tone bursts and linear frequency modulated sweeps. The effects of serotonin are complex, and vary among neurons. How serotonin transforms the responses to spectrotemporally complex sounds of the type normally heard in natural settings has been poorly examined. To explore this issue further, the effects of iontophoretically applied serotonin on the responses of individual inferior colliculus neurons to a variety of recorded species-specific vocalizations were examined. These experiments were performed in the Mexican free-tailed bat, a species that uses a rich repertoire of vocalizations for the purposes of communication as well as echolocation. Serotonin frequently changed the number of recorded calls that were capable of evoking a response from individual neurons, sometimes increasing (15% of serotonin-responsive neurons), but usually decreasing (62% of serotonin-responsive neurons), this number. A functional consequence of these serotonin-evoked changes would be to change the population response to species-specific vocalizations.     

Frequency tuning and response latencies at three levels in the brainstem of the echolocating bat,Eptesicus fuscus

To determine the level at which certain response characteristics originate, we compared monaural auditory responses of neurons in ventral cochlear nucleus, nuclei of lateral lemniscus and inferior colliculus. Characteristics examined were sharpness of frequency tuning, latency variability for individual neurons and range of latencies across neurons.Exceptionally broad tuning curves were found in the nuclei of the lateral lemniscus, while exceptionally narrow tuning curves were found in the inferior colliculus. Neither specialized tuning characteristic was found in the ventral cochlear nuclei.All neurons in the columnar division of the ventral nucleus of the lateral lemniscus maintained low variability of latency over a broad range of stimulus conditions. Some neurons in the cochlear nucleus (12%) and some in the inferior colliculus (15%) had low variability in latency but only at best frequency.Range of latencies across neurons was small in the ventral cochlear nucleus (1.3–5.7 ms), intermediate in the nuclei of the lateral lemniscus (1.7–19.8 ms) and greatest in the inferior colliculus (2.9–42.0 ms).We conclude that, in the nuclei of the lateral lemniscus and in the inferior colliculus, unique tuning and timing properties are built up from ascending inputs.Abbreviations AVCN anteroventral cochlear nucleus - BF best frequency - CV coefficient of variation - DCN dorsal cochlear nucleus - FM frequency modulation - IC inferior colliculus - NLL nuclei of lateral lemniscus - PSTH post stimulus time histogram - PVCN posteroventral cochlear nucleus - SD standard deviation - SPL sound pressure level - VCN ventral cochlear nuclei - VNLLc ventral nucleus of the lateral lemniscus, columnar division     

Auditory Cortical Areas Activated by Slow Frequency-Modulated Sounds in Mice

Species-specific vocalizations in mice have frequency-modulated (FM) components slower than the lower limit of FM direction selectivity in the core region of the mouse auditory cortex. To identify cortical areas selective to slow frequency modulation, we investigated tonal responses in the mouse auditory cortex using transcranial flavoprotein fluorescence imaging. For differentiating responses to frequency modulation from those to stimuli at constant frequencies, we focused on transient fluorescence changes after direction reversal of temporally repeated and superimposed FM sweeps. We found that the ultrasonic field (UF) in the belt cortical region selectively responded to the direction reversal. The dorsoposterior field (DP) also responded weakly to the reversal. Regarding the responses in UF, no apparent tonotopic map was found, and the right UF responses were significantly larger in amplitude than the left UF responses. The half-max latency in responses to FM sweeps was shorter in UF compared with that in the primary auditory cortex (A1) or anterior auditory field (AAF). Tracer injection experiments in the functionally identified UF and DP confirmed that these two areas receive afferent inputs from the dorsal part of the medial geniculate nucleus (MG). Calcium imaging of UF neurons stained with fura-2 were performed using a two-photon microscope, and the presence of UF neurons that were selective to both direction and direction reversal of slow frequency modulation was demonstrated. These results strongly suggest a role for UF, and possibly DP, as cortical areas specialized for processing slow frequency modulation in mice.     

GABA能抑制调制大棕蝠下丘神经元对串声刺激的强度敏感性

本文采用不同重复率的串声刺激,模拟大棕蝠回声定位不同阶段听到的调频声纳信号,利用电生理方法和微电泳技术研究不同重复率串刺激条件下GABA能抑制对下丘神经元强度敏感性的影响。结果发现,随串刺激重复率的增加,有的神经元强度敏感性增强,有的神经元强度敏感性则降低。在不同串刺激条件下,微电泳荷包牡丹碱,神经元放电率均增加,随重复率增加强度敏感性增强或减弱的趋势消失,提示GABA能抑制调制下丘神经元对不同重复率串刺激反应的强度敏感性。串刺激强度在最低闽值附近时,微电泳荷包牡丹碱导致放电率增加的百分率最大,随串刺激强度增加,放电率增加的百分率逐渐减小。提示刺激强度较低时,GABA能抑制对下丘神经元强度敏感性的影响更有效。     

Bicuculline对小鼠中脑下丘听神经元

采用微电泳技术考察了GABAA受体拮抗剂荷包牡丹碱（bicuculline）,对小鼠中脑下丘听神经元强度-放电率曲线、频率调谐曲线和听空间反应域的影响。结果表明,微电泳bicuculline使听神经元的放电率显著提高,多数神经元的强度-放电率曲线变为单调型;听神经元频率调谐曲线加宽,并且对曲线上部的作用更加明显;听神经元的听空间反应域增大,方向敏感性降低。实验结果提示了GABA能抑制在下丘听信息处理中的重要作用。     

Spectrotemporal Response Properties of Core Auditory Cortex Neurons in Awake Monkey

So far, most studies of core auditory cortex (AC) have characterized the spectral and temporal tuning properties of cells in non-awake, anesthetized preparations. As experiments in awake animals are scarce, we here used dynamic spectral-temporal broadband ripples to study the properties of the spectrotemporal receptive fields (STRFs) of AC cells in awake monkeys. We show that AC neurons were typically most sensitive to low ripple densities (spectral) and low velocities (temporal), and that most cells were not selective for a particular spectrotemporal sweep direction. A substantial proportion of neurons preferred amplitude-modulated sounds (at zero ripple density) to dynamic ripples (at non-zero densities). The vast majority (>93%) of modulation transfer functions were separable with respect to spectral and temporal modulations, indicating that time and spectrum are independently processed in AC neurons. We also analyzed the linear predictability of AC responses to natural vocalizations on the basis of the STRF. We discuss our findings in the light of results obtained from the monkey midbrain inferior colliculus by comparing the spectrotemporal tuning properties and linear predictability of these two important auditory stages.     

Neural modelling of the encoding of fast frequency modulation

Frequency modulation (FM) is a basic constituent of vocalisation in many animals as well as in humans. In human speech, short rising and falling FM-sweeps of around 50 ms duration, called formant transitions, characterise individual speech sounds. There are two representations of FM in the ascending auditory pathway: a spectral representation, holding the instantaneous frequency of the stimuli; and a sweep representation, consisting of neurons that respond selectively to FM direction. To-date computational models use feedforward mechanisms to explain FM encoding. However, from neuroanatomy we know that there are massive feedback projections in the auditory pathway. Here, we found that a classical FM-sweep perceptual effect, the sweep pitch shift, cannot be explained by standard feedforward processing models. We hypothesised that the sweep pitch shift is caused by a predictive feedback mechanism. To test this hypothesis, we developed a novel model of FM encoding incorporating a predictive interaction between the sweep and the spectral representation. The model was designed to encode sweeps of the duration, modulation rate, and modulation shape of formant transitions. It fully accounted for experimental data that we acquired in a perceptual experiment with human participants as well as previously published experimental results. We also designed a new class of stimuli for a second perceptual experiment to further validate the model. Combined, our results indicate that predictive interaction between the frequency encoding and direction encoding neural representations plays an important role in the neural processing of FM. In the brain, this mechanism is likely to occur at early stages of the processing hierarchy.     

GABA-mediated echo duration selectivity of inferior collicular neurons of Eptesicus fuscus, determined with single pulses and pulse–echo pairs

When insectivorous bats such as Eptesicus fuscus emit ultrasonic signals and analyze the returning echoes to hunt insects, duration selectivity of auditory neurons plays an important role in echo recognition. The success of prey capture indicates that they can effectively encode progressively shortened echo duration throughout the hunting process. The present study examines the echo duration selectivity of neurons in the central nucleus of the bat inferior colliculus (IC) under stimulation conditions of single pulses and pulse–echo (P–E) pairs. This study also examines the role of gamma-aminobutyric acid (GABA)ergic inhibition in shaping echo duration selectivity of IC neurons. The data obtained show that the echo duration selectivity of IC neurons is sharper when determined with P–E pairs than with single pulses. Echo duration selectivity also sharpens with shortening of pulse duration and P–E gap. Bicuculline application decreases and GABA application increases echo duration selectivity of IC neurons. The degree of change in echo duration selectivity progressively increases with shortening of pulse duration and P–E gap during bicuculline application while the opposite is observed during the GABA application. These data indicate that the GABAergic inhibition contributes to sharpening of echo duration selectivity of IC neurons and facilitates echo recognition by bats throughout different phases of hunting.