Neural representations of the axis of acoustic particle motion in nucleus centralis of the torus semicircularis of the goldfish, <Emphasis Type="Italic">Carassius auratus</Emphasis>

Experiments examined differential coding of acoustic particle motion axis in the auditory midbrain of goldfish. Animals were exposed to vibratory stimuli varying in axis orientation as action potentials were recorded from single units in the central neuropil of nucleus centralis in the torus semicircularis. Response magnitudes as a function of stimulation axis were visualized in three dimensional plots called directional response profiles. These are generally comparable to directional responses observed among primary saccular afferents in having substantially vertical orientations. Distortions in shape from the peripheral patterns indicate neural information processing. A three-dimensional model was used to evaluate the hypothesis that responses in the auditory midbrain reflect the convergence of excitatory and inhibitory primary afferent-like responses. Model afferent inputs were generated and combined arithmetically. This analysis gives insight into the mechanisms of information processing that appear to occur in brainstem nuclei. The lack of diversity in best axis directions suggests that this mechanism alone cannot account for directional hearing abilities in this species. The roles that this directional representation and processing may play in directional hearing and sound source localization are not yet clear. Implications of these data on current models of fish directional hearing are discussed.     

Influence of descending forebrain projections on processing of acoustic signals and audiomotor integration in the anuran midbrain

We tested the role of descending projections for auditory processing and audiomotor integration in the anuran torus semicircularis. Intracellular recordings were made from isolated brain preparations, impaled neurons were stained. Auditory neurons responded to electrical stimulation of striatum and/or dorsal thalamus, they integrated forebrain and auditory nerve inputs. High frequency stimulation in striatum or thalamus changed the auditory response of torus neurons located in the laminar subnucleus. Our results suggest that the laminar nucleus is the primary target of forebrain projections, which provides a basis for modulation of acoustically guided behaviour.     

Postmetamorphic changes in auditory sensitivity of the bullfrog midbrain

During metamorphosis, the lateral line system of ranid frogs (Rana catesbeiana) degenerates and an auditory system sensitive to airborne sounds develops. We examined the onset of function and developmental changes in the central auditory system by recording multi-unit activity from the principal nucleus of the torus semicircularis (TSp) of bullfrogs at different postmetamorphic stages in response to tympanically-presented auditory stimuli. No responses were recorded to stimuli of up to 95 dB SPL from latemetamorphic tadpoles, but auditory responses were recorded within 24 hours of completion of metamorphosis. Audiograms from froglets (SVL < 5.5 cm) were relatively flat in shape with high thresholds, and showed a decrease in most sensitive frequency (MSF) from about 2500 Hz to about 1500 Hz throughout the first 7–10 days after completion of metamorphosis. Audiograms from frogs larger than 5.5 cm showed continuous downward shifts in MSF and thresholds, and increases in sharpness around MSF until reaching adult-like values. Spontaneous activity in the TSp increased throughout postmetamorphic development. The torus increased in volume by approximately 50% throughout development and displayed changes in cell density and nuclear organization. These observations suggest that the onset of sensitivity to tympanically presented airborne sounds is limited by peripheral, rather than central, auditory maturation.Abbreviations CF characteristic frequency - MSF most sensitive frequency - PB phasic burst - PL primary like - S sustained - SVL snout-vent length - TS torus semicircularis - TSl laminar nucleus of TS - TSm magnocellular nucleus of TS - TSp principal nucleus of TS - TW tympanic width     

Sharpening of directional responses along the auditory pathway of the oyster toadfish, <Emphasis Type="Italic">Opsanus tau</Emphasis>

Our previous studies have shown that the peripheral auditory system of the toadfish encodes the direction of a sound source. Here, we compare directional responses of peripheral saccular afferents, cells in the descending octaval nucleus (DON) of the medulla, and the torus semicircularis (TS) of the midbrain. Recording locations in the brain were labeled with neurobiotin to confirm the site. To compare directional responses among cells, we calculated an index [sharpening ratio (SR)] that weights the relative strength of responses to the best direction for that cell and to the adjacent stimulus angles tested. Unsharpened saccular afferents tend to have a cosinusoidal directional response pattern (DRP) with an expected SR of 0.87. In DON, more than 60% of the cells exhibited directional sharpening (defined as SR <0.8). In TS, more than 80% of the cells exhibited directional sharpening. We conclude that directional auditory sharpening first occurs in DON and some additional sharpening occurs in the ascending pathway to the midbrain, particularly in azimuth. The sharpening of directional selectivity is likely to be an important component of the neural computations underlying directional hearing.     

The cytoarchitecture of the torus semicircularis in the teleost Barbus meridionalis

The torus semicircularis of Barbus meridionalis is composed of two nuclei, the nucleus centralis and nucleus lateralis. Its cytoarchitecture was studied in sections stained by Nissl and Golgi-Colonnier techniques. In the nucleus lateralis two portions were identified: the ‘pars lateralis’ and the ‘pars medialis.’ Cytoarchitecturally, both portions are identical. They exhibit a layered structure in which there is an alternation of cell-poor and cell-rich laminae designated as: (1) the subependymal layer; (2) the layer of small cells; (3) the fibrillar layer; and (4) the layer of disperse cells. The subependymal layer consists of fine fibers and some small rounded-ovoid cells whose dendritic prolongations course horizontally or ventrally. The second layer has small, densely-packed cells with rounded-ovoid and triangular somata and a main dendritic trunk that courses ventrally. The third layer contains dendritic fields of the cells of layer two and of cells from layer four. The fourth layer is composed of fusiform neurons with two dendritic trunks of equal thickness, rounded-ovoid neurons with one or two main dendritic trunks and multipolar triangular stellate neurons with equal dendritic trunks. The nucleus centralis comprises a fibrillar cortex with a structure identical to that of the subependymal layer. There is also a cellular region with the same cell types as those found in the nucleus lateralis. These two nuclei thus compose the torus semicircularis of the barbel. They exhibit the same cytological characteristics and both are differentiated by their cytoarchitectural and functional orders.     

Acoustic response properties of single units in the torus semicircularis of the goldfish,Carassius auratus

Single units of the goldfish torus semicircularis (TS) were recorded in response to pure tones. Response areas (RA) were obtained by recording the number of spikes evoked by tones in a range of frequencies and levels within the units' dynamic range. RAs gave estimates of best sensitivity (BS), characteristic frequency (CF), most excitatory frequency at each level (BF), and Q_10dB. Peri-stimulus-time histograms (PSTH), interspike interval histograms (ISIH), and period histograms were obtained at various frequencies and levels to describe the units' temporal response patterns.The distribution of CF is nonuniform with modes at 155, 455, and 855 Hz. The distribution of the coefficient of synchronization to standard tones is also nonuniform, revealing a dichotomy between units with little or no phase-locking and those that phase-lock strongly. PSTHs for units without significant phase-locking vary widely and include patterns resembling those of the mammalian auditory brainstem. Compared with saccular afferents, torus units tend to have lower spontaneous rates, greater sensitivity, and sharper tuning. Unlike saccular afferents, BF is independent of level for most torus units. Some torus units are similar to saccular afferents while others reveal significant transformations of information between the periphery and the midbrain.Abbreviations BF best frequency - BS best sensitivity - CF characteristic frequency - ISIH inter-spike interval histogram - PSTH peri-stimulus-time histogram - RA response area - TS torus semicircularis     

Does the magnocellular octaval nucleus process auditory information in the toadfish, Opsanus tau?

Previous work on auditory processing in Opsanus tau has focused on the descending octaval nucleus; however, the magnocellular octaval nucleus receives similar inputs from the otolithic endorgans. The purpose of this study was to assess whether cells in any of the three subdivisions of the magnocellular nucleus respond to auditory frequencies and encode sound source direction. Extracellular recording sites were chosen based on anatomical landmarks, and neurobiotin injections confirmed the location of auditory sites in subdivisions of the magnocellular nucleus. In general, the auditory cells in M2 and M3 responded best to frequencies at or below 100 Hz. Most auditory cells responded well to directional stimuli presented along axes in the horizontal plane. Cells in M3 (not M2) also responded to lateral line stimulation, consistent with otolithic endorgan and lateral line inputs to M3. The convergence of auditory and lateral line inputs in M3, the lack of Mauthner cells in this species, and previous evidence that the magnocellular nucleus does not contribute to ascending auditory pathways suggest to us that the large cells of M3 may play a role in rapid behavioral responses to particle motion stimuli in oyster toadfish.     

Integration of ascending and descending inputs in the auditory midbrain of anurans.

In anurans, the midbrain torus semicircularis is involved in auditory processing and audio-motor integration. In this study, we examined the influence of descending forebrain projections on the auditory response properties and hence the audiomotor transmission of mesencephalic interface neurons. In order to investigate response integration, we performed intracellular recordings from torus neurons in an isolated brain preparation of Discoglossus pictus and Bombina orientalis and stimulated the auditory nerve, striatum, and the dorsal thalamus electrically with single pulses. Stimulation of all three sites could evoke responses in torus neurons that were either excitatory, inhibitory, or a mixture of both, with durations of up to several hundred milliseconds. Further, striatum and thalamus were activated by pulse trains (10-20 Hz, 50 pulses) immediately before stimulating the auditory nerve with single pulses. Thus, responses of torus neurons to "auditory" input were facilitated or suppressed for up to 2 min by striatum stimulation or only suppressed by thalamus stimulation. Intracellular labeling of recorded neurons revealed that response modulation by descending input mostly occurred in laminar nucleus neurons. These results suggest that descending forebrain projections to mesencephalic audiomotor interface neurons may play an important role in modifying acoustically guided behavior in anurans.     

不同物种雄性鸣声刺激下雌性凹耳蛙脑组织中ZENK的表达

通过免疫组化SABC法,研究在背景噪音刺激、雄性凹耳蛙(Odorrana tormota)声音刺激以及雄性大绿臭蛙(O. graminea)声音刺激下ZENK蛋白在雌性凹耳蛙脑部的表达。结果显示,上述三种声音刺激下雌性凹耳蛙的端脑、间脑和小脑区域均未发现ZENK蛋白的阳性细胞。在雄性凹耳蛙声音刺激下,雌性凹耳蛙中脑的视顶盖、脚间核、中脑深部核、被盖、半环隆枕以及延脑的孤束核、中缝核、网状核、网状结构、上橄榄核部位均有阳性细胞分布,在大绿臭蛙声音刺激下,这些部位的阳性细胞数量明显减少。本实验中ZENK蛋白阳性细胞表达部位的一部分是位于与两栖类处理听觉信息有关的中脑半环隆枕、被盖以及延脑的上橄榄核,由此可推断,在雌性凹耳蛙ZENK蛋白与听觉信息处理有着密不可分的联系。     

Efferent neurons of the auditory center in the midbrain of the frog Rana ridibunda

Individual cells which produce projections from the torus semicircularis in the frog have been visualized after injection of horseradish peroxidase (HRP) to various thalamic and isthmal areas. Labeled toral cells were observed if HRP had been injected to the posterodorsal areas of the thalamus or to the isthmal areas where lateral lemniscus fibers and cells of the premature lateral lemniscal nucleus are situated. Medium and large size cells in the rostrolateral torus semicircularis were mainly labeled. Thalamic injections of the HRP produced more labeled cells in the lateral part of the magnocellular nucleus, whereas isthmal injections produced labeled cells mainly in the lateral part of the laminar nucleus. A few HRP containing cells were observed in the principal nucleus of the torus. Specificity of the neuronal organisation of the auditory pathway in amphibians is discussed.     

Responses of auditory brainstem neurons in the grassfrog to clicks

Single unit recordings were made in the dorsal medullary nucleus and in the torus semicircularis of the immobilized grassfrog. The natural calls have a periodic pulsatile structure. To investigate the coding of pulse repetition rate periodic click trains with varying pulse repetition rate and an ensemble of clicks distributed randomly in time were used as stimuli. In the dorsal medullary nucleus strong time-locking to clicks was found. Most units showed an activation followed by suppression response. Some units showed a preference for pulse repetition rates matching their low-frequency sensitivity. In the torus semicircularis part of the units showed responses similar to dorsal medullary nucleus units. Other response types were activation irrespective of pulse repetition rate, and suppression followed by activation. The responses to the two stimulus ensembles were more compatible in the dorsal medullary nucleus than in the torus semicircularis.     

The jamming avoidance response ofEigenmannia: properties of a diencephalic link between sensory processing and motor output

Summary Brain regions participating in the control ofEigenmannia's electric organ discharge frequency were localized by electrical microstimulation and anatomically identified by means of horseradish peroxidase deposition. A diencephalic region was found which, when stimulated, caused electric organ discharge (EOD) frequency increases of similar magnitude and time course as the frequency increases seen during the jamming avoidance response. Single unit recordings from this region revealed one cell type which preferentially responded to stimuli that cause the acceleration phase of the jamming avoidance response (electric organ discharge frequency increase). A second cell type responded preferentially to stimuli which cause EOD frequency decrease, and both cell types were tuned to stimuli which evoked maximal jamming avoidance behaviors.The results of the horseradish peroxidase experiments showed that the recording and stimulation sites correspond to the previously described nucleus electrosensorius. Our results confirm the earlier finding that this nucleus receives output from the torus semicircularis and we also found that the N. electrosensorius projects to the mesencephalic prepacemaker nucleus. The prepacemaker projects to the medullary pacemaker nucleus which generates the commands that evoke electric organ discharges.The anatomical and physiological results described here establish this diencephalic region as a link between the major sensory processing region for the jamming avoidance response, the torus semicircularis, and a mesencephalic pre-motor region, the prepacemaker nucleus.Abbreviations AM amplitude modulation - DF Delta F - ELLL electrosensory lateral line lobe - EOD electric organ discharge - JAR jamming avoidance response - NE nucleus electrosensorius - PPN prepacemaker nucleus - PN pacemaker nucleus     

Developmental changes in cell proliferation in the auditory midbrain of the bullfrog, Rana catesbeiana

We examined patterns of cell proliferation in the auditory midbrain (torus semicircularis) of the bullfrog, Rana catesbeiana, over larval and early postmetamorphic development, by visualizing incorporation of 5-bromo-2'-deoxyuridine (BrdU) in cycling cells. At all developmental stages, BrdU-labeled cells were concentrated around the optic ventricle. BrdU-labeled cells also appeared within the torus semicircularis itself, in a stage-specific manner. The mitotic index, quantified as the percent of BrdU-positive cells outside the ventricular zone per total cells available for label, varied over larval development. Mitotic index was low in hatchling, early larval, and late larval stages, and increased significantly in deaf period, metamorphic climax, and froglet stages. Cell proliferation was higher in metamorphic climax than at other stages, suggesting increased cell proliferation in preparation for the transition from an aquatic to an amphibious existence. The change in mitotic index over development did not parallel the change in the total numbers of cells available for label. BrdU incorporation was additionally quantified by dot-blot assay, showing that BrdU is available for label up to 72 h postinjection. The pattern of change in cell proliferation in the torus semicircularis differs from that in the auditory medulla (dorsal medullary nucleus and superior olivary nucleus), suggesting that cell proliferation in these distinct auditory nuclei is mediated by different underlying mechanisms.     

Evaluation of Postspike Changes in Neuronal Excitability by Comparing Ordinary and Shuffled Autocorrelation Functions

Responses of single neurons to tonal signals amplitude-modulated by repeating segments of lowfrequency noise were studied in the dorsal (cochlear) medullary nucleus and midbrain auditory center (torus semicircularis) of the grass frog Rana temporaria. An autocorrelation function of the response to a total presentation and a shuffled autocorrelation function were derived. The latter was obtained by correlating the impulse response to each segment of the modulated signal with responses to all other segments with the exception of the initial one. After the necessary normalization, the function differed from the initial autocorrelation only in lacking postspike changes in excitability. A delay dependence of the ratio of the two functions directly demonstrated the time course of the postspike change in excitability of the studied cell. The majority of second-order neurons, which are in the dorsal nucleus of the medulla oblongata, were characterized only by brief intervals of absolute and relative refractoriness. However, cells with excitability that was markedly facilitated immediately after the refractory period were observed even in this nucleus. Neurons with a complex pattern of postspike changes in excitability were detected in the torus semicircularis. In these cells, a comparatively long postspike decrease in excitability was usually interrupted by intervals in which the neuron sensitivity was significantly higher than normal. The results demonstrate that spike generation has a marked effect on subsequent activity in brainstem auditory units. The effects may play an important role in the formation of the temporal pattern of neuronal responses to auditory signals.     

Directional and frequency response characteristics in the descending octaval nucleus of the toadfish (Opsanus tau)

This study is a continuation of a long-term investigation of the auditory circuit in the oyster toadfish, Opsanus tau. Input from the auditory periphery projects to the ipsilateral descending octaval nucleus (DON). Ipsilateral and contralateral DONs project to the auditory midbrain, where a previous study indicated that both frequency tuning and directional sharpening are present. To better understand the transformation of auditory information along the auditory pathway, we have examined over 400 units in the DON to characterize frequency and directional information encoded in the dorsolateral division of the nucleus. Background activity was primarily low (<10 spikes/s) or absent. The maximum coefficient of synchronization was equivalent to the periphery (R = 0.9) and substantially better than in the midbrain. The majority of DON units (79%) responded best to stimulus frequencies of 84-141 Hz and were broadly tuned. DON cells retain or enhance the directional character of their peripheral input (s); however, characteristic axes were distributed in all quadrants around the fish, providing further evidence that binaural computations may first occur in the DON of this species.     

Auditory responses in the torus semicircularis of the cane toad, Bufo marinus. I. Field potential studies

Field potentials have been recorded in the torus semicircularis of the toad, Bufo marinus, in response to brief tones presented in the free field. The amplitude of the potentials varied with the frequency of the stimulus and location of the electrode along the rostro-caudal axis of the torus. All frequencies in the auditory range evoked largest potentials when the stimulus was located in the contralateral auditory field. Potentials evoked by low to mid frequencies were largest when the stimulus was located near the line orthogonal to the long axis of the animal. For progressively higher frequencies, the optimal stimulus position was progressively more anterior in the contralateral field. In animals in which one eighth nerve had been sectioned, field potentials evoked by tones of low to mid frequency were less sensitive to changes in stimulus direction than in normal animals. However, the directional sensitivity of field potentials evoked by mid to high frequencies was similar in monaural and normal animals. These observations suggest that binaural neural integration is important in determining the directional sensitivity of field potentials in the torus evoked by low to mid frequencies but not for potentials evoked by mid to high frequencies.     

鲇鱼(Parasilurus Asotus)中脑电感受单元方向选择

本文分析了鲇中脑半园丘(Torns semicircularis TS)侧核深层中的一种瞬时单元(phasic unit)对电场方向的选择,这种方向选择取决于感受野的位置和单元是内向的(inward)抑或是外向的(outward).     

Acoustic response properties of single neurons in the central posterior nucleus of the thalamus of the goldfish,Carassius auratus

Acoustic responses were recorded extracellularly from single neurons in the thalamic central posterior nucleus (CP). Spontaneous activity, best sensitivity, and sharpness of tuning (Q_10db) of CP neurons ranged from 0 to 36 spikes/s, -40 to 5 dB re: 1 dyne/cm², and 0.18 to 1.80, respectively. The distribution of characteristic frequency (CF) was nonuniform with a mode at 195 Hz. Temporal response patterns of CP neurons (N = 60) were categorized into three groups: phasic (25%), tonic chopper-like (22%), and tonic nonchopper-like (53%) on the basis of peri-stimulus time and inter-spike interval histograms. Most CP neurons (90%) did not phase-lock to tones, and none phase-locked strongly. The properties of CP neurons are similar to those of the midbrain torus semicircularis neurons in spontaneous rates, best sensitivities, nonuniform CF distributions, and in exhibiting level-independent best frequencies. Both CP and toral neurons show a diversity of response patterns resembling those found in the mammalian central auditory system. However, CP neurons have broader tuning and less phase-locking than toral neurons, suggesting different roles in auditory processing. While peripheral frequency analysis is enhanced at the midbrain level, the integration of frequency-selective channels in the thalamus may function in the processing of wideband spectra characteristic of natural sound sources.Abbreviations BF best frequency - BS best sensitivity - CF characteristic frequency - CP central posterior nucleus - ISIH inter-spike interval histogram - PSTH peri-stimulus-time histogram - RA response area     

The coding of signals in the electric communication of the gymnotiform fish Eigenmannia: From electroreceptors to neurons in the torus semicircularis of the midbrain

Summary In the context of aggression and courtship, Eigenmannia repeatedly interrupts its electric organ discharges (EODs) These interruptions (Fig. 1) contain low-frequency components as well as high-frequency transients and, therefore, stimulate ampullary and tuberous electroreceptors, respectively (Figs. 2, 3). Information provided by these two classes of receptors is relayed along separate pathways, via the electrosensory lateral line lobe (ELL) of the hindbrain, to the dorsal torus semicircularis (TSd) of the midbrain. Some neurons of the torus receive inputs from both types of receptors (Figs. 14, 15), and some respond predominantly to EOD interruptions while being rather insensitive to other forms of signal modulations (Figs. 12, 13). This high selectivity appears to result from convergence and gating of inputs from individually less selective neurons.Abbreviations CP central posterior thalamic nucleus - Df frequency difference between neighbor's EOD and fish's own - DPn dorsal posterior nucleus (thalamus) - EOD electric organ discharge - ELL electrosensory lateral line lobe - JAR jamming avoidance response - LMR lateral mesencephalic reticular formation - nE nucleus electrosensorius - nEb nucleus electrosensorius, beat-related area - nE nucleus electrosensorius, area causing rise of EOD frequency - nE nucleus electrosensorius, area causing fall of EOD frequency - nEar nucleus electrosensorius-acusticolateralis area - NPd nucleus praeeminentialis, pars dorsalis - PPn prepacemaker nucleus - PT pretectal nucleus - SE nucleus subelectrosensorius - TeO optic tectum - TSd dorsal (electrosensory) torus semicircularis - TSv ventral (mechano-sensory and auditory) torus semicircularis     

Gonadotropin-releasing hormone (GnRH) modulates auditory processing in the fish brain

Gonadotropin-releasing hormone 1 (GnRH1) neurons control reproductive activity, but GnRH2 and GnRH3 neurons have widespread projections and function as neuromodulators in the vertebrate brain. While these extra-hypothalamic GnRH forms function as olfactory and visual neuromodulators, their potential effect on processing of auditory information is unknown. To test the hypothesis that GnRH modulates the processing of auditory information in the brain, we used immunohistochemistry to determine seasonal variations in these neuropeptide systems, and in vivo single-neuron recordings to identify neuromodulation in the midbrain torus semicircularis of the soniferous damselfish Abudefduf abdominalis. Our results show abundant GnRH-immunoreactive (-ir) axons in auditory processing regions of the midbrain and hindbrain. The number of extra-hypothalamic GnRH somata and the density of GnRH-ir axons within the auditory torus semicircularis also varied across the year, suggesting seasonal changes in GnRH influence of auditory processing. Exogenous application of GnRH (sGnRH and cGnRHII) caused a primarily inhibitory effect on auditory-evoked single neuron responses in the torus semicircularis. In the majority of neurons, GnRH caused a long-lasting decrease in spike rate in response to both tone bursts and playbacks of complex natural sounds. GnRH also decreased response latency and increased auditory thresholds in a frequency and stimulus type-dependent manner. To our knowledge, these results show for the first time in any vertebrate that GnRH can influence context-specific auditory processing in vivo in the brain, and may function to modulate seasonal auditory-mediated social behaviors.