Changes in RNA content in neurons of the superior olivary complex during monaural and binaural stimulation

The character of binaural competitive connections at the level of the superior olive was investigated cytospectrophotometrically in cats. As a result of monaural stimulation for 2 h or binaural stimulation for 1.5 h by rhythmic noise signal the RNA content in the neurons of the ipsilateral and contralateral medial and lateral nuclei increased significantly. The volume of functioning neurons in the nuclei studied either increased or remained the same as in the control. The increase in the RNA content in neurons of both the ipsilateral and contralateral medial and lateral nuclei suggests a uniform distribution of binaurally converging connections on the neurons of these nuclei. The results also suggest that the accumulation of cytoplasmic RNA takes place in response not only to excitation, but also to inhibition.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 10, No. 1, pp. 67–74, January–February, 1978.     

Topical organization of the cat inferior colliculus under conditions of the simulated sound source motion

Binaurally presented click trains with gradually changing ITDs were used to imitate the sound source motion. Focal EP-trains from the cat inferior colliculus, IC, were recorded, and changing amplitude of the successive EPs following successive clicks was taken for estimation of the motion effect (ME). It was shown that a) in 25% of cases the ME depends on the motion direction; b) the ME is better presented ipsilaterally in respect to the recording point, with equal dependence on motion velocity both on the left and on the right; c) the ME phenomenon is connected with prevalence of contralateral afferentation as compared to the ipsilateral one; d) the ME is connected with inhibitory phenomena such as binaural suppression of monaural afferentation; e) the ME of the best amplitude is most pronounced in the ventrolateral area of the IC central nucleus, its localisation being near to that of the great multipolar neurones, whereas slightly and moderately pronounced MEs are presented rather uniformly over the central nucleus like the "basic" neurones are; MEs of different pronouncement are supposed to depend differently on effectiveness of ipsi- and contralateral stimulation--in accordance with properties of multipolar and "basic" neurons.     

Evoked potentials recorded from brain-stem nuclei in awake cat: interaction of tone bursts and continuous tone

Previous work indicated that components of the auditory thalamocortical potential evoked by a brief binaural tone burst could be enhanced by certain stimulus combinations, e.g., a brief tone burst in the presence of a continuous tone. The principal questions of the present study were whether enhaced components could be obtained caudal to thalamocortex and whether monaural stimuli would be effective in producing enhancement. Eight cats received electrodes in cochlear nucleus and the nucleus of the inferior colliculus. Custom earmolds were made for each ear of each animal. The median attenuation produced by the earmolds was 35 dB and the use of a single earmold approximated monaural stimulation. Auditory evoked potentials were recorded from the electrodes while the animals were unanesthetized but comfortably restrained. Brief 6.25 kHz tone bursts were presented against a background of silence or of a 4.96 kHz continuous tone. In the presence of the continuous tone, enhanced components were obtained from a majority of the electrodes in inferior colliculus but from none of the electrodes in cochlear nucleus. The late negative component in the colliculus potential was increased in amplitude while other components were reduced in amplitude by the continuous tone. The latencies of all components from all electrodes were increased by the presence of the continuous tone. It was concluded that enhancement effects could be obtained at the level of inferior colliculus, and that binaural stimulation does not appear to be necessary to produce enhanced components.     

Regularly firing neurons in the inferior colliculus have a weak interaural intensity difference sensitivity

The spike discharge regularity may be important in the processing of information in the auditory pathway. It has already been shown that many cells in the central nucleus of the inferior colliculus fire regularly in response to monaural stimulation by the best frequency tones. The aim of this study was to find how the regularity of units was affected by adding ipsilateral tone, and how interaural intensity difference sensitivity is related to regularity. Single unit recordings were performed from 66 units in the inferior colliculus of the anaesthetized guinea pig in response to the best frequency tone. Regularity of firing was measured by calculating the coefficient of variation as a function of time of a unit’s response. There was a positive correlation between coefficient of variation and interaural intensity difference sensitivity, indicating that highly regular units had very weak and irregular units had strong interaural intensity difference sensitivity responses. Three effects of binaural interaction on the sustained regularity were observed: constant coefficient of variation despite change in rate (66% of the units), negative (20%) and positive (13%) rate–CV relationships. A negative rate-coefficient of variation relationship was the dominant pattern of binaural interaction on the onset regularity.     

Structural Changes and Lack of HCN1 Channels in the Binaural Auditory Brainstem of the Naked Mole-Rat (Heterocephalus glaber)

Naked mole-rats (Heterocephalus glaber) live in large eu-social, underground colonies in narrow burrows and are exposed to a large repertoire of communication signals but negligible binaural sound localization cues, such as interaural time and intensity differences. We therefore asked whether monaural and binaural auditory brainstem nuclei in the naked mole-rat are differentially adjusted to this acoustic environment. Using antibody stainings against excitatory and inhibitory presynaptic structures, namely the vesicular glutamate transporter VGluT1 and the glycine transporter GlyT2 we identified all major auditory brainstem nuclei except the superior paraolivary nucleus in these animals. Naked mole-rats possess a well structured medial superior olive, with a similar synaptic arrangement to interaural-time-difference encoding animals. The neighboring lateral superior olive, which analyzes interaural intensity differences, is large and elongated, whereas the medial nucleus of the trapezoid body, which provides the contralateral inhibitory input to these binaural nuclei, is reduced in size. In contrast, the cochlear nucleus, the nuclei of the lateral lemniscus and the inferior colliculus are not considerably different when compared to other rodent species. Most interestingly, binaural auditory brainstem nuclei lack the membrane-bound hyperpolarization-activated channel HCN1, a voltage-gated ion channel that greatly contributes to the fast integration times in binaural nuclei of the superior olivary complex in other species. This suggests substantially lengthened membrane time constants and thus prolonged temporal integration of inputs in binaural auditory brainstem neurons and might be linked to the severely degenerated sound localization abilities in these animals.     

Response properties of auditory activated cells in the occipital cortex of the blind mole rat: an electrophysiological study

Previous studies have demonstrated that despite its blindness, the subterranean blind mole rat (Spalax ehrenbergi) possesses a noticeable lateral geniculate nucleus and a typical cyto-architectural occipital cortex that are reciprocally connected. These two areas, as revealed by the metabolic tracer 2-deoxyglucose, are activated by auditory stimuli. Using single unit recordings, we show that about 57% of 325 cells located within the occipital cortex of anesthetized mole rats responded to at least one of the following auditory stimuli — white noise, pure tones, clicks, and amplitude modulated tones — with the latter two being the most effective. About 85% of cells driven by either contralateral or ipsilateral stimulation also responded to binaural stimulation; about 13% responded only to binaural stimulation; and 2% were driven exclusively by contralateral stimulation. Comparing responsiveness and response strength to these three modes of stimulation revealed a contralateral predominance. Mean latency (±SD) of ipsilateral and contralateral responses were 48.5±32.6 ms and 33.5±9.4 ms, respectively. Characteristic frequencies could be divided into two distinct subgroups ranging between 80 and 125 Hz and between 2,500 and 4,400 Hz, corresponding to the most intensive spectral components of the vibratory intraspecific communication signals and airborne vocalizations.Abbreviations BMF best modulation frequency - CF characteristic frequency - 2-DG 2-deoxyglucose - dLGN dorsal lateral geniculate nucleus - IC inferior colliculus - LGN lateral geniculate nucleus - OC occipital cortex - MTF modulation transfer function - SAM sinusoidally amplitude modulation - SC superior colliculus     

Ascending and descending projections to the inferior colliculus in the rat

The ascending and descending projections to the central nucleus of the inferior colliculus (IC) were studied with the aid of retrograde transport of horseradish peroxidase (HRP). HRP-labelled cells were found in contralateral cochlear nuclei, where the majority of different cell types was stained. Few labelled cells were observed in the ipsilateral cochlear nuclei. HRP-positive neurones were found in all nuclei of the superior olivary complex on the ipsilateral side with the exception of the medial nucleus of the trapezoid body, which was never labelled either ipsilaterally or contralaterally. The largest concentration of HRP-labelled cells was usually observed in the ipsilateral superior olivary nucleus. Smaller numbers of labelled cells were present in contralateral nuclei of the superior olivary complex. Massive projections to the inferior colliculus were found from the contralateral and ipsilateral dorsal nucleus of the lateral lemniscus and ipsilateral ventral nucleus of the lateral lemniscus. Many neurones of the central and external nuclei of the contralateral inferior colliculus were labelled with HRP. Topographic organisation of the pathways ascending to the colliculus was expressed in the cochlear nuclei, lateral superior olivary nucleus and in the dorsal nucleus of the lateral lemniscus. HRP--positive cells were found in layer V of the ipsilateral auditory cortex, however, the evidence for topographic organisation was lacking.     

单耳堵塞对蝙蝠下丘GABA阳性反应神经元的影响

已有的研究表明,单耳堵塞可以使中脑下丘神经元的听空间反应特性改变,而神经元听空间特性的形成与中枢中的周边抑制有关,是神经元的兴奋性中心区域与抑制性周边区域相互作用的结果［１］。γ－氨基丁酸（Ｇａｍｍａ－ａｍｉｎｏｂｙｔｒｉｃａｃｉｄ,ＧＡＢＡ）作为中...     

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     

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

Summary 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).Abbreviations BF best frequency - Bic bicuculline - C control - CF constant frequency - CN cochlear nucleus - DNLL dorsal nucleus of the lateral lemniscus - FM frequency modulation - GABA gamma amino butyric acid - IC inferior colliculus - LSO lateral superior olive - Str strychnine     

A cytochemical study of binaurally converging connections in the nuclei of the cat superior olive]

Experiments of binaurally converging connections on the level of the superior olive nuclei was carried out on anesthetized cats by the method of quantitative cytospectrometry. A considerable increase in the cytoplasmic RNA content in the neurons of the ipsi- and contralateral medial and lateral nuclei was observed following a 2-hour monaural stimulation by rhythmical sound (noise) signals. The volume of the functioning neurons of the nuclei under study showed either no significant change or increased. The data obtained demonstrated the nature of functional organization of the binaurally converging connections.     

The reflection of the simulated movement of a sound source in the evoked potentials of the inferior colliculus in the cat

EP series from the cat's inferior colliculus were recorded following binaural stimulation with click series imitating sound source movement due to variation of the interaural time delay (and thus evoking in man the sensation of the moving fused auditory image, FI). The "movement effect" was evaluated as the change in the EP amplitude during the series. The movement effect itself as well as its predominance under conditions of the ipsilateral FI movement as compared to those of the contralateral movement, proved to be connected with greater effectiveness of the contralateral stimulation relative the ipsilateral one.     

成对短声不同间隔对单、双耳听觉脑干反应的影响——衍生听觉脑干反应方法的应用

采用具有不同间隔(0～32ms)的65dB nHL(正常听力水平)的成对短声刺激,记录20名正常人的单侧耳和两耳交替刺激的听觉脑干反应(ABR)。用计算机从成对短声反应中减去单一短声反应以提取衍生ABR。结果表明,单、双耳的衍生ABR V波振幅在成对短声间隔为0.2～1.5ms时,受到明显影响。单耳的减小54％～65％(P<0.01),两耳的减少46％～53％(P<0.01),但单、双耳的衍生ABR I波振幅未显示显著差异(P>0.05)。该结果说明,高位脑干通路在成对短声间隔为0.2～1.5ms时,不但对同侧耳的第2个短声反应能力降低,而且对来自对侧耳的第2个短声也如此。从而推断,在两耳交替刺激的耳间短声间隔小于2ms范围时,在下丘部位可能存在两耳交互作用。结果还提示,临床检查ABR时,采用的短声刺激间隔至少不应小于30ms。     

Effect of destruction of the inferior colliculus on function of the echolocation system in horseshoe bats

The effect of unilateral and bilateral destruction of the inferior colliculus on the sensitivity of the auditory system, on parameters of the sonor signals, and on Doppler shift compensation in echo signals was studied in experiments on horseshoe bats (Rhinolophus ferrum-equinum). The results show that complete bilateral destruction of the inferior colliculus in bats does not lead to total disturbance of function of the auditory system but it sharply reduces the sensitivity of that system, as shown by a decrease in the maximal obstacle detection range and inability to respond to an insect emitting a feeble sound. It can also be concluded that the inferior colliculus plays a direct part in maintenance of the emission frequency and that different parts of the inferior colliculus play different roles in this process. The Doppler shift compensation effect requires preservation of the integrity of not less than half of the central nucleus of at least one inferior colliculus.A. A. Ukhtomskii Physiological Institute, A. A. Zhdanov State University, Leningrad. Translated from Neirofiziologiya, Vol. 12, No. 4, pp. 375–381, July–August, 1980.     

Response of Neurons of The Inferior Colliculi of Cats To Changes in the Intensity of One of The Monaural Components of Binaurally Presented Stimuli

In spite of the large quantity of psychophysiological investigations of binaural hearing, a systematic study of its concrete mechanisms has only been begun comparatively recently (cf. review [1]). In particular, changes in neuron response under conditions of lateral differentiation brought about by varying the intensity of one of the monaural components of binaurally presented stimuli have been studied in detail only at the site of the superior olivary body (3, 4, 6) — the first section of the auditory system — where afferent fibers from the right and left cochlea converge (7, 9). Published data provide evidence that variations in the intensity of one of the monaural components of binaurally presented signals has an appreciable effect on the firing rate of single neurons of the inferior colli cuius (5, 8), which is situated directly behind the superior olivary body of the auditory system. in the present report, the types of response of neurons in the inferior colliculus to variations in the intensity of one of the monaural components of binaurally presented stimuli are systematically investigated.     

Monaural phasic sensitivity of auditory system neurons in the cat

During monaural stimulation, studies have been made of impulse reactions in neurons from the cochlear nucleus and inferior colluculus to changes in the phase value (psi) of one of the harmonics in two-tone signals. It was shown that about 70% of the neurons from the cochleaur nucleus with a tonic discharge pattern may sharply change their activity (from maximum one up to a complete inhibition) due to changes in psi value. Unlike the cochlear nucleus units, neurons from the inferior colliculus change their tonic or burst-like activity rather seldomly and to a slight extent. At the same time, their phasic on- and especially off-responses exhibit high sensitivity to changes in psi value.     

Commissural connections mediate inhibition for the computation of interaural level difference in the barn owl

Summary In the barn owl (Tyto alba), the posterior nucleus of the ventral lateral lemniscus (VLVp) is the first site of binaural convergence in the pathway that processes interaural level difference (ILD), an important sound-localization cue. The neurons of VLVp are sensitive to ILD because of an excitatory input from the contralateral ear and an inhibitory input from the ipsilateral ear. A previously described projection from the contralateral cochlear nucleus, can account for the excitation. The present study addresses the source of the inhibitory input.We demonstrate with standard axonal transport methods that the left and right VLVps are interconnected via fibers of the commissure of Probst. We further show that the anesthetization of one VLVp renders ineffective the inhibition that is normally evoked by stimulation of the ipsilateral ear. Thus, one cochlear nucleus (driven by the ipsilateral ear) appears to provide inhibition to the ipsilateral VLVp by exciting commissurally-projecting inhibitory neurons in the contralateral VLVp.Abbreviations ABL average binaural level - CP commissure of Probst - DNLL dorsal nucleus of the lateral lemniscus - IC inferior colliculus - ILD interaural level difference - IPc nucleus isthmi, pars parvocellularis - ITD interaural time difference - LSO lateral superior olive - MNTB medial nucleus of the trapezoid body - NA nucleus angularis - SL nucleus semilunaris - VLVa nucleus ventralis lemnisci lateralis, pars anterior - VLVp nucleus ventralis lemnisci lateralis, pars posterior     

Fast-acting excitatory amino acids are involved in the enhancement of the aversiveness of the electrical stimulation of the inferior colliculus by systemic injections of muscimol

Gradual increases in the electrical stimulation of the inferior colliculus produces progressive aversive responses from vigilance, through freezing, until escape. These responses are probably mediated by excitatory amino acids (EAA) mechanisms as microinjection of glutamate into the inferior colliculus can trigger freezing responses while microinjections of NMDA cause a mixture of immobility and escape responses. Moreover, it has been shown that the neural substrates for defensive behavior in this structure are regulated by GABA-benzodiazepine mechanisms. Indeed, these responses are depressed by muscimol and midazolam locally injected into the inferior colliculus. In this work we were interested in knowing how GABAergic mechanisms interact with the EAA-mediated neural substrates of aversion generated at the inferior colliculus level. We found that while intraperitoneal injections of muscimol caused the expected antiaversive effects, unexpectedly systemic injections of muscimol enhanced the aversive reactions induced by electrical stimulation of the inferior colliculus of rats. Local injections into the central nucleus of the inferior colliculus of GDEE-an AMPA/kainate receptor antagonist-inhibited whereas AP7-a NMDA receptor antagonist-did not influence these responses. It is suggested that systemic injections of muscimol inhibit GABAergic inputs to the inferior colliculus. The removal of these inhibitory influences reduce the well-known tonic inhibitory control exerted by GABAergic mechanisms on the neural substrates of aversion of the inferior colliculus. Activation of these neural substrates by fast-acting AMPA/kainate receptors trigger the initial steps of the defense reaction in the central nucleus of the inferior colliculus.     

Effect of transient neonatal hypothyroidism on the free aspartic acid, glutamic acid, and GABA content of different central auditory regions in the rat

The aspartic acid, glutamic acid, and gamma-aminobutyric acid (GABA) contents were determined in four central auditory system regions in rats with transient neonatal hypothyroidism compared with control ones: the ventral and dorsal parts of the cochlear nucleus, the central nucleus of the inferior colliculus, the auditory cortex, and in an extra-auditory structure, the substantia nigra pars reticulata. The animals were sacrificed at 50 days of age, brain tissue samples were taken out by microdissection, and the free amino acids were extracted. The amino acid content was assessed by double-isotope labelling following two-dimensional thin-layer chromatography separation. GABA content was significantly decreased in both cochlear nucleus regions and glutamic acid was elevated in the inferior colliculus. Neonatal hypothyroidism had no significant effect on the aspartic acid levels in the regions studied. The results suggest an effect of neonatal hypothyroidism on regional contents of free amino acids known as candidate neurotransmitters in the auditory system.     

Coding of sound location and frequency in the auditory midbrain of diurnal birds of prey,families accipitridae and falconidae

Summary The coding of sound frequency and location in the avian auditory midbrain nucleus (nMLD) was examined in three diurnal raptors: the brown falcon (Falco berigora), the swamp harrier (Circus aeruginosus) and the brown goshawk (Accipiter fasciatus). Previously this nucleus has been studied with free field stimuli in only one other species, the barn owl (Tyto alba).We found some parallels between the organisation of nMLD in the diurnal raptors and that reported in the barn owl in that the central region of nMLD was tonotopically organised and contained cells that did not encode location, and the lateral region (nMLDl) contained cells which were sensitive to stimulus position. However, unlike the barn owl, which has units with circumscribed receptive fields, cells sensitive to stimulus location had large receptive fields which were restricted in azimuth but not in elevation (hemifield units). Such cells could not provide an acoustic space map in which both azimuthal and elevational dimensions were represented, but there was a tendency for units with contralateral borders to be found superficially, and those with ipsilateral borders to be found deep, in nMLDl. Hemifield units displayed receptive field properties consistent with the directional properties of the tympana in the presence of sound transmission through the interaural canal, if there is a central mechanism which is sensitive to interaural intensity differences.Abbreviations nMLD nucleus mesencephalicus lateralis pars dorsalis - SPL sound pressure level re 20 Pa - nMLDl lateral region of nMLD - ICC central nucleus of the inferior colliculus - ICX external nucleus of the inferior colliculus - IID interaural intensity difference - EI excitatory inhibitory