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Walter Metzner

Publications and source records attributed to Walter Metzner.

8 recordsLinked to original sources

A mechanism for vocal-respiratory coupling in the mammalian parabrachial nucleus.

Mammalian vocalizations require the precise coordination of separate laryngeal and respiratory motor pathways. Precisely how and where in the brain vocal motor patterns interact with respiratory rhythm control is unknown. The parabrachial nucleus (PB) is known to mediate key respiratory reflexes and is also considered a principle component of the mammalian vocal motor pathway, making it a likely site for vocal-respiratory interactions, yet a specific role for the PB in vocalizing has yet to be demonstrated. To investigate the role of the PB in vocal-respiratory coordination, we pharmacologically manipulated synaptic activity in the PB while spontaneously vocalizing horseshoe bats were provoked to emit either short, single syllable or long, multisyllabic vocal motor patterns. Iontophoresis of the GABAA agonist muscimol (MUS) into the lateral PB extended expiratory durations surrounding all vocalizations and increased mean call durations. Alternatively, application of the GABAA antagonist bicuculline methiodide (BIC) shortened expirations and call durations. In addition, BIC eliminated the occurrence of multisyllabic vocalizations. BIC caused a mild increase in quiet breathing rates, whereas MUS tended to slow quiet breathing. The results indicate that GABAA receptor-mediated inhibition in the lateral PB modulates the time course of respiratory phase switching during vocalizing, and is needed for proper coordination of calling and breathing in mammals. We hypothesize that vocal-respiratory rhythm entrainment is achieved at least in part via mechanisms similar to other forms of locomotor-respiratory coupling, namely somatosensory feedback influences on respiratory phase-switching in the lateral PB.

Animals↗

Vocal communication in adult greater horseshoe bats, Rhinolophus ferrumequinum.

Whereas echolocation in horseshoe bats is well studied, virtually nothing is known about characteristics and function of their communication calls. Therefore, the communication calls produced by a group of captive adult greater horseshoe bats were recorded during various social interactions in a free-flight facility. Analysis revealed that this species exhibited an amazingly rich repertoire of vocalizations varying in numerous spectro-temporal aspects. Calls were classified into 17 syllable types (ten simple syllables and seven composites). Syllables were combined into six types of simple phrases and four combination phrases. The majority of syllables had durations of more than 100 ms with multiple harmonics and fundamental frequencies usually above 20 kHz, although some of them were also audible to humans. Preliminary behavioral observations indicated that many calls were emitted during direct interaction with and in response to social calls from conspecifics without requiring physical contact. Some echolocation-like vocalizations also appeared to clearly serve a communication role. These results not only shed light upon a so far widely neglected aspect of horseshoe bat vocalizations, but also provide the basis for future studies on the neural control of the production of communicative vocalizations in contrast to the production of echolocation pulse sequences.

Age Factors↗

Spontaneous, generalized lipidosis in captive greater horseshoe bats (Rhinolophus ferrumequinum).

During a routine 6-month quarantine period, 3 of 34 greater horseshoe bats (Rhinolophus ferrumequinum) captured in mainland China and transported to the United States for use in echolocation studies were found dead with no prior history of illness. All animals were in good body condition at the time of death. At necropsy, a large amount of white fat was found within the subcutis, especially in the sacrolumbar region. The liver, kidneys, and heart were diffusely tan in color. Microscopic examination revealed that hepatocytes throughout the liver were filled with lipid, and in some areas, lipid granulomas were present. renal lesions included moderate amounts of lipid in the cortical tubular epithelium and large amounts of protein and lipid within Bowman's capsules in the glomeruli. In addition, one bat had large lipid vacuoles diffusely distributed throughout the myocardium. The exact pathologic mechanism inducing the hepatic, renal, and cardiac lipidosis is unknown. The horseshoe bats were captured during hibernation and immediately transported to the United States. It is possible that the large amount of fat stored coupled with changes in photoperiod, lack of exercise, and/or the stress of captivity might have contributed to altering the normal metabolic processes, leading to anorexia and consequently lipidosis in these animals.

Animals↗

Auditory-feedback control of temporal call patterns in echolocating horseshoe bats.

During flight, auditory feedback causes horseshoe bats to adjust the duration and repetition rate of their vocalizations in a context-dependent manner. As these bats approach a target, they make finely graded adjustments in call duration and interpulse interval (IPI), but their echolocation behavior is also characterized by abrupt transitions in overall temporal calling patterns. We investigated the relative contributions of two prominent acoustic cues, echo frequency and delay, toward the control of both graded and transitional changes in call duration and IPI. Echoes returning at frequencies above the emitted call frequency caused bats to switch from long single calls to pairs of short calls (doublets). Alternatively, increasing echo delay caused progressive increases in IPI but caused no accompanying changes in call duration. When frequency shifts were combined with changing echo delays, echo delay altered the IPIs occurring between doublets but not the IPI within a doublet. When the echo mimic was replaced by presentation of either an artificial constant-frequency (CF) stimulus or a frequency-modulated (FM) stimulus, each designed to mimic major components of the echo acoustic structure, we found that CF stimuli could trigger the switch to doublets, but changing CF delay had no influence on IPI, whereas the timing of an FM-sweep presentation had a strong effect on IPI. Because CF and FM sounds are known to be processed separately in the bat auditory system, the results indicate that at least two distinct neural feedback pathways may be used to control the temporal patterns of vocalization in echolocating horseshoe bats.

Animals↗

A neural basis for auditory feedback control of vocal pitch.

Hearing one's own voice is essential for the production of correct vocalization patterns in many birds and mammals, including humans. Bats, for instance, adjust temporal, spectral, and intensity parameters of their echolocation calls by precisely monitoring the characteristics of the returning echo signals. However, neuronal substrates and mechanisms for auditory feedback control of vocalizations are still mostly unknown in any vertebrate. We used echolocating horseshoe bats to investigate the role of the midbrain and hindbrain tegmentum for the control of call frequencies in response to changing auditory feedback. These bats accurately control the frequency of their echolocation calls through auditory feedback both when the bat is at rest [resting frequency (RF)] and when it is flying and compensating for changes in echo frequency caused by flight-induced Doppler shifts [Doppler shift compensation (DSC)]. We iontophoretically injected various GABAergic and glutamatergic transmitter agonists and antagonists into the brainstem tegmentum. We found that within the parabrachial nuclei and the immediately adjacent tegmentum, excitatory effects caused by application of the glutamate agonist AMPA or the GABA(A) antagonist bicuculline raised RF and the frequency of calls emitted during DSC. Bicuculline application routinely blocked DSC altogether. Alternately, inhibitory effects caused by application of either the GABA(A) agonist muscimol or the AMPA antagonist CNQX lowered call frequencies emitted at rest and during DSC. Such an audio-vocal feedback mechanism might share basic aspects with audio-vocal feedback controlling the pitch of vocalizations in other mammals, including the involuntary response to "pitch-shifted feedback" in humans.

6-Cyano-7-nitroquinoxaline-2,3-dione↗

Effects of echo intensity on Doppler-shift compensation behavior in horseshoe bats.

Echolocating horseshoe bats respond to flight-speed induced shifts in echo frequency by adjusting the frequency of subsequent calls. Under natural conditions, Doppler effects may force the frequency of a returning echo several kilohertz above the original emission frequency. By lowering subsequent call frequencies, the bat can return echo frequencies to within a narrow spectral bandwidth to which its highly specialized auditory system is most sensitive. While Doppler-shift compensation (DSC) behavior specifically refers to frequency compensation, other parameters of the returning echo, such as delay, duration, and interaural time and intensity differences have been shown to influence DSC performance. Understanding the nature of these influences has already led to a better appreciation of the neurophysiology of DSC. Here we provide a quantitative analysis of the effects of a prominent feature of the returning echo, its intensity, on DSC performance in horseshoe bats. Although DSC performance generally tolerates echo attenuation up to approximately 40 dB relative to the outgoing emission intensity, a systematic decline in DSC performance can be observed over this range. Generally, the effects of echo attenuation are characterized by a reduction in 1) the overall amount of compensation relative to the size of the shift in echo frequency and 2) the rate at which the bat responds to perceived echo shifts. These effects appear to be the consequence of a systematic shift in the range of echo frequencies capable of inducing DSC behavior. In particular, the reference frequency (the minimum shift in echo frequency that will elicit DSC behavior) appears to be highly sensitive to echo intensity. Every 10-dB reduction in echo intensity shifts the reference upward nearly 250 Hz. Our results indicate that, even at the highest intensity levels, relatively minor changes in echo intensity critically influence frequency compensation during normal DSC. We conclude with a discussion of how these results might impact echolocation behavior of horseshoe bats under natural and experimental conditions.

Adaptation, Physiological↗

Stimulus encoding and feature extraction by multiple sensory neurons.

Neighboring cells in topographical sensory maps may transmit similar information to the next higher level of processing. How information transmission by groups of nearby neurons compares with the performance of single cells is a very important question for understanding the functioning of the nervous system. To tackle this problem, we quantified stimulus-encoding and feature extraction performance by pairs of simultaneously recorded electrosensory pyramidal cells in the hindbrain of weakly electric fish. These cells constitute the output neurons of the first central nervous stage of electrosensory processing. Using random amplitude modulations (RAMs) of a mimic of the fish's own electric field within behaviorally relevant frequency bands, we found that pyramidal cells with overlapping receptive fields exhibit strong stimulus-induced correlations. To quantify the encoding of the RAM time course, we estimated the stimuli from simultaneously recorded spike trains and found significant improvements over single spike trains. The quality of stimulus reconstruction, however, was still inferior to the one measured for single primary sensory afferents. In an analysis of feature extraction, we found that spikes of pyramidal cell pairs coinciding within a time window of a few milliseconds performed significantly better at detecting upstrokes and downstrokes of the stimulus compared with isolated spikes and even spike bursts of single cells. Coincident spikes can thus be considered "distributed bursts." Our results suggest that stimulus encoding by primary sensory afferents is transformed into feature extraction at the next processing stage. There, stimulus-induced coincident activity can improve the extraction of behaviorally relevant features from the stimulus.

Action Potentials↗

Doppler-shift compensation behavior in horseshoe bats revisited: auditory feedback controls both a decrease and an increase in call frequency.

Among mammals, echolocation in bats illustrates the vital role of proper audio-vocal feedback control particularly well. Bats adjust the temporal, spectral and intensity parameters of their echolocation calls depending on the characteristics of the returning echo signal. The mechanism of audio-vocal integration in both mammals and birds is, however, still largely unknown. Here, we present behavioral evidence suggesting a novel audio-vocal control mechanism in echolocating horseshoe bats (Rhinolophus ferrumequinum). These bats compensate for even subtle frequency shifts in the echo caused by flight-induced Doppler effects by adjusting the frequency of their echolocation calls. Under natural conditions, when approaching background targets, the bats usually encounter only positive Doppler shifts. Hence, we commonly believed that, during this Doppler-shift compensation behavior, horseshoe bats use auditory feedback to compensate only for these increases in echo frequency (=positive shifts) by actively lowering their call frequency below the resting frequency (the call frequency emitted when not flying and not experiencing Doppler shifts). Re-investigation of the Doppler-shift compensation behavior, however, shows that decreasing echo frequencies (=negative shifts) are involved as well: auditory feedback from frequencies below the resting frequency, when presented at similar suprathreshold intensity levels as higher echo frequencies, cause the bat's call frequency to increase above the resting frequency. However, compensation for negative shifts is less complete than for positive shifts (22% versus 95%), probably because of biomechanical restrictions in the larynx of bats. Therefore, Doppler-shift compensation behavior involves a quite different neural substrate and audio-vocal control mechanism from those previously assumed. The behavioral results are no longer consistent with solely inhibitory feedback originating from frequencies above the resting frequency. Instead, we propose that auditory feedback follows an antagonistic push/pull principle, with inhibitory feedback lowering and excitatory feedback increasing call frequencies. While the behavioral significance of an active compensation for echo frequencies below RF remains unclear, these behavioral results are crucial for determining the neural implementation of audio-vocal feedback control in horseshoe bats and possibly in mammals in general.

Animals↗