Increasing Stimulus Amplitude Research Articles

Student Competition (Technology Innovation) ID 1984861

Backgrounds/Objective Intraspinal microstimulation (ISMS) is a neuromodulation technique for restoring walking after spinal cord injury. The objective of this study was to fabricate a stretchable ISMS device suitable for pigs, a clinically-relevant animal model. Methods Polyimide-insulated microwires (50µm, Pt-Ir, 80%/20%) were used for fabrication of electrodes. Their tips were de-insulated (∼0.15mm2) and sharpened using nanosecond and femtosecond UV lasers. Microcoils were fabricated from 25μm microwires (Pt-Ir, 80%/20%) to add stretchability to the lead wires. Sixteen microelectrode-leads were connected to a custom, wirelessly controlled stimulator using Medtronic extension cables (Model 37081). The implants were tested in seven domestic pigs and current pulse trains were delivered to various rostro-caudal regions of the lumbar spinal cord (1s, 40Hz, 50µA-300µA) to activate locomotor-related muscle synergies. The kinematics and isometric joint forces of the evoked hindlimb responses were recorded. Results Graded joint movements were evoked with increasing stimulus amplitude. Changes in the hip, knee, and ankle joints angles evoked by ISMS at 300µA were 17.9±1˚, 28.1±1˚, and 21.6±2˚, respectively. Isometric joint forces evoked by ISMS at 300µA were 12.21±0.91N, 7.4±0.71N, and 1.7±0.15N for knee extension, hip flexion, and ankle flexion, respectively. Conclusion The movements evoked using the developed ISMS implant could generate full ranges of motion in the joints. The graded responses imply a near-physiological recruitment order of motoneurons, which is necessary for achieving long walking distances without muscle fatigue. The results show the capability of the developed ISMS device in generating movements in pigs, and the implants’ potential for future use in humans.

Neural substrates of perception in the vestibular thalamus during natural self-motion: A review.

Accumulating evidence across multiple sensory modalities suggests that the thalamus does not simply relay information from the periphery to the cortex. Here we review recent findings showing that vestibular neurons within the ventral posteriolateral area of the thalamus perform nonlinear transformations on their afferent input that determine our subjective awareness of motion. Specifically, these neurons provide a substrate for previous psychophysical observations that perceptual discrimination thresholds are much better than predictions from Weber's law. This is because neural discrimination thresholds, which are determined from both variability and sensitivity, initially increase but then saturate with increasing stimulus amplitude, thereby matching the previously observed dependency of perceptual self-motion discrimination thresholds. Moreover, neural response dynamics give rise to unambiguous and optimized encoding of natural but not artificial stimuli. Finally, vestibular thalamic neurons selectively encode passively applied motion when occurring concurrently with voluntary (i.e., active) movements. Taken together, these results show that the vestibular thalamus plays an essential role towards generating motion perception as well as shaping our vestibular sense of agency that is not simply inherited from afferent input.

Block Phenomena During Electric Micro-Stimulation of Pyramidal Cells and Retinal Ganglion Cells.

Electric micro-stimulation of the nervous system is a means to restore various body functions. The stimulus amplitude necessary to generate action potentials, the lower threshold (LT), is well characterized for many neuronal populations. However, electric overstimulation above an upper threshold (UT) prevents action potential generation and therefore hinders optimal neuro-rehabilitation. Previous studies demonstrated the impact of the UT in micro-stimulation of retinal ganglion cells (RGCs). The observed phenomenon is mostly explained by (i) reversed sodium ion flow in the soma membrane, and (ii) anodal surround block that hinders spike conduction in strongly hyperpolarized regions of the axon at high stimulus intensities. However, up to now, no detailed study of the nature of these phenomena has been presented, particularly for different cell types. Here, we present computational analyses of LT and UT for layer 5 pyramidal cells (PCs) as well as alpha RGCs. Model neurons were stimulated in close vicinity to the cell body and LTs and UTs as well as the ratio UT/LT were compared. Aside from a simple point source electrode and monophasic stimuli also realistic electrode and pulse configurations were examined. The analysis showed: (i) in RGCs, the soma contributed to action potential initiation and block for small electrode distances, whereas in PCs the soma played no role in LTs or UTs. (ii) In both cell types, action potential always initiated within the axon initial segment at LT. (iii) In contrast to a complete block of spike conductance at UT that occurred in RGCs, an incomplete block of spiking appeared in PC axon collaterals. (iv) PC axon collateral arrangement influenced UTs but had small impact on LTs. (v) Population responses of RGCs change from circular regions of activation to ring-shaped patterns for increasing stimulus amplitude. A better understanding of the stimulation window that can reliably activate target neurons will benefit the future development of neuroprostheses.

Feed-forward information and zero-lag synchronization in the sensory thalamocortical circuit are modulated during stimulus perception

The direction of functional information flow in the sensory thalamocortical circuit may play a role in stimulus perception, but, surprisingly, this process is poorly understood. We addressed this problem by evaluating a directional information measure between simultaneously recorded neurons from somatosensory thalamus (ventral posterolateral nucleus, VPL) and somatosensory cortex (S1) sharing the same cutaneous receptive field while monkeys judged the presence or absence of a tactile stimulus. During stimulus presence, feed-forward information (VPL → S1) increased as a function of the stimulus amplitude, while pure feed-back information (S1 → VPL) was unaffected. In parallel, zero-lag interaction emerged with increasing stimulus amplitude, reflecting externally driven thalamocortical synchronization during stimulus processing. Furthermore, VPL → S1 information decreased during error trials. Also, VPL → S1 and zero-lag interaction decreased when monkeys were not required to report the stimulus presence. These findings provide evidence that both the direction of information flow and the instant synchronization in the sensory thalamocortical circuit play a role in stimulus perception.

Segmental trunk and head dynamics during frontal plane tilt stimuli in healthy sitting adults

A more detailed understanding of trunk behavior during upright sitting is needed to create a foundation to address functional posture impairments. Therefore, we characterized the dynamics of the trunk and head during perturbed sitting. A three-link inverted pendulum model of head and trunk segments was used to analyze kinematics of eight healthy sitting adults. Magnetic sensors were placed at the head and two locations of the trunk (C7 and T7). Six surface tilt stimuli (two spontaneous sway tests [no surface stimulus; eyes open, EO/eyes closed, EC] and four tests with continuous pseudorandom surface tilts [2 peak-to peak amplitudes of 2° or 8°; EO/EC]) were applied in the frontal plane. We used frequency-response functions (FRFs) to analyze sway across ~0.045–3Hz and found systematic differences in sway dynamics across segments. Superior segments exhibited larger fluctuations in gain and phase values across frequencies. FRF gains in superior segments were attenuated compared to other segments only at low frequencies but were larger at the higher frequencies. We also tested the influence of stimulus amplitude and visual availability on FRFs. Across all segments, increasing stimulus amplitude and visual availability (EO) resulted in lower gains, however, these effects were most prominent in superior segments. These changes in gain were likely influenced by changes in sensory reliance across test conditions. In conclusion, these results provide a benchmark for future comparisons to segmental responses from individuals with impaired trunk control. We suggest that a frequency-based approach provides detail needed to characterize multi-segment dynamics related to sensorimotor control.

Stance width changes how sensory feedback is used for multisegmental balance control.

A multilink sensorimotor integration model of frontal plane balance control was developed to determine how stance width influences the use of sensory feedback in healthy adults. Data used to estimate model parameters came from seven human participants who stood on a continuously rotating surface with three different stimulus amplitudes, with eyes open and closed, and at four different stance widths. Dependent variables included lower body (LB) and upper body (UB) sway quantified by frequency-response functions. Results showed that stance width had a major influence on how parameters varied across stimulus amplitude and between visual conditions. Active mechanisms dominated LB control. At narrower stances, with increasing stimulus amplitude, subjects used sensory reweighting to shift reliance from proprioceptive cues to vestibular and/or visual cues that oriented the LB more toward upright. When vision was available, subjects reduced reliance on proprioception and increased reliance on vision. At wider stances, LB control did not exhibit sensory reweighting. In the UB system, both active and passive mechanisms contributed and were dependent on stance width. UB control changed across stimulus amplitude most in wide stance (opposite of the pattern found in LB control). The strong influence of stance width on sensory integration and neural feedback control implies that rehabilitative therapies for balance disorders can target different aspects of balance control by using different stance widths. Rehabilitative strategies designed to assess or modify sensory reweighting will be most effective with the use of narrower stances, whereas wider stances present greater challenges to UB control.

Embryonic assembly of auditory circuits: spiral ganglion and brainstem

During early development, peripheral sensory systems generate physiological activity prior to exposure to normal environmental stimuli. This activity is thought to facilitate maturation of these neurons and their connections, perhaps even promoting efficacy or modifying downstream circuitry. In the mammalian auditory system, initial connections form at embryonic ages, but the functional characteristics of these early neural connections have not been assayed. We investigated processes of embryonic auditory development using a whole-head slice preparation that preserved connectivity between peripheral and brainstem stations of the auditory pathway. Transgenic mice expressing fluorescent protein provided observation of spiral ganglion and cochlear nucleus neurons to facilitate targeted electrophysiological recording. Here we demonstrate an apparent peripheral-to-central order for circuit maturation. Spiral ganglion cells acquire action potential-generating capacity at embryonic day 14 (E14), the earliest age tested, and action potential waveforms begin to mature in advance of comparable states for neurons of the ventral cochlear nucleus (VCN) and medial nucleus of the trapezoid body (MNTB). In accordance, auditory nerve synapses in the VCN are functional at E15, prior to VCN connectivity with the MNTB, which occurs at least 1 day later. Spiral ganglion neurons exhibit spontaneous activity at least by E14 and are able to drive third-order auditory brainstem neurons by E17. This activity precedes cochlear-generated wave activity by 4 days and ear canal opening by at least 2 weeks. Together, these findings reveal a previously unknown initial developmental phase for auditory maturation, and further implicate the spiral ganglion as a potential controlling centre in this process.

Influence of Stance Width on Frontal Plane Postural Dynamics and Coordination in Human Balance Control

The influence of stance width on frontal plane postural dynamics and coordination in human bipedal stance was studied. We tested the hypothesis that when subjects adopt a narrow stance width, they will rely heavily on nonlinear control strategies and coordinated counter-phase upper and lower body motion to limit center-of-mass (CoM) deviations from upright; as stance increases, the use of these strategies will diminish. Freestanding frontal plane body sway was evoked through continuous pseudorandom rotations of the support surface on which subjects stood with various stimulus amplitudes. Subjects were either eyes open (EO) or closed (EC) and adopted various stance widths. Upper body, lower body, and CoM kinematics were summarized using root-mean-square and peak-to-peak measures, and dynamic behavior was characterized using frequency-response and impulse-response functions. In narrow stance, CoM frequency-response function gains were reduced with increasing stimulus amplitude and in EO compared with EC; in wide stance, gain reductions were much less pronounced. Results show that the narrow stance postural system is nonlinear across stimulus amplitude in both EO and EC conditions, whereas the wide stance postural system is more linear. The nonlinearity in narrow stance is likely caused by an amplitude-dependent sensory reweighting mechanism. Finally, lower body and upper body sway were approximately in-phase at low frequencies (<1 Hz) and out-of-phase at high frequencies (>1 Hz) across all stance widths, and results were therefore inconsistent with the hypothesis that subjects made greater use of coordinated counter-phase upper and lower body motion in narrow compared with wide stance conditions.

Spike-Firing Resonance in Hypoglossal Motoneurons

During an inspiration the output of hypoglossal (XII) motoneurons (HMs) in vitro is characterized by synchronous oscillatory firing in the 20- to 40-Hz range. To maintain synchronicity it is important that the cells fire with high reliability and precision. It is not known whether the intrinsic properties of HMs are tuned to maintain synchronicity when stimulated with time-varying inputs. We intracellularly recorded from HMs in an in vitro brain stem slice preparation from juvenile mice. Cells were held at or near spike threshold and were stimulated with steady or swept sine-wave current functions (10-s duration; 0- to 40-Hz range). Peristimulus time histograms were constructed from spike times based on threshold crossings. Synaptic transmission was suppressed by including blockers of GABAergic, glycinergic, and glutamatergic neurotransmission in the bath solution. Cells responded to sine-wave stimulation with bursts of action potentials at low (<3- to 5-Hz) sine-wave frequency, whereas they phase-locked 1:1 to the stimulus at intermediate frequencies (3-25 Hz). Beyond the 1:1 frequency range cells were able to phase-lock to subharmonics (1:2, 1:3, or 1:4) of the input frequency. The 1:1 phase-locking range increased with increasing stimulus amplitude and membrane depolarization. Reliability and spike-timing precision were highest when the cells phase-locked 1:1 to the stimulus. Our findings suggest that the coding of time-varying inspiratory synaptic inputs by individual HMs is most reliable and precise at frequencies that are generally lower than the frequency of the synchronous inspiratory oscillatory activity recorded from the XII nerve.

Multimodal gain control at the hippocampal Schaffer collateral-CA1 synapse

Information processing at central nervous system synapses is shaped by long-lasting modifications, such as long-term potentiation and short-lived and putatively synapse-specific modifications by various forms of short-term plasticity, such as facilitation, potentiation, and depression. Using an extracellular paired-pulse facilitation (PPF) protocol at the Schaffer collateral-CA1 (SC) connection in acute hippocampal slices in mice, we extend previous reports of optimal signal gain at intermediate interpulse intervals obtained at single SC synapses to the network level. Moreover, maximum signal gain changed when the input intensity was altered. We found further that facilitation decreased with increasing stimulus amplitude and duration in an exact exponential fashion when varied at a fixed interpulse interval. Variation of these intensity parameters accounted for significant changes in PPF adding a spatial dimension to time-based synaptic filter characteristics. Thus, this synapse functions as an amplitude window discriminator with a low-level aperture in combination with a band-pass frequency filter. By providing mathematical functions for the characteristic presynaptic parameters frequency, stimulus amplitude, and pulse duration at the network level our results lay ground for future studies on pharmacologically, genetically, or otherwise altered animal models.

Cerebral activation using a MR-compatible piezoelectric actuator with adjustable vibration frequencies and in vivo wave propagation control

Functional magnetic resonance imaging (fMRI) studies are increasingly used in patients with brain tumors near the sensory motor cortex for planning of therapy. Passive stimuli can be helpful for reproducible results. The purpose of our study was to investigate frequency and amplitude dependencies of cerebral activation patterns using a vibratory stimulus that involves sensory and motor function and allows exact adjustment of vibratory frequencies and direct control of penetration depth into the tissue. Fifteen volunteers were studied with fMRI during vibratory stimulation of the right biceps muscle utilizing a block design (frequencies: 150 and 300 Hz, amplitudes: 400, 600, and 800 μm). In addition, visualization of the wave propagation into the biceps tissue itself was performed with a modified phase contrast sequence. A specially developed MR-compatible mechanical oscillator was used to apply the vibrotactile sensations. fMRI revealed activation of the left primary somatosensory cortex during application of both vibratory frequencies. Additionally, activity of the primary and supplementary motor cortex was revealed using 150-Hz stimuli, while only minimal at 300 Hz. The activity strength correlated with increasing stimulus amplitudes and the visualized penetration depth. Activation of motor areas was more pronounced at the beginning of the rest period. In conclusion, sensory motor areas can be activated using a piezoelectric actuator, with less pronounced motor area activation at higher frequencies. Our setup allowed local control of stimulus penetration through the tissue correlated to central activation, providing objective stimulus control. The pronounced activation of the motor cortex during the rest condition may reflect the subjective feeling of arm movement after the end of the stimulus.

Age-Dependent Impairment of Somatosensory Response in the Amyloid Precursor Protein 23 Transgenic Mouse Model of Alzheimer's Disease

Quantitative functional magnetic resonance imaging was applied to characterize brain function in amyloid precursor protein 23 (APP23) transgenic mice, which reproduce the neuropathological alterations associated with Alzheimer's disease. Electrical stimulation of the paw led to cerebral blood volume increases in the contralateral somatosensory cortex. In APP23 mice this hemodynamic response decreased with increasing age of the animal and with increasing stimulus amplitude as compared with wild-type animals. The age-dependent dysfunction in APP23 mice may be attributed in part to a compromised cerebrovascular reactivity. Quantitative functional brain mapping that uses standardized sensory inputs should allow for assessment of disease progression and therapy response (e.g., passive immunization against beta-amyloid) in patients also.

The steady-state postural response to continuous sinusoidal galvanic vestibular stimulation

Galvanic vestibular stimulation (GVS) applied between the mastoids during quiet standing elicits postural sway. The aim of this study was to characterize the postural sway response to continuous sinusoidal GVS across various stimulus frequencies and amplitudes. Binaural bipolar sinusoidal GVS was applied to the skin overlying the mastoid processes of 10 subjects while they stood on a force plate with eyes closed. The position of the center of pressure (COP) at the feet was recorded from a forceplate, while the head displacement was measured with a magnetic position tracking system. The stimulus conditions included four frequencies (0.1, 0.25, 0.45, and 1.1 Hz) and five peak amplitudes (0.05, 0.1, 0.25, 0.5, and 1.0 mA). Each subject experienced one trial at each amplitude-frequency pair. Additionally, each subject underwent three trials in which a dual-frequency stimulus (0.1 plus 0.45 Hz at a peak of 0.5 mA each) was presented. The stimuli elicited sway in the frontal plane in all subjects, as evidenced by changes in the displacement of the COP and head. Sway magnitude decreased with increasing stimulus frequency and increased with increasing stimulus amplitude. However, the response magnitude saturated at higher stimulus amplitudes. Phase lag increased with increasing stimulus frequency. The response to the dual-frequency stimulus was reduced at 0.1 Hz and nearly equal at 0.45 Hz in comparison with the single-frequency responses. This study suggests that the postural sway response is nonlinear due to saturation and violation of the principle of superposition.

Sensorimotor integration in human postural control.

It is generally accepted that human bipedal upright stance is achieved by feedback mechanisms that generate an appropriate corrective torque based on body-sway motion detected primarily by visual, vestibular, and proprioceptive sensory systems. Because orientation information from the various senses is not always available (eyes closed) or accurate (compliant support surface), the postural control system must somehow adjust to maintain stance in a wide variety of environmental conditions. This is the sensorimotor integration problem that we investigated by evoking anterior-posterior (AP) body sway using pseudorandom rotation of the visual surround and/or support surface (amplitudes 0.5-8 degrees ) in both normal subjects and subjects with severe bilateral vestibular loss (VL). AP rotation of body center-of-mass (COM) was measured in response to six conditions offering different combinations of available sensory information. Stimulus-response data were analyzed using spectral analysis to compute transfer functions and coherence functions over a frequency range from 0.017 to 2.23 Hz. Stimulus-response data were quite linear for any given condition and amplitude. However, overall behavior in normal subjects was nonlinear because gain decreased and phase functions sometimes changed with increasing stimulus amplitude. "Sensory channel reweighting" could account for this nonlinear behavior with subjects showing increasing reliance on vestibular cues as stimulus amplitudes increased. VL subjects could not perform this reweighting, and their stimulus-response behavior remained quite linear. Transfer function curve fits based on a simple feedback control model provided estimates of postural stiffness, damping, and feedback time delay. There were only small changes in these parameters with increasing visual stimulus amplitude. However, stiffness increased as much as 60% with increasing support surface amplitude. To maintain postural stability and avoid resonant behavior, an increase in stiffness should be accompanied by a corresponding increase in damping. Increased damping was achieved primarily by decreasing the apparent time delay of feedback control rather than by changing the damping coefficient (i.e., corrective torque related to body-sway velocity). In normal subjects, stiffness and damping were highly correlated with body mass and moment of inertia, with stiffness always about 1/3 larger than necessary to resist the destabilizing torque due to gravity. The stiffness parameter in some VL subjects was larger compared with normal subjects, suggesting that they may use increased stiffness to help compensate for their loss. Overall results show that the simple act of standing quietly depends on a remarkably complex sensorimotor control system.

Experimental and computational analysis of monkey smooth pursuit eye movements.

Smooth pursuit eye movements are guided by visual feedback and are surprisingly accurate despite the time delay between visual input and motor output. Previous models have reproduced the accuracy of pursuit either by using elaborate visual signals or by adding sources of motor feedback. Our goal was to constrain what types of signals drive pursuit by obtaining data that would discriminate between these two modeling approaches, represented by the "image motion model" and the "tachometer feedback" model. Our first set of experiments probed the visual properties of pursuit with brief square-pulse and sine-wave perturbations of target velocity. Responses to pulse perturbations increased almost linearly with pulse amplitude, while responses to sine wave perturbations showed strong saturation with increasing stimulus amplitude. The response to sine wave perturbations was strongly dependent on the baseline image velocity at the time of the perturbation. Responses were much smaller if baseline image velocity was naturally large, or was artificially increased by superimposing sine waves on pulse perturbations. The image motion model, but not the tachometer feedback model, could reproduce these features of pursuit. We used a revision of the image motion model that was, like the original, sensitive to both image velocity and image acceleration. Due to a saturating nonlinearity, the sensitivity to image acceleration declined with increasing image velocity. Inclusion of this nonlinearity was motivated by our experimental results, was critical in accounting for the responses to perturbations, and provided an explanation for the unexpected stability of pursuit in the presence of perturbations near the resonant frequency. As an emergent property, the revised image motion model was able to reproduce the frequency and damping of oscillations recorded during artificial feedback delays. Our second set of experiments replicated prior recordings of pursuit responses to multiple-cycle sine wave perturbations, presented over a range of frequencies. The image motion model was able to reproduce the responses to sine wave perturbations across all frequencies, while the tachometer feedback model failed at high frequencies. These failures resulted from the absence of image acceleration signals in the tachometer model. We conclude that visual signals related to image acceleration are important in driving pursuit eye movements and that the nonlinearity of these signals provides stability. Smooth pursuit thus illustrates that a plausible neural strategy for combating natural delays in sensory feedback is to employ information about the derivative of the sensory input.

Facilitatory and inhibitory frequency tuning of combination-sensitive neurons in the primary auditory cortex of mustached bats.

Mustached bats, Pteronotus parnellii parnellii, emit echolocation pulses that consist of four harmonics with a fundamental consisting of a constant frequency (CF(1-4)) component followed by a short, frequency-modulated (FM(1-4)) component. During flight, the pulse fundamental frequency is systematically lowered by an amount proportional to the velocity of the bat relative to the background so that the Doppler-shifted echo CF(2) is maintained within a narrowband centered at approximately 61 kHz. In the primary auditory cortex, there is an expanded representation of 60.6- to 63. 0-kHz frequencies in the "Doppler-shifted CF processing" (DSCF) area where neurons show sharp, level-tolerant frequency tuning. More than 80% of DSCF neurons are facilitated by specific frequency combinations of approximately 25 kHz (BF(low)) and approximately 61 kHz (BF(high)). To examine the role of these neurons for fine frequency discrimination during echolocation, we measured the basic response parameters for facilitation to synthesized echolocation signals varied in frequency, intensity, and in their temporal structure. Excitatory response areas were determined by presenting single CF tones, facilitative curves were obtained by presenting paired CF tones. All neurons showing facilitation exhibit at least two facilitative response areas, one of broad spectral tuning to frequencies centered at BF(low) corresponding to a frequency in the lower half of the echolocation pulse FM(1) sweep and another of sharp tuning to frequencies centered at BF(high) corresponding to the CF(2) in the echo. Facilitative response areas for BF(high) are broadened by approximately 0.38 kHz at both the best amplitude and 50 dB above threshold response and show lower thresholds compared with the single-tone excitatory BF(high) response areas. An increase in the sensitivity of DSCF neurons would lead to target detection from farther away and/or for smaller targets than previously estimated on the basis of single-tone responses to BF(high). About 15% of DSCF neurons show oblique excitatory and facilitatory response areas at BF(high) so that the center frequency of the frequency-response function at any amplitude decreases with increasing stimulus amplitudes. DSCF neurons also have inhibitory response areas that either skirt or overlap both the excitatory and facilitatory response areas for BF(high) and sometimes for BF(low). Inhibition by a broad range of frequencies contributes to the observed sharpness of frequency tuning in these neurons. Recordings from orthogonal penetrations show that the best frequencies for facilitation as well as excitation do not change within a cortical column. There does not appear to be any systematic representation of facilitation ratios across the cortical surface of the DSCF area.

Regulation of muscle stiffness during periodic length changes in the isolated abdomen of the hermit crab.

Reflex activation of the ventral superficial muscles (VSM) in the abdomen of the hermit crab, Pagurus pollicarus, was studied using sinusoidal and stochastic longitudinal vibration of the muscle while recording the length and force of the muscle and the spike times of three exciter motoneurons. In the absence of vibration, the interspike interval histograms of the two larger motoneurons were bimodal; cutting sensory nerves containing most of the mechanoreceptor input removed the short interval peak in the histogram, indicating that the receptors are important in maintaining tonic firing. Vibration of the muscle evoked a reflex increase in motoneuron frequency that habituated after an initial peak but remained above control levels for the duration of stimulation. Motoneuron frequency increased with root mean square (rms) stimulus amplitude. Average stiffness during stimulation was about two times the stiffness of passive muscle. The reflex did not alter muscle dynamics. Estimated transfer functions were calculated from the fast Fourier transform of length and force signals. Coherence was >0.9 for the frequency range of 3-35 Hz. Stiffness magnitude gradually increased over this range in both reflex activated and passive muscle; phase was between 10 and 20 degrees. Reflex stiffness decreased with increasing stimulus amplitudes, but at larger amplitudes, this decrease was much less pronounced; in this range stiffness was regulated by the reflex. The sinusoidal frequency at which reflex bursts were elicited was approximately 6 Hz, consistent with previous measurements using ramp stretch. During reflex excitation, there was an increase in amplitude of the short interval peak in the interspike interval histogram; this was reduced when the majority of afferent pathways was removed. A phase histogram of motoneuron firing during sinusoidal vibration had a peak at approximately 110 ms, also suggesting that an important component of the reflex is via direct projections from the mechanoreceptors. These results are consistent with the hypothesis that a robust feedforward regulation of abdominal stiffness during continuous disturbances is achieved by mechanoreceptors signalling the absolute value of changing forces; habituation of the reflex, its high-threshold for low frequency disturbances and the activation kinetics of the muscle further modify reflex dynamics.

A dynamic characterization of postural stability using time-frequency analysis

Galvanic vestibular stimulation (GVS) applied between the mastoids during quiet standing elicits postural sway. The aim of this study was to characterize the postural sway response to continuous sinusoidal GVS across various stimulus frequencies and amplitudes. Binaural bipolar sinusoidal GVS was applied to the skin overlying the mastoid processes of 10 subjects while they stood on a force plate with eyes closed. The position of the center of pressure (COP) at the feet was recorded from a forceplate, while the head displacement was measured with a magnetic position tracking system. The stimulus conditions included four frequencies (0.1, 0.25, 0.45, and 1.1 Hz) and five peak amplitudes (0.05, 0.1, 0.25, 0.5, and 1.0 mA). Each subject experienced one trial at each amplitude–frequency pair. Additionally, each subject underwent three trials in which a dual-frequency stimulus (0.1 plus 0.45 Hz at a peak of 0.5 mA each) was presented. The stimuli elicited sway in the frontal plane in all subjects, as evidenced by changes in the displacement of the COP and head. Sway magnitude decreased with increasing stimulus frequency and increased with increasing stimulus amplitude. However, the response magnitude saturated at higher stimulus amplitudes. Phase lag increased with increasing stimulus frequency. The response to the dual-frequency stimulus was reduced at 0.1 Hz and nearly equal at 0.45 Hz in comparison with the single-frequency responses. This study suggests that the postural sway response is nonlinear due to saturation and violation of the principle of superposition.

Gain Control in the Femur–Tibia Feedback System of the Stick Insect

ABSTRACT This paper presents a quantitative description of the variability and the adaptive properties of information processing in the femur-tibia feedback system of the stick insect. The gain of this proprioceptive feedback system is determined by external stimuli changing the behavioural state of the animal and by internal properties that make it dependent on different parameters of the stimulus programme, e.g. stimulus frequency and amplitude, repetition rate and resting pauses. The gain of the feedback loop in the inactive animal was investigated under open-loop conditions by applying mechanical sine-wave stimuli to the femoral chordotonal organ (fCO). The resistance movement of the tibia caused by these stimuli was measured with a new optoelectronic device. A large increase or decrease in gain (up to a factor of 50) can be induced by stimulation, but also occurs spontaneously. The system shows habituation and sensitization. The initial gain can be decreased by repetitive sine-wave stimulation of the fCO. Disturbance of the animal (e.g. by tactile stimuli) increases the gain. The gain of the system decreases with increasing stimulus amplitude. The described nonlinearities form a system which adjusts gain to a value that permits effective feedback and prevents instability. This was verified by closed-loop experiments.

Optokinetic torsion: Dynamics and relation to circularvection

Continuous records of optokinetic torsion to sinusoidal inputs were obtained using the electromagnetic scleral search-coil technique. We measured the gain and phase lag of optokinetic torsion in response to a spherical visual display rotating steadily at various angular velocities and sinusoidally at frequencies from 0.2 to 2.0 Hz and at amplitudes from 10 to 80 deg. Gain (peak slow-phase eye velocity over stimulus angular velocity) of up to 0.12 were obtained with stimulus frequencies of 0.2 Hz and declined to an average value of about 0.02 at a frequency of 2.0 Hz. Phase lag was virtually zero at a frequency of 0.2 Hz and increased to over 80 deg at 2.0 Hz. The records from the sinsoidal stimuli show very few quick phases. With increasing stimulus amplitudes, the amplitude of the response increased but its gain declined. We found no evidence of torsional after-nystagmus nor any relation between the torsional response and reports of vection or sensation of body tilt induced by the rotating display. Torsional optokinetic nystagmus is most suited to compensate for low-amplitude, low-frequency stimulus rotation and normally supplements torsion induced by head tilt.