Unusual mating behavior of the Malaysian treefrogs, Polypedates leucomystax.
Explore the source record for details and available documents.
Animals↗
Biomedical subjects
A S Feng
Publications and source records attributed to A S Feng.
Information processing in the auditory brainstem.
The past year has seen significant advances in our understanding of the structural (circuitry and chemistry of synaptic connections) and functional characteristics of the auditory brainstem. Some of the findings that shed light on the mechanisms underlying complex auditory information processing are highlighted.
Animals↗
Temporal coding in the frog auditory midbrain: the influence of duration and rise-fall time on the processing of complex amplitude-modulated stimuli.
1. Single-unit recordings were made in the auditory midbrain, the torus semicircularis (TS) of the northern leopard frog, to independently characterize the processing of different temporal attributes (signal duration, rise-fall time, and rate of amplitude modulation) of natural sounds and to investigate how these temporal variables interact to produce the observed responses to complex amplitude-modulated (AM) signals. Response functions, on the basis of mean spike count, were derived and categorized to describe the unit's temporal response characteristics to each of the variables. 2. To characterize the duration response functions, tone bursts of different durations (stimuli repeated at a constant repetition rate) at the unit's characteristic frequency (CF) and 10 dB above minimum threshold at CF (MT) were presented monaurally to the contralateral ear. The duration response function of a TS neuron was often related to the temporal discharge characteristics of the neuron. Increases in stimulus duration elicited an increase in spike counts (therefore, long-pass response function) from most neurons (74%) in the TS; 91% of these neurons showed tonic discharge patterns. Phasic-burst (PB) cells that were rapidly adapting showed long-pass duration response functions that were highly nonlinear, having peaks and notches embedded within the functions. On the other hand, one-third of phasic neurons tended to be insensitive to stimulus duration, giving similar spike counts in response to stimuli of greatly different durations (i.e., all pass). In the TS, some neurons (9%) only responded to a limited range of durations (i.e., band-duration pass), and still others showed a preference for shorter durations (9%; i.e., short pass); these were exhibited primarily by phasic and PB neurons. 3. To characterize the rise-fall time response functions, tone bursts having different rise-fall times were presented. The rise-fall time response functions of TS neurons had two distinct characteristics. The majority of tonic cells (91%), as well as some PB (38%) and phasic (29%) neurons, gave essentially invariant spike counts for all stimulus rise-fall times (i.e., all pass; 73% of neurons). Despite the relatively stable spike counts of neurons showing all-pass functions, the peristimulus time histograms (PSTHs) deriving from responses to slower rise-fall time stimuli exhibited a longer and somewhat more variable onset latency. About one-fourth (27%) of TS neurons, mostly phasic and PB neurons, showed higher spike counts for signals with rapid rise-fall times.(ABSTRACT TRUNCATED AT 400 WORDS)
Acoustic Stimulation↗
Differential innervation patterns of three divisions of frog auditory midbrain (torus semicircularis).
The connectivity pattern of the laminar, principal, and magnocellular nuclei of the frog torus semicircularis (TS) was investigated. A small amount of horseradish peroxidase was injected focally into individual divisions of the TS and anterograde and retrograde transport patterns were observed. Results of our tracing study showed that these divisions of the TS possessed distinct innervation patterns. The principal nucleus appeared to be the primary input port of the TS receiving extensive inputs from all caudal brainstem auditory nuclei bilaterally, but especially from the contralateral dorsal medullary nucleus and the ipsilateral superior olivary and lateral lemniscus nuclei. Descending projection to this nucleus was limited to that from the posterior thalamic nucleus. In contrast, the laminar nucleus, but even more markedly the magnocellular nucleus, received extensive descending inputs from numerous structures in the dorsal thalamus and less pronounced ascending afferents from caudal brainstem auditory nuclei. Similar to the afferent connection patterns, the efferent projections originating from these three toral divisions differed substantially. The principal nucleus gave restricted ascending projections, limited mainly to the caudal region of the posterior thalamic nucleus, a region important in processing spectral information of complex sounds, and the pretectal gray. Its descending projection was also somewhat restricted, being limited to the superior olivary and lateral lemniscus nuclei. The laminar nucleus and especially the magnocellular nucleus gave robust descending as well as ascending projections; these nuclei serve as the main output paths for the TS and provide the main routes by which auditory input reaches the central thalamic nucleus, a structure previously shown to be involved in temporal information processing.
Afferent Pathways↗
Processing of behaviorally relevant temporal parameters of acoustic stimuli by single neurons in the superior olivary nucleus of the leopard frog.
Response characteristics of 130 single neurons in the superior olivary nucleus of the northern leopard frog (Rana pipiens pipiens) were examined to determine their selectivity to various behaviorally relevant temporal parameters [rise-fall time, duration, and amplitude modulation (AM) rate] of acoustic signals. Response functions were constructed with respect to each of these variables. Neurons with different temporal firing patterns such as tonic, phasic or phasic-burst firing patterns, participated in time domain analysis in specific manners. Phasic neurons manifested preferences for signals with short rise-fall times, thus possessing low-pass response functions with respect to this stimulus parameter; conversely, tonic and phasic-burst units were non-selective and possessed all-pass response functions. A distinction between temporal firing patterns was also observed for duration coding. Whereas phasic units showed no change in the mean spike count with a change in stimulus duration (i.e., all-pass duration response functions), tonic and phasic-burst units gave higher mean spike counts with an increase in stimulus duration (i.e., primary-like high-pass response functions). Phasic units manifested greater response selectivity for AM rate than did tonic or phasic-burst units, and many phasic units were tuned to a narrow range of modulation rates (i.e., band-pass). The results suggest that SON neurons play an important role in the processing of complex acoustic patterns; they perform extensive computations on AM rate as well as other temporal parameters of complex sounds. Moreover, the response selectivities for rise-fall time, duration, and AM rate could often be shown to contribute to the differential responses to complex synthetic and natural sounds.
Acoustic Stimulation↗
Coding of temporal parameters of complex sounds by frog auditory nerve fibers.
1. Physiological recordings were made from single auditory fibers in the frog eighth nerve to determine quantitatively how the different behaviorally relevant temporal parameters (the signal rise-fall time, duration, and rate of amplitude modulation) of complex sounds are encoded in the auditory periphery. Individual temporal parameters were varied. Response functions (RFs) were constructed with respect to each of these parameters using each unit's best excitatory frequency (BF) as the carrier. 2. In response to a change in signal rise-fall time, auditory nerve fibers showed little change in the mean spike count or firing rate, i.e., all fibers displayed ALL-PASS RFrfts. But the transient components, particularly the early phasic component, of responses varied with rise-fall times; these components were more pronounced in the responses to stimuli with shorter rise-fall times. 3. In response to an increase in signal duration, auditory nerve fibers showed a corresponding increase in firing duration and thus in the mean spike count, giving rise to HIGH-PASS RFdurs. The shape of response curves differed among fibers; the difference appeared to be related to the fiber's temporal adaptation characteristic. When the firing rate was measured, all fibers displayed higher mean firing rates in response to shorter duration stimuli than they did to longer duration stimuli, thus giving rise to LOW-PASS response functions. 4. To determine the response transfer functions to modulation rate, pulsed (PAM) and sinusoidally (SAM) amplitude-modulated signals were used. These signals differed substantially in terms of their envelopes and how they varied with AM rate. Data were analyzed by 1) plotting spike counts against the AM rate to derive modulation transfer functions (MTFspks) and 2) plotting synchronization coefficients (SCs) against the AM rate to generate MTFscs. 5. In response to PAM stimuli, all fibers showed an increase in mean spike count with modulation frequency over the range examined, giving rise to HIGH-PASS MTFspks. 6. For SAM stimuli, the average energy and duty cycle are independent of AM rate. Most (79%) auditory fibers showed little selectivity for AM rate over a range of 5-400 Hz, giving rise to ALL-PASS MTFspks. The remaining auditory fibers displayed LOW-PASS MTFspks, i.e., there was a distinct decline in the mean spike count with increasing AM rate. 7. In response to PAM stimuli, most fibers showed good response synchrony at low AM rates but the SC declined with an increase in the AM rate (i.e., LOW-PASS MTFscs). The cut-off frequency was typically very high, averaging 90 pulses/s.(ABSTRACT TRUNCATED AT 400 WORDS)
Acoustic Stimulation↗
Temporal processing in the dorsal medullary nucleus of the Northern leopard frog (Rana pipiens pipiens).
1. Single-unit responses to different temporal acoustic parameters were characterized in the dorsal medullary nucleus (DMN) of the Northern leopard frog, Rana pipiens pipiens. Our goal was to provide both a quantitative and a qualitative assessment of the neural representation of behaviorally relevant temporal acoustic patterns in the frog's DMN. 2. Acoustic stimuli included tone bursts having different durations, rise times, or rates of amplitude modulation (AM). Several metrics were used to compute temporal response functions for each of these, including mean spike count, average firing rate, and/or peak firing rate. Synchronization coefficients were also used to characterize responses to stimuli presented at different AM rates. 3. On the basis of mean spike count, the temporal response functions of DMN neurons with respect to signal rise time could be characterized as 1) all-pass, in which the mean spike count was largely independent of rise time, or 2) fast-pass, in which the mean spike count decreased with increasing rise time. Fast-pass response functions were of two types, those that decayed rapidly and those that decayed gradually from their peak values. 4. The minimum threshold varied with signal rise time for cells showing fast-pass but not all-pass response functions. Minimum response thresholds for fast-pass neurons were typically higher with slower signal rise time. 5. The filtering characteristics of cells displaying fast-pass rise time response functions were dependent on signal level, becoming all-pass when signal levels exceeded 30-40 dB above the minimum threshold. 6. Approximately 44% of DMN neurons exhibiting fast-pass response functions for signal rise time showed all-pass filtering characteristics when broadband noise rather than best frequency tones were used, thereby signifying an influence of signal spectrum on the pass-band characteristics of these cells. 7. All DMN neurons, regardless of discharge pattern, showed maximal instantaneous firing rates to signals having short (less than 25 ms) rise times. Response functions based on instantaneous firing rate were, therefore, fast-pass in nature. These responses were independent of signal level and spectrum. 8. There was an ordinal relationship between signal duration and the duration of tonic but not phasic unit discharges. This relationship was not intensity dependent. 9. On the basis of mean spike count, the temporal response functions of DMN neurons with respect to signal duration were characterized as 1) all-pass, in which the mean spike count was largely independent of signal duration, or 2) long-pass, in which the mean spike count increased with increasing signal duration.(ABSTRACT TRUNCATED AT 400 WORDS)
Acoustic Stimulation↗
Functional organization of ascending and descending connections of the cochlear nucleus of horseshoe bats.
The ascending projections of the cochlear nucleus (CN) and the sources of descending inputs to the CN were investigated in horseshoe bats (Rhinolophus rouxi) by tracing the anterograde and retrograde transport of horseradish peroxidase (HRP or WGA-HRP) injected into the CN. The tracer was iontophoretically deposited into physiologically characterized regions of the cochlear nucleus (Feng and Vater, '85). We report the course and termination of pathways arising from the anteroventral (AVCN), posteroventral (PVCN), and dorsal (DCN) cochlear nucleus. The projection fields within the auditory brainstem centers (superior olivary complex [SOC]; lateral lemniscus complex [LLC]; and inferior colliculus [IC]) and their tonotopic organization according to the frequency representations at the injection sites are described. While the projection pattern is generally in accordance with other mammals, several species-characteristic features are noted: i) the lateral superior olive (LSO) receives tonotopically organized input from both the AVCN and PVCN; ii) the CN-projections to medial nuclear groups of the SOC located between the LSO and the medial nucleus of the trapezoid body do not support previously suggested homologies; iii) the ventral nucleus of the LLC can be subdivided into two divisions with distinct input patterns from the AVCN and PVCN, respectively.
Animals↗
Neural basis of sound pattern recognition in anurans.
Explore the source record for details and available documents.
Animals↗
Classification of the temporal discharge patterns of single auditory neurons in the dorsal medullary nucleus of the northern leopard frog.
1. The dorsal medullary nucleus (DMN) of frogs is the presumed homolog of the mammalian cochlear nucleus (CN). Like the CN, the DMN is the sole target of centrally projecting primary auditory-nerve fibers and the first central auditory-processing center. To study the transformation of acoustic information in the DMN, we have utilized relatively simple stimuli--tone bursts--to detail the temporal discharge patterns of DMN neurons that can be compared with those shown by auditory-nerve fibers. 2. Based on the shape of poststimulus time (PSTH) and interspike interval (ISIH) histograms, we observed six distinctive discharge patterns to tone bursts presented at the best excitatory frequency (BEF), 10 dB above threshold. Four of these (primary-like type 1-4) resembled discharge patterns seen at the level of the auditory nerve, whereas two (phasic and phasic burst) were only observed in the DMN. 3. At stimulus levels of 20-30 dB above BEF threshold several phasic neurons became tonic responders, whereas several primary-like type-2 cells gave "pauser" discharges. The response patterns of the remaining cells were intensity independent. 4. We further showed that many of the single-unit discharge patterns were related to other neuronal response properties; specifically, spontaneous firing rate, intensity-rate functions, threshold, latency, BEF, and sharpness of tuning (Q10). 5. The implications of our findings are discussed with respect to 1) the transformation of acoustic information as it is passed from the auditory nerve to the DMN, and 2) the functional organization of the DMN.
Acoustic Stimulation↗
Influence of envelope rise time on neural responses in the auditory system of anurans.
The influence of envelope rise time on neural responses was investigated in the central auditory pathway of frogs. Single unit and evoked potential recordings were made from the dorsal medullary nucleus (DMN) and thalamus, respectively. It was found that phasic neurons (15% of the population) in the DMN responded preferentially to stimuli with rapid (less than 25 ms) rise times. Acoustically evoked potentials (AEPs) recorded from the thalamus, specifically the Nucleus of Bellonci, also showed more pronounced responses to stimuli with rapid, rather than slower, envelope rise times. Interestingly, the leopard frog mating call, which has a rapid onset, elicited strong neuronal discharge both within the DMN and thalamus. In contrast, the mating call of bullfrogs, a species sympatric with leopard frogs, has a characteristically slow (greater than 100 ms) envelope rise time and elicited little, if any, response. These findings indicate the presence of neural specializations within the frog's auditory pathway for the optimal detection of the conspecific mating call. The relevance of these findings for stimulus coding in the auditory pathway of other vertebrates is discussed.
Animals↗
Evidence for parallel processing in the frog's auditory thalamus.
We have conducted anatomical and physiological experiments to investigate the functional organization of the dorsal thalamus in the northern leopard frog (Rana pipiens pipiens). Our studies provide evidence for parallel auditory processing at this level of the frog's brain. Acoustically evoked potentials were recorded from the posterior and central thalamic nuclei and several differences in sound-evoked activity were noted between them: the amplitude of acoustically evoked potentials (AEPs), in response to a standard search stimulus, was always greater in the central, as opposed to the posterior, nucleus; the posterior, but not central, nucleus exhibited the phenomenon of nonlinear summation when 350-Hz and 1,700-Hz tones were presented simultaneously rather than individually; and the central, but not posterior, nucleus showed selectivity for the repetition rate of pulsed sound signals. The posterior and central thalamic nuclei also possessed distinct innervation patterns as revealed by the HRP transport patterns arising from these structures. The central nucleus was reciprocally connected with the major auditory relay stations along the frog's central auditory pathway including the superior olive, nucleus of the lateral lemniscus, and the torus semicircularis. Major projections to the lateral thalamic nucleus, ventral hypothalamus, and the telencephalic striatal complex were also observed. The posterior nucleus, on the other hand, established reciprocal connections primarily with the medial reticular nucleus, ventral midbrain tegmentum, and structures constituting of the ventral thalamic nuclei, particularly the nucleus of Bellonci. Thus, time and frequency cues contained within the species mating call, and conveying information concerning species identity, appear to be processed independently within the frog's thalamus with separate neural channels for each.
Animals↗
Autoradiographic study of descending pathways from the pontine reticular formation and the mesencephalic trigeminal nucleus in the rat.
Descending projections were studied in autoradiographically prepared material after injections of tritiated leucine in the pontine tegmentum of rats. Injections involving the medial pontine reticular formation resulted not only in labeling commissural fibers, the medial reticulospinal tract, and the dorsal cap of the inferior olive, but also, in two cases, in labeling a cerebellar projection that originated from a region near the midline and clearly dorsal to the nucleus reticularis tegmenti pontis. The labeled fibers passed ventral in the midline to the pontine gray, then laterally through the gray and into the middle cerebellar peduncle to terminate as mossy fibers primarily in the flocculus, lobulus simplex, and Crus I of the ansiform lobule. Injections involving the mesencephalic nucleus of the trigeminal nerve (Vmes), resulted in labeling of Probst's tract, which descends in the dorsolateral reticular formation. Probst's tract gave off extensive terminal branches to the lateral medullary reticular formation and weaker projections to restricted portions of the descending trigeminal nucleus, the solitary nucleus, and the hypoglossal nucleus. In one case, fibers could be traced into the dorsal horn of the upper cervical cord.
Abducens Nerve↗
Afferent and efferent innervation patterns of the cochlear nucleus (dorsal medullary nucleus) of the leopard frog.
The afferent and efferent innervation patterns of the frog dorsal medullary nucleus (DMN; anuran homolog of the cochlear nucleus) were examined by studying the anterograde and retrograde transport patterns of horseradish peroxidase injected focally into the nucleus. It was found that this structure projected bilaterally to the superior olivary nuclei (SON) and dorsal midbrain tegmental nuclei, and contralaterally to the opposite DMN, the lateral lemniscus nucleus (LLN) and the torus semicircularis (TS). The termination sites in the TS were restricted to the laminar and principal nuclei. The DMN in turn received projections from these structures with the exception of the TS and dorsal tegmental nuclei. The projection to the ipsilateral LLN and TS was not pronounced. In addition to the above findings, the ascending projection to the DMN, SON and TS, as well as the centrifugal projection from the SON, were found to be organized tonotopically.
Animals↗
Neural analysis of temporally patterned sounds in the frog's thalamus: processing of pulse duration and pulse repetition rate.
We conducted a single unit study of auditory neurons in the central thalamic nucleus of the northern leopard frog. These cells, unlike those of the posterior thalamic nucleus (J. Comp. Physiol., 150 (1983) 333-344), were broadly tuned and sensitive to one or more temporal features (pulse duration and repetition rate) of complex sound signals. When compared with previous studies, these results suggest that temporal and spectral information may be processed separately by distinct populations of neurons within the frog's thalamus.
Acoustic Stimulation↗
Afferent and efferent innervation patterns of the superior olivary nucleus of the leopard frog.
An HRP study of the frog's superior olivary nucleus (SON) revealed that it (1) receives reciprocal tonotopic projections from dorsal medullary nuclei and principal nuclei of the torus semicircularis (TS) bilaterally; (2) receives bilateral projections from caudalis nuclei and brainstem reticular nuclei, and unilateral projections from the ipsilateral ventral tegmental nuclei and the contralateral SON; (3) is reciprocally connected with the ipsilateral laminar and magnocellular nuclei of the TS, dorsal tegmental nuclei and the posterior thalamic nucleus; (4) projects directly to the ipsilateral central thalamic nucleus.
Animals↗
Functional organization of the cochlear nucleus of rufous horseshoe bats (Rhinolophus rouxi): frequencies and internal connections are arranged in slabs.
The functional organization of the cochlear nucleus (CN) was studied with physiological recording and anatomical tracing techniques. Recordings were made from single CN neurons to examine their temporal firing patterns to tone burst stimuli and their frequency tuning characteristics. Recording loci of individual neurons were carefully monitored in order to understand how the functional properties of a cell relate to its location within the CN. We found that tonal frequencies were systematically represented in each of the three CN divisions (anteroventral, AVCN; posteroventral, PVCN; dorsal, DCN). Eight temporal response patterns were observed in CN neurons when stimulated at units' best excitatory frequencies (BF). With a few exceptions, neurons in each CN division could generate all eight firing patterns with different distributions for the three division. A focal injection of horseradish peroxidase (HRP), at the end of the physiological study, to a group of neurons possessing a similar BF in one CN division resulted in anterograde labeling of nerve terminals in the other two divisions at precisely the areas where the same frequency band was processed in these divisions. Labeled terminals in each division were closely congregated in the form of a thin slab. The slab orientation was division specific whereas its location was frequency specific, which could be predicted on the basis of physiological data. HRP injections into the DCN also resulted in retrograde labeling of somata in the AVCN and PVCN. On the other hand, only DCN neurons were retrogradely labeled when HRP was injected into the AVCN or the PVCN. These data showed how the three CN divisions are internally connected. Furthermore, retrogradely labeled cells occupied the same slabs where we found anterogradely labeled nerve terminals. Additionally, in a group of bats, HRP was injected into various functionally (i.e., BF) identified regions of the central nucleus of the inferior coliculus (IC) to clarify the type and location of CN projecting neurons. Retrogradely labeled cells in individual CN divisions likewise were arranged in slabs whose locations in the CN nuclei depended on the BFs of neurons at the injection site in the IC. These results show that slabs represent units of functional organization (i.e., tonal frequency, local connection and central projection) in the CN.
Animals↗
An HRP-study of the frequency-place map of the horseshoe bat cochlea: morphological correlates of the sharp tuning to a narrow frequency band.
The frequency-place map of the horseshoe bat cochlea was studied with the horseradish peroxidase (HRP) technique involving focal injections into various, physiologically defined regions of cochlear nucleus (CN). The locations of labeled spiral ganglion cells and their termination sites on inner hair cells of the organ of Corti from injections into CN-regions responsive to different frequencies were analyzed in three dimensional reconstructions of the cochlea. Horseshoe bats from different geographical populations were investigated. They emit orientation cells with constant frequency (CF) components around 77 kHz (Rhinolophus rouxi from Ceylon) and 84 kHz (Rhinolophus rouxi from India) and their auditory systems are sharply tuned to the respective CF-components. The HRP-map shows that in both populations: the frequency range around the CF-component of the echolocation signal is processed in the second half-turn of the cochlea, where basilar membrane (BM) is not thickened, secondary spiral lamina (LSS) is still present and innervation density is maximal; frequencies more than 5 kHz above the CF-component are processed in the first half-turn, where the thickened BM is accompanied by LSS and innervation density is low; frequencies below the spectral content of the orientation call are represented in apical turns showing no morphological specializations. The data demonstrate that the cochlea of horseshoe bats is normalized to the frequency of the individual specific CF-component of the echolocation call. The HRP-map can account for the overrepresentation of neurons sharply tuned to the CF-signal found in the central auditory system. A comparison of the HRP-map with a map derived with the 'swollen nuclei technique' following loud sound exposure (Bruns 1976b) reveals that the latter is shifted towards cochlear base by about 4 mm. This discrepancy warrants a new interpretation of the functional role of specialized morphological structures of the cochlea within the mechanisms giving rise to the exceptionally high frequency selectivity of the auditory system.
Animals↗