Ontogenesis of auditory fovea representation in the inferior colliculus of the Sri Lankan rufous horseshoe bat,Rhinolophus rouxi

Summary This report describes the ontogenesis of tonotopy in the inferior colliculus (IC) of the rufous horseshoe bat (Rhinolophus rouxi). Horseshoe bats are deaf at birth, but consistent tonotopy with a low-to-high frequency gradient from dorsolateral to ventromedial develops from the 2nd up to the 5th week. The representation of the auditory fovea is established in ventro-mediocaudal parts of the IC during the 3rd postnatal week (Fig. 3). Then, a narrow frequency band 5 kHz in width, comprising 16% of the bat's auditory range, captures 50–60 vol% of the IC (Fig. 3c). However, foveal tuning is 10–12 kHz (1/3 octave) lower than in adults; foveal tuning in females (65–68 kHz) is 2–3 kHz higher than in males (62–65 kHz). Thereafter, foveal tuning increases by 1–1.5 kHz per day up to the 5th postnatal week, when the adult hearing range is established (Figs. 4, 5). The increase of sensitivity and of tuning sharpness of single units also follows a low-to-high frequency gradient (Fig. 6).Throughout this development the foveal tuning matches the second harmonic of the echolocation pulses vocalised by these young bats. The results confirm the hypothesis of developmental shifts in the frequency-place code for the foveal high frequency representation in the IC.Abbreviations BF best frequency - CF constant frequency - FM frequency modulation - IC inferior colliculus - IHC inner hair cell; - OHC outer hair cell - RR Rhinolophus rouxi     

Corticofugal regulation of auditory sensitivity in the bat inferior colliculus

Under free-field stimulation conditions, corticofugal regulation of auditory sensitivity of neurons in the central nucleus of the inferior colliculus of the big brown bat, Eptesicus fuscus, was studied by blocking activities of auditory cortical neurons with Lidocaine or by electrical stimulation in auditory cortical neuron recording sites. The corticocollicular pathway regulated the number of impulses, the auditory spatial response areas and the frequency-tuning curves of inferior colliculus neurons through facilitation or inhibition. Corticofugal regulation was most effective at low sound intensity and was dependent upon the time interval between acoustic and electrical stimuli. At optimal interstimulus intervals, inferior colliculus neurons had the smallest number of impulses and the longest response latency during corticofugal inhibition. The opposite effects were observed during corticofugal facilitation. Corticofugal inhibitory latency was longer than corticofugal facilitatory latency. Iontophoretic application of γ-aminobutyric acid and bicuculline to inferior colliculus recording sites produced effects similar to what were observed during corticofugal inhibition and facilitation. We suggest that corticofugal regulation of central auditory sensitivity can provide an animal with a mechanism to regulate acoustic signal processing in the ascending auditory pathway. Accepted: 15 July 1998     

The inferior colliculus of the mustached bat has the frequency-vs-latency coordinates

In the mustached bat, the central auditory system contains FM–FM (delay-tuned) neurons which are specialized for processing target-distance information carried by echo delays. Mechanisms for creating the FM–FM neurons involve delay lines, coincidence detection and amplification. A neural basis for delay lines can be a map representing response latencies. The aim of the present study is to explore whether the central nucleus of the inferior colliculus has a latency axis incorporated into iso-best frequency slabs. Responses of single or multiple neurons were recorded from the inferior colliculus of unanesthetized mustached bats with tungsten-wire electrodes, and their response latencies were measured with tone bursts at their best frequencies and best amplitudes or 65 dB SPL. In the dorsoventral electrode penetrations across the inferior colliculus, response latency systematically shortens from ˜12 to ˜4␣ms. Tonotopic representation in the inferior colliculus is somewhat complex. Iso-best frequency slabs are tilted and/or curved, but they orient more or less ventrodorsally. Nevertheless, the latency axis is evident in each iso-best frequency slab, regardless of best frequency. The inferior colliculus has the frequency-vs-latency coordinates. Accepted: 2 October 1996     

生后发育过程中大鼠下丘脑NMDA受体各亚单位mRNA基因表达

中枢NMDA受体通道具有一种独特的门控方式,既受配基门控又受电压门控,而这种独特的门控通道又必须由各种亚单位形成的多样化的异寡聚体才具有一定的活性和功能.从胚胎形成到生后发育,中枢NMDA受体各亚单位及其所组成的异寡聚体具有各自不同的时空特性,其功能以及药理学特性也随发育过程而变化.本研究应用RT-PCR技术,研究在生后发育过程中听觉中枢神经系统下丘NMDA受体NR1、NR2A、NR2B和NR2CmRNA基因的表达,以期为在分子水平上揭示其在生后发育过程中的规律.     

The effect of monaural middle ear destruction on postnatal development of mouse inferior colliculus

This study examined the effect of monaural middle ear destruction on postnatal development of inferior collicular (IC) neurons of the laboratory mouse, Mus musculus. Monaural middle ear destruction was performed on juvenile mice and the density, number and size of IC neurons were determined at different postoperative ages. For electrophysiological study, collicular auditory response properties were always examined four weeks after operation. Monaural middle ear destruction produced larger neurons in the ipsilateral IC (relative to the operated ear) and smaller neurons in the contralateral IC of experimental mice in comparison with IC neurons of control mice. IC neurons of control mice typically had lower minimum thresholds and greater Q10 values than IC neurons of experimental mice. In experimental mice, neurons in the contralateral IC typically had longer latencies and higher minimum thresholds than neurons in the ipsilateral IC. Clear tonotopic organization was only observed for IC neurons of control mice. Possible mechanisms for these different observations are discussed.     

Encoding repetition rate and duration in the inferior colliculus of the big brown bat,Eptesicus fuscus

1. Encoding of temporal stimulus parameters by inferior collicular (IC) neurons of Eptesicus fuscus was studied by recording their responses to a wide range of repetition rates (RRs) and durations at several stimulus intensities under free field stimulus conditions. 2. The response properties of 424 IC neurons recorded were similar to those reported in previous studies of this species. 3. IC neurons were classified as low-pass, band-pass, and high-pass according to their preference for RRs and/or durations characteristic of, respectively, search, approach, or terminal phases of echolocation. These neurons selectively process stimuli characteristic of the various phases of hunting. 4. Best RRs and best durations were not correlated with either the BFs or recording depths This suggests that each isofrequency lamina is capable of processing RRs and durations of all hunting phases. 5. Responses of one half of IC neurons studied were correlated with the stimulus duty cycle. These neurons may preferentially process terminal phase information when the bat's pulse emission duty cycle increases. 6. While the stimulus RR affected the dynamic range and overall profile of the intensity rate function, only little effect was observed with different stimulus durations.     

Neurons in the inferior colliculus of the big brown bat show maximal amplitude sensitivity at the best duration

The big brown bats, Eptesicus fuscus, emit ultrasonic signals and analyze the returning echoes in multi-parametric domains to extract target features. The variation of different pulse parameters during hunting predicts that analysis of an echo parameter by bats is inevitably affected by other co-varying echo parameters. In this study, we presented data to show that the bat inferior collicular (IC) neurons have maximal amplitude sensitivity at the best duration (BD). A family of rate-amplitude function (RAF) of each IC neuron is plotted with the BD and non-BD sound pulses. The RAF plotted with BD pulses has sharper slope (SL) and smaller dynamic range (DR) than the RAF plotted with non-BD pulses has. All RAFs can be described as monotonic, saturated or non-monotonic. IC neurons with monotonic RAF are mostly recorded at deeper IC and they have the largest average BD, best amplitude (BA) and DR. Conversely, IC neurons with non-monotonic RAF are mostly recorded at upper IC and they have the smallest average BD, BA and DR. Low best frequency (BF) neurons at upper IC have shorter BD, smaller BA and DR than high BF neurons at deeper IC have. These data suggest that IC neurons that tune to an echo duration also have the greatest sensitivity to echo amplitude. These data also suggest that sensitivity in frequency, duration and amplitude appears to be orderly represented along the dorso-ventral axis of the IC.     

Coding of sinusoidally amplitude modulated acoustic stimuli in the inferior colliculus of the rufous horseshoe bat,Rhinolophus rouxi

Summary Single neuron responses to sinusoidally amplitude modulated (SAM) signals were studied in the inferior colliculus of the horseshoe bat,Rhinolophus rouxi.57% of the neurons responded to SAM stimuli with periodical discharges synchronized to the modulation cycle. The proportion of cells driven by amplitude modulated signals was independent of the best frequency of the neurons. Best modulation frequencies were at or below 100 Hz in about 70% of the neurons. Synchronized activity could be elicited by modulation frequencies up to 400 Hz.Best SAM responses were observed at stimulus intensities 10 dB above threshold. Generally the BMF of a neuron did not change with intensity. The BMF decreased with decreasing modulation depth of the amplitude modulation.A trend for a topographical organization of neurons according to best modulation frequencies was detected. The results did not reveal any significant specialization of the bat's auditory system for coding of amplitude modulations as compared to other mammals.Abbreviations BF best frequency - BMF best modulation frequency - CF constant frequency - FM frequency modulation - IC inferior colliculus - SAM sinusoidal amplitude modulation - SFM sinusoidal 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.     

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.     

Ontogenesis of the echolocation system in the rufous horseshoe bat,Rhinolophus rouxi (Audition and vocalization in early postnatal development)

1. The development of vocalization and hearing was studied in Sri Lankan horseshoe bats (Rhinolophus rouxi) during the first postnatal month. The young bats were caught in a nursing colony of rhinolophids in which birth took place within a two week period. 2. The new-born bats emitted isolation calls through the mouth. At the beginning these calls consisted of pure tones with frequencies below 10 kHz (Fig. 1). During the first postnatal week the call frequency increased to about 15 kHz, and the fundamental was augmented by two to four harmonics. No evoked potentials to pure tone stimuli could be elicited in the inferior colliculus of this age group, i.e., auditory processing at the midbrain level was not demonstrable. 3. Evoked potentials were first recorded in the second week, broadly tuned to 15-45 kHz, with a maximum sensitivity between 15-25 kHz. In the course of the second week, however, higher frequencies up to 60 kHz became progressively incorporated into the audiogram (Fig. 3). The fundamental frequency of the multiharmonic isolation calls, emitted strictly through the mouth, increased to about 20 kHz. 4. In the bats' third postnatal week an increased hearing sensitivity (auditory filter) emerged, sharply tuned at frequencies between 57 and 60 kHz (Fig. 4e). The same individuals were also the first to emit long constant frequency echolocation calls through the nostrils (Fig. 4c). The energy of the calls was arranged in harmonic frequency bands with the second harmonic exactly tuned to the auditory filter. These young bats continued to emit isolation calls through the mouth, which were, however, not harmonically related to the echolocation calls (Fig. 4b, d). 5. During the fourth week, both the auditory filter and the matched echolocation pulses (the second harmonic) shifted towards higher frequencies (Fig. 5). During the fifth week the fundamental frequency of the calls was progressively attenuated, and both the second harmonic of the pulses and the auditory filter reached the frequency range typical for adult bats of 73-78 kHz (Fig. 6). 6. The development of audition and vocalization is discussed with regard to possible interactions of both subsystems, and their incorporation into the active orientation system of echolocation.     

Multiple mechanisms shape selectivity for FM sweep rate and direction in the pallid bat inferior colliculus and auditory cortex

The inferior colliculus and auditory cortex of the pallid bat contain a large percentage of neurons that are highly selective for the direction and rate of the downward frequency modulated (FM) sweep of the bat’s echolocation pulse. Approximately 25% of neurons tuned to the echolocation pulse respond exclusively to downward FM sweeps. This review focuses on the finding that this selectivity is generated by multiple mechanisms that may act alone or in concert. In the inferior colliculus, selectivity for sweep rate is shaped by at least three mechanisms: shortpass or bandpass tuning for signal duration, delayed high-frequency inhibition that prevents responses to slow sweep rates, and asymmetrical facilitation that occurs only when two tones are presented at appropriate delays. When acting alone, the three mechanisms can produce essentially identical rate selectivity. Direction selectivity can be produced by two mechanisms: an early low-frequency inhibition that prevents responses to upward sweeps, and the same asymmetrical two-tone inhibition that shapes rate tuning. All mechanisms except duration tuning are also present in the auditory cortex. Discussion centers on whether these mechanisms are redundant or complementary.     

Ontogenesis of alpha-amylase in rat parotid gland during postnatal development

Changes in alpha-amylase (alpha-1,4-glucan-4-glucanohydrolase, EC 3.2.1.1) of parotid gland were investigated during postnatal development of the rat. Modifications in amylase activity after birth allow the distinction of three stages which can be correlated with the morphologic development of the parotid gland. Significant sexual differences in the evolution of alpha-amylase activity were found. During the first stage (from birth to the 20th day) there is a higher increase in females, while males have a more pronounced increment in the second stage (from the 20th to the 30th day). By means of gel electrophoresis of parotid extracts, four molecular forms of amylase can be separated. The slowest migrating band (Form 1) is not detected at the initial stage.     

Frequency tuning, latencies, and responses to frequency-modulated sweeps in the inferior colliculus of the echolocating bat, Eptesicus fuscus

Neurons in the inferior colliculus (IC) of the awake big brown bat, Eptesicus fuscus, were examined for joint frequency and latency response properties which could register the timing of the bat's frequency-modulated (FM) biosonar echoes. Best frequencies (BFs) range from 10 kHz to 100 kHz with 50% tuning widths mostly from 1 kHz to 8 kHz. Neurons respond with one discharge per 2-ms tone burst or FM stimulus at a characteristic latency in the range of 3–45 ms, with latency variability (SD) of 50 μs to 4–6 ms or more. BF distribution is related to biosonar signal structure. As observed previously, on a linear frequency scale BFs appear biased to lower frequencies, with 20–40 kHz overrepresented. However, on a hyperbolic frequency (linear period) scale BFs appear more uniformly distributed, with little overrepresentation. The cumulative proportion of BFs in FM₁ and FM₂ bands reconstructs a scaled version of the spectrogram of FM broadcasts. Correcting FM latencies for absolute BF latencies and BF time-in-sweep reveals a subset of IC cells which respond dynamically to the timing of their BFs in FM sweeps. Behaviorally, Eptesicus perceives echo delay and phase with microsecond or even submicrosecond accuracy and resolution, but even with use of phase-locked FM and tone-burst stimuli the cell-by-cell precision of IC time-frequency registration seems inadequate by itself to account for the temporal acuity exhibited by the bat. Accepted: 21 June 1997     

抑制性频谱整合对大棕蝠下丘神经元声强敏感性的影响

自由声场条件下 ,采用特定双声刺激方法研究了不同频率通道之间的非线性整合对下丘神经元声强敏感性的调制作用。实验在 1 2只麻醉与镇定的大棕蝠 (Eptesicusfuscus)上进行 ,双电极同步记录 2个配对神经元的声反应动作电位。主要结果如下 :1 )所获 1 1 0个 (5 5对 )配对神经元中 ,85 5 %表现为抑制性频谱整合作用 ,其余 1 4 5 %为易化性频谱整合 ;2 )阈上 1 0dB (SPL)放电率抑制百分比与神经元最佳频率 (BF)及记录深度呈负相关 ;3)抑制效率随声刺激强度升高而逐步下降 ;4 )当掩蔽声分别位于神经元兴奋性频率调谐曲线(FTC)内 (MSin) /外 (MSout)时 ,其抑制效率存在差异。前者的放电率抑制百分比及声反应动力学范围(DR)下降百分比均显著高于后者 ;5 )抑制性频谱整合导致 3类DR改变 :6 1 6 %为下降、 1 0 9%增加、另有2 7 5 %变化小于 1 0 %。本结果进一步支持如下设想 :下丘不同频率通道之间的抑制性频谱整合参与了对强度编码的主动神经调制活动     

Analysis of ERK1/2 kinases in the inferior colliculus of rats genetically prone to audiogenic seizures during postnatal development

The activity of ERK1/2 kinases in the quadrigemina inferior colliculus of Krushinsky–Molodkina rats of different age, which are characterized by an increased seizure readiness compared to Wistar rats, was analyzed. An increased (probably genetically determined) activity of these enzymes during the development of epileptiform activity in ontogeny was found, which may be the cause of abnormalities in the neurotransmitter system functioning.