Stimulus Rise Time Research Articles

Vestibular Testing-New Physiological Results for the Optimization of Clinical VEMP Stimuli.

Both auditory and vestibular primary afferent neurons can be activated by sound and vibration. This review relates the differences between them to the different receptor/synaptic mechanisms of the two systems, as shown by indicators of peripheral function-cochlear and vestibular compound action potentials (cCAPs and vCAPs)-to click stimulation as recorded in animal studies. Sound- and vibration-sensitive type 1 receptors at the striola of the utricular macula are enveloped by the unique calyx afferent ending, which has three modes of synaptic transmission. Glutamate is the transmitter for both cochlear and vestibular primary afferents; however, blocking glutamate transmission has very little effect on vCAPs but greatly reduces cCAPs. We suggest that the ultrafast non-quantal synaptic mechanism called resistive coupling is the cause of the short latency vestibular afferent responses and related results-failure of transmitter blockade, masking, and temporal precision. This "ultrafast" non-quantal transmission is effectively electrical coupling that is dependent on the membrane potentials of the calyx and the type 1 receptor. The major clinical implication is that decreasing stimulus rise time increases vCAP response, corresponding to the increased VEMP response in human subjects. Short rise times are optimal in human clinical VEMP testing, whereas long rise times are mandatory for audiometric threshold testing.

Electrets: A Remedy for Partial Discharge Caused by Power Electronics Switching

In this article, a novel approach that mitigates partial discharge (PD) by neutralizing local electric fields with the incorporation of electrets is presented. The surface charge density requirement of electret films that enable electric field neutralization is derived and numerical analysis is performed to validate them. The fabrication process of electrets is introduced, and we experimentally demonstrate the electret films reducing PD caused by the high <italic xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink">dv/dt</i> of power electronics switching voltage stimuli for the first time. The PD measurements of an uncharged polyvinylidene fluoride (PVDF) film and a charged PVDF (electret) film are compared as a function of the rise time of square voltage stimuli. In addition, we compare the PD inception voltage of the two films while maintaining <italic xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink">dv/dt</i> at a constant value to regulate the impact of capacitive current pulses falsely being detected as PD. The results show great promise in the electret-based approach and suggest that further development of the technology may be able to completely solve PD that has been both chronic and emerging issues in the power industry.

The startle reflex in echolocating odontocetes: basic physiology and practical implications.

ABSTRACTThe acoustic startle reflex is an oligo-synaptic reflex arc elicited by rapid-onset sounds. Odontocetes evolved a range of specific auditory adaptations to aquatic hearing and echolocation, e.g. the ability to downregulate their auditory sensitivity when emitting clicks. However, it remains unclear whether these adaptations also led to changes of the startle reflex. We investigated reactions to startling sounds in two bottlenose dolphins (Tursiops truncatus) and one false killer whale (Pseudorca crassidens). Animals were exposed to 50 ms, 1/3 octave band noise pulses of varying levels at frequencies of 1, 10, 25 and 32 kHz while positioned in a hoop station. Startle responses were quantified by measuring rapid muscle contractions using a three-dimensional accelerometer attached to the dolphin. Startle magnitude increased exponentially with increasing received levels. Startle thresholds were frequency dependent and ranged from 131 dB at 32 kHz to 153 dB at 1 kHz (re. 1 µPa). Startle thresholds only exceeded masked auditory AEP thresholds of the animals by 47 dB but were ∼82 dB above published behavioural audiograms for these species. We also tested the effect of stimulus rise time on startle magnitude using a broadband noise pulse. Startle responses decreased with increasing rise times from 2 to 100 ms. Models suggested that rise times of 141–220 ms were necessary to completely mitigate startle responses. Our data showed that the startle reflex is conserved in odontocetes and follows similar principles as in terrestrial mammals. These principles should be considered when assessing and mitigating the effects of anthropogenic noise on marine mammals.

Effects of noise burst rise time and level on bottlenose dolphin (Tursiops truncatus) auditory brainstem responses.

Although the auditory brainstem response (ABR) is known to be an onset response, specific features of acoustic stimuli that affect the morphology of the ABR are not well understood. In this study, the effects of stimulus onset properties were investigated by measuring ABRs in seven bottlenose dolphins while systematically manipulating stimulus rise time and the amplitude of the sound pressure temporal envelope plateau. Stimuli consisted of spectrally pink (i.e., equal mean-square pressure in proportional frequency bands) noise bursts with linear rise (and fall) envelopes and frequency content from 10 to 160 kHz. Noise burst rise times varied from 32 μs to 4 ms and plateau sound pressure levels varied from 96 to 150 dB re 1 μPa. ABR peak latency was found to be a function of the rate of change of the sound pressure envelope, while ABR peak amplitude was a function of the envelope sound pressure at the end of a fixed integration window. The data support previous single-unit and nearfield response data from terrestrial mammals and a model where the rate of change of envelope sound pressure is integrated across a time window aligned with stimulus onset.

Heterogeneous Cytoskeletal Force Distribution Delineates the Onset Ca2+ Influx Under Fluid Shear Stress in Astrocytes.

Mechanical perturbations increase intracellular Ca2+ in cells, but the coupling of mechanical forces to the Ca2+ influx is not well understood. We used a microfluidic chamber driven with a high-speed pressure servo to generate defined fluid shear stress to cultured astrocytes, and simultaneously measured cytoskeletal forces using a force sensitive actinin optical sensor and intracellular Ca2+. Fluid shear generated non-uniform forces in actinin that critically depended on the stimulus rise time emphasizing the presence of viscoelasticity in the activating sequence. A short (ms) shear pulse with fast rise time (2 ms) produced an immediate increase in actinin tension at the upstream end of the cell with minimal changes at the downstream end. The onset of Ca2+ rise began at highly strained areas. In contrast to stimulus steps, slow ramp stimuli produced uniform forces throughout the cells and only a small Ca2+ response. The heterogeneity of force distribution is exaggerated in cells having fewer stress fibers and lower pre-tension in actinin. Disruption of cytoskeleton with cytochalasin-D (Cyt-D) eliminated force gradients, and in those cells Ca2+ elevation started from the soma. Thus, Ca2+ influx with a mechanical stimulus depends on local stress within the cell and that is time dependent due to viscoelastic mechanics.

Effect of Stimulus Rise-Time on the Ocular Vestibular-Evoked Myogenic Potential to Bone-Conducted Vibration

The negative potential at 10 msec (called n10) of the ocular vestibular-evoked myogenic potential (oVEMP) recorded beneath the eyes in response to bone-conducted vibration (BCV) delivered to the skull at the midline in the hairline (Fz) is a new indicator of otolithic, and in particular utricular, function. Our aim is to find the optimum combination of frequency and rise-time for BCV stimulation, to improve the sensitivity of oVEMP testing in the clinic. We tested 10 healthy subjects with 6 msec tone bursts of BCV at three stimulus frequencies, 250, 500, and 750 Hz, at rise-times ranging between 0 and 2 msec. The BCV was delivered at Fz. The n10 response was significantly larger at the shorter rise-times, being largest at zero rise-time. In addition, we examined the effect of stimulus frequency in these same subjects by delivering 6 msec tone bursts at zero rise-time at a range of frequencies from 50 to 1200 Hz. The main effect of rise-time was significant with shorter rise-times leading to larger n10 responses and the Rise-Time × Frequency interaction was significant so that at low frequencies (100 Hz) shorter rise-times had a modest effect on n10 whereas at high frequencies (750 Hz) shorter rise-times increased n10 amplitude substantially. The main effect of frequency was also significant: The n10 response tended to be larger at lower frequency, being largest between 250 and 500 Hz. In summary, in this sample of healthy subjects, the most effective stimulus for eliciting oVEMP n10 to BCV at Fz was found to be a tone burst with a rise-time of 0 msec at low stimulus frequency (250 or 500 Hz).

A postural reflex evoked by brief axial accelerations

Impulsive stimuli were used to evoke postural reflexes in healthy subjects (n = 10) and avestibular patients (n = 2). Electromyographic (EMG) activity was recorded with subjects standing erect, feet together with eyes closed and leaning forward to activate their leg muscles. EMG was recorded bilaterally from over the soleus muscles, rectified and averaged. The evoked response was biphasic and consisted of a short latency (SL) and a smaller medium latency (ML) response, at approximately 60 and 80 ms post-stimulus, respectively. Reflex amplitude was expressed as a percentage of the prestimulus level. Stimulus rise times of 10 and 14 ms evoked the largest responses. SL responses were largest when stimulating at the vertebra prominens (C7) or over the mid-thoracic region (C7: 56 ± 42%, mid-thoracic: 56 ± 51%). Positive (push) stimuli evoked larger responses than negative stimuli which evoked an initial inhibition (C7 positive: 56 ± 42%, C7 negative: -12 ± 5%). Similarly, increasing intensity also produced larger responses (standard intensity: 56 ± 42%; increased intensity: 80 ± 53%). There was no apparent EMG response in the sitting condition, despite activation of calf muscles, indicating the reflex was dependent on posture. Avestibular patients had well-formed responses similar to normals (patients: SL = 137.5 and 84.5%, ML = -11.6 and -16.2%) with mildly prolonged latencies. This study has described the basic properties of a reflex evoked by impulsive accelerations of the trunk that we propose is not primarily dependent upon vestibular afferents and is likely to have a role in normal postural stability. A truncal origin of the receptors mediating the response best fits our observations and is an alternative explanation for some previous reports about postural reflexes in response to limb displacements.

Latency of tone-burst-evoked auditory brain stem responses and otoacoustic emissions: Level, frequency, and rise-time effects

Simultaneous measurement of auditory brain stem response (ABR) and otoacoustic emission (OAE) delays may provide insights into effects of level, frequency, and stimulus rise-time on cochlear delay. Tone-burst-evoked ABRs and OAEs (TBOAEs) were measured simultaneously in normal-hearing human subjects. Stimuli included a wide range of frequencies (0.5-8 kHz), levels (20-90 dB SPL), and tone-burst rise times. ABR latencies have orderly dependence on these three parameters, similar to previously reported data by Gorga et al. [J. Speech Hear. Res. 31, 87-97 (1988)]. Level dependence of ABR and TBOAE latencies was similar across a wide range of stimulus conditions. At mid-frequencies, frequency dependence of ABR and TBOAE latencies were similar. The dependence of ABR latency on both rise time and level was significant; however, the interaction was not significant, suggesting independent effects. Comparison between ABR and TBOAE latencies reveals that the ratio of TBOAE latency to ABR forward latency (the level-dependent component of ABR total latency) is close to one below 1.5 kHz, but greater than two above 1.5 kHz. Despite the fact that the current experiment was designed to test compatibility with models of reverse-wave propagation, existing models do not completely explain the current data.

Vestibular-dependent spinal reflexes evoked by brief lateral accelerations of the heads of standing subjects

An impulsive acceleration stimulus, previously shown to activate vestibular afferents, was applied to the mastoid. Evoked EMG responses from the soleus muscles in healthy subjects (n = 10) and patients with bilateral vestibular dysfunction (n = 3) were recorded and compared with the effects of galvanic stimulation (GVS). Subjects were stimulated while having their eyes closed, head rotated, and while tonically activating their soleus muscles. Rectified EMG responses were recorded from the leg contralateral to the direction of head rotation. Responses were characterized by triphasic potentials that consisted of short-latency (SL), medium-latency (ML), and long-latency (LL) components beginning at (mean ± SD) 54.2 ± 4.8, 88.4 ± 4.7, and 121 ± 7.1 ms, respectively. Mean amplitudes for the optimum stimulus rise times were 9.05 ± 3.44% for the SL interval, 16.70 ± 4.41% for the ML interval, and 9.75 ± 4.89% for the LL interval compared with prestimulus values. Stimulus rise times of 14 and 20 ms evoked the largest ML amplitudes. GVS evoked biphasic responses (SL and ML) with similar latencies. Like GVS, the polarity of the initial interval was determined by the polarity of the stimulus and the evoked EMG response was attenuated when subjects were seated. There was no significant EMG response evoked when subjects were stimulated using 500-Hz vibration or in patients with bilateral vestibular dysfunction. Our study demonstrates that a brief lateral acceleration, likely to activate the utricle, can evoke spinal responses with properties similar to those previously shown for vestibular activation by GVS. The triphasic nature of the responses may allow the nervous system to respond differently to short compared with long-duration linear accelerations, consistent with their differing significance.

The sender-receiver matching hypothesis: support from the peripheral coding of acoustic features in songbirds

The sender-receiver matching hypothesis predicts that species-specific features of vocalizations will be reflected in species-specific auditory processing. This hypothesis has most often been invoked to explain correlations between vocal frequency ranges and the frequency range of auditory sensitivity; however, it could apply to other structural features, such as the rise time of stimuli. We explored this hypothesis in five songbird species that vary in the rise times and frequency range of their vocalizations. We recorded auditory evoked potentials (AEPs) to onset and sustained portions of stimuli that varied in both frequency and rise time. AEPs are gross potentials generated in the auditory nerve and brainstem and measured from the scalp. We found that species with shorter rise times in their vocalizations had greater amplitude and shorter latency onset AEPs than species with longer rise times. We also found that species with lower frequency and/or more tonal vocalizations had stronger sustained AEPs that follow the sound pressure changes in the stimulus (i.e. frequency following responses) than species with higher frequency and/or less tonal vocalizations. This is the first study in songbirds to show that acoustic features such as rise time and tonality are reflected in peripheral auditory processing.

Encoding of amplitude modulations by auditory neurons of the locust: influence of modulation frequency, rise time, and modulation depth

Using modulation transfer functions (MTF), we investigated how sound patterns are processed within the auditory pathway of grasshoppers. Spike rates of auditory receptors and primary-like local neurons did not depend on modulation frequencies while other local and ascending neurons had lowpass, bandpass or bandstop properties. Local neurons exhibited broader dynamic ranges of their rate MTF that extended to higher modulation frequencies than those of most ascending neurons. We found no indication that a filter bank for modulation frequencies may exist in grasshoppers as has been proposed for the auditory system of mammals. The filter properties of half of the neurons changed to an allpass type with a 50% reduction of modulation depths. Contrasting to reports for mammals, the sensitivity to small modulation depths was not enhanced at higher processing stages. In ascending neurons, a focus on the range of low modulation frequencies was visible in the temporal MTFs, which describe the temporal locking of spikes to the signal envelope. To investigate the influence of stimulus rise time, we used rectangularly modulated stimuli instead of sinusoidally modulated ones. Unexpectedly, steep stimulus onsets had only small influence on the shape of MTF curves of 70% of neurons in our sample.Electronic supplementary materialThe online version of this article (doi:10.1007/s00359-010-0587-4) contains supplementary material, which is available to authorized users.

Temporal Features of Spectral Integration in the Inferior Colliculus: Effects of Stimulus Duration and Rise Time

This report examines temporal features of facilitation and suppression that underlie spectrally integrative responses to complex vocal signals. Auditory responses were recorded from 160 neurons in the inferior colliculus (IC) of awake mustached bats. Sixty-two neurons showed combination-sensitive facilitation: responses to best frequency (BF) signals were facilitated by well-timed signals at least an octave lower in frequency, in the range 16-31 kHz. Temporal features and strength of facilitation were generally unaffected by changes in duration of facilitating signals from 4 to 31 ms. Changes in stimulus rise time from 0.5 to 5.0 ms had little effect on facilitatory strength. These results suggest that low frequency facilitating inputs to high BF neurons have phasic-on temporal patterns and are responsive to stimulus rise times over the tested range. We also recorded from 98 neurons showing low-frequency (11-32 kHz) suppression of higher BF responses. Effects of changing duration were related to the frequency of suppressive signals. Signals<23 kHz usually evoked suppression sustained throughout signal duration. This and other features of such suppression are consistent with a cochlear origin that results in masking of responses to higher, near-BF signal frequencies. Signals in the 23- to 30-kHz range-frequencies in the first sonar harmonic-generally evoked phasic suppression of BF responses. This may result from neural inhibitory interactions within and below IC. In many neurons, we observed two or more forms of the spectral interactions described here. Thus IC neurons display temporally and spectrally complex responses to sound that result from multiple spectral interactions at different levels of the ascending auditory pathway.

Differentiation between protective reflexes: Cardiac defense and startle

Rise time and duration are two parametric characteristics of the eliciting stimulus frequently used to differentiate among psychophysiological reflexes. The present research varied the duration (study 1) and rise time (study 2) of an intense acoustic stimulus to dissociate cardiac defense and cardiac startle using the eyeblink response as the external criterion of startle. In each study, 100 participants were presented with five white noise stimuli of 105 dB under one of five duration (50, 100, 250, 500, and 1000 ms) or rise time (0, 24, 48, 96, and 240 ms) conditions. Cardiac defense was affected by stimulus duration, present only in the 500- and 1000-ms conditions, but not by stimulus rise time, present in all rise time conditions. Rise time affected blink startle, but did not selectively alter the short latency accelerative component of the heart rate response, thus questioning whether it reflects startle.

Detection of high-frequency spectral notches as a function of level

High-frequency spectral notches are important cues for sound localization. Our ability to detect them must depend on their representation as auditory nerve (AN) rate profiles. Because of the low threshold and the narrow dynamic range of most AN fibers, these rate profiles deteriorate at high levels. The system may compensate by using onset rate profiles whose dynamic range is wider, or by using low-spontaneous-rate fibers, whose threshold is higher. To test these hypotheses, the threshold notch depth necessary to discriminate between a flat spectrum broadband noise and a similar noise with a spectral notch centered at 8 kHz was measured at levels from 32 to 100 dB SPL. The importance of the onset rate-profile representation of the notch was estimated by varying the stimulus duration and its rise time. For a large proportion of listeners, threshold notch depth varied nonmonotonically with level, increasing for levels up to 70-80 dB SPL and decreasing thereafter. The nonmonotonic aspect of the function was independent of notch bandwidth and stimulus duration. Thresholds were independent of stimulus rise time but increased for the shorter noise bursts. Results are discussed in terms of the ability of the AN to convey spectral notch information at different levels.

Age reduces response latency of mouse inferior colliculus neurons to AM sounds.

Age and stimulus rise time (RT) effects on response latency were investigated for inferior colliculus (IC) neurons in young-adult and old CBA mice. Single-unit responses were recorded to unmodulated and sinusoidal amplitude modulated (SAM) broadband noise carriers, presented at 35 to 80 dB SPL. Data from 63 young-adult and 76 old phasic units were analyzed to identify the time interval between stimulus onset and driven-response onset (latency). When controlling for stimulus sound level and AM frequency, significant age-related changes in latency were identified. Absolute latency decreased with age at all stimulus AM frequencies, significantly so for equivalent rise times (RT) < or = 12.5 ms. The linear correlation of latency with AM stimulus RT was significant for both young-adult and old units, and increased significantly with age. It is likely that both the decrease in absolute latency and the increase in latency/RT correlation with age are consistent with a reduction of inhibitory drive with age in the IC. These latency changes will result in age-related timing variations in brainstem responses to stimulus onsets, and therefore affect the encoding of complex sounds.

Differential effect of current rise time on short and medium latency vestibulospinal reflexes.

To investigate the effect of varying current rise time on galvanic-evoked short (SL) and medium (ML) latency vestibulospinal reflexes. We recorded the soleus EMG of standing subjects in response to 3 mA direct current transmastoid stimulation with a series of current ramps with rise times of 0-300 ms. Longer current rise times significantly delayed the onset of both SL (P<<0.001) and ML (P<<0.001) vestibulospinal responses, by approximately 20 and 39 ms, respectively. The SL response amplitude was reduced with increasing rise time (P<<0.001), whereas the ML response amplitude was relatively unaffected by stimulus rise time. With very slow rise times a prolonged ML response alone was evoked. Both SL and ML reflexes can be evoked by changes in vestibular activity produced by transmastoid galvanic stimulation with a ramp onset. We found a differential effect of current rise time on SL and ML vestibulospinal reflexes, suggesting different potential functional roles for the two reflexes. SL reflexes can participate in the response to abrupt disturbances only. ML reflexes are evoked by both fast and slow changes in vestibular discharge and may be particularly effective for slowly-changing disturbances.

Sensitivity of unanesthetized chinchilla auditory system to noise burst onset, and the effects of carboplatin

The gross near-field responses of the auditory nerve and inferior colliculus to noise burst stimuli were recorded through intracranially implanted electrodes in six unanesthetized chinchillas. Responses were studied as a function of stimulus plateau amplitude and rise time, both before and after a systemic dose of 75 mg/kg of carboplatin. Both recording sites showed sensitivity to stimulus level and rise time. Increases in stimulus level and decreases in stimulus rise time each produced increases in the response magnitude, and decreases in response latency. When the stimuli were re-specified as rate of pressure change at sound onset (Pa/s), the amplitude and latency of responses at each site were found to be a direct function of rate of sound pressure change. These data provide the first confirmation in unanesthetized animals of previous single unit observations in barbiturate-anesthetized cats. Carboplatin treatment resulted in a 20–80% loss of inner hair cells, a modest threshold elevation, and a 50–75% reduction in peak response amplitudes. The general patterns of sensitivity to stimulus level and rise time were not markedly affected by carboplatin, nor was the fashion in which response parameters (amplitude and latency) were ruled by rate of pressure change at sound onset.

Directional hearing is only weakly dependent on the rise time of acoustic stimuli

First-spike latency differences between left and right auditory-nerve fibers have been proposed as one of the physiological cues for sound localization. Since first-spike latency depends not only on stimulus intensity, but also on the steepness of the amplitude rise of a sound stimulus, differences in first-spike latency are not a simple function of interaural level differences but also a function of stimulus rise time. We therefore investigated whether rise time influences human directional hearing in a localization paradigm. Subjects tended to localize a 3-kHz tone pulse with a long (18-ms) rise time further to the side than one with a short (2-ms) rise time delivered from the same source. The small size of this effect and its large inter-individual variability, however, suggest that it is of minor importance for human directional hearing.

Short-term and long-term habituation of the acoustic startle response as a function of stimulus rise time in rats

In three experiments, short-term habituation (STH) and long-term habituation (LTH) of the acoustic startle response (ASR) were assessed as a function of the rise time of the startle-inducing stimulus. In Experiment 1, presentations of a 300-msec rise time stimulus (RTS) produced neither a startle response nor response decrement to a 0.4-msec RTS when switched to that stimulus within a session (STH). When tested over days (LTH) training with the 300-msec RTS produced significant response decrements to the 0.4-msec RTS. Stimulus-induced freezing and vocalization were inversely related to rise time, and these emotional responses habituated over days. Experiment 2 showed that STH was inversely related to the rise time of the training stimulus, not simply to stimulus change. Experiment 3 replicated the LTH results of Experiment 1. These data show that STH of ASR is inversely related to stimulus rise time, whereas LTH is relatively independent of rise time.

Is odor processing related to oral breathing?

This paper addresses two questions related to the inherent association between breathing and odor perception: Does central nervous processing of odors change when an artificial breathing technique (velopharyngeal closure) is introduced and secondly, does odor processing vary with the oral breathing phase (inhalation or exhalation)? Chemosensory event-related potentials (CSERP) were obtained from eight female subjects while they were smelling an odor mixture (citral, eugenol, linalool, menthol and isoamylacetate). Each subject was required to perform spontaneous mouth breathing (120 trials) as well as the velopharyngeal closure technique (120 trials). Simultaneously, a thermistor monitored the phase of the respiratory cycle. The results reveal that the central nervous correlates of odor processing change with the breathing technique but not with the oral breathing cycle. The findings that early stimulus processing is faster (N1 latency) and late stimulus processing more pronounced (P3 amplitudes) when the subjects are breathing spontaneously are discussed with regard to attentional effects. The reduction of the N1 amplitude during the spontaneous breathing condition may be caused by larger latency variations and longer stimulus rise-times. Furthermore, it is concluded that the oral breathing cycle is less important than the nasal breathing cycle for olfactory information transmission.