Simulation of vestibular semicircular canal responses during righting movements of a freely falling cat

The righting maneuver of a freely falling cat was filmed at 1000 pictures per second, and the head position about the roll axis was digitized from each film frame using a graphics input tablet. The head angular velocity and acceleration were computed from the roll axis position trajectory. Head acceleration trajectories approximated two periods of a damped sinusoid at a frequency of 26 Hz. Head acceleration peak amplitudes exceeded 120,000 deg/s². These trajectories were used as stimuli for the horizontal semicircular canals in a computer simulation of first-order afferent responses during the fall. Linear system afferent response dynamics, characterized in a previous study of the cat horizontal canal using pseudorandom rotations, provided the basis for linear predictions of falling cat afferent responses. Results showed predicted single afferent firing rates that exceeded physiological values; and variations in afferent sensitivities and phase were predicted among different neurons. Fast head movement information could be carried by ensemble populations of vestibular neurons, and a phase-locking encoding hypothesis is proposed which accomplishes this. Implications for central program versus peripheral vestibular feedback strategies for motor control during falling are presented and discussed.     

Recovering from Laboratory-Induced slips and trips causes high levels of lumbar muscle activity and spine loading

Slips, trips, and falls are some of the most substantial and prevalent causes of occupational injuries and fatalities, and these events may contribute to low-back problems. We quantified lumbar kinematics (i.e., lumbar angles relative to pelvis) and kinetics during unexpected slip and trip perturbations, and during normal walking, among 12 participants (6F, 6 M). Individual anthropometry, lumbar muscle geometry, and lumbar angles, along with electromyography from 14 lumbar muscles were used as input to a 3D, dynamic, EMG-based model of the lumbar spine. Results indicated that, in comparison with values during normal walking, lumbar range of motion, lumbosacral (L5/S1) loads, and lumbar muscle activations were all significantly higher during the slip and trip events. Maximum L5/S1 compression forces exceeded 2700 N during slip and trip events, compared with ∼ 1100 N during normal walking. Mean values of L5/S1 anteroposterior (930 N), and lateral (800 N) shear forces were also substantially larger than the shear force during the normal walking (230 N). These observed levels of L5/S1 reaction forces, along with high levels of bilateral lumbar muscle activities, suggest the potential for overexertion injuries and tissue damage during unexpected slip and trip events, which could contribute to low back injuries. Outcomes of this study may facilitate the identification and control of specific mechanisms involved with low back disorders consequent to slips or trips.     

EMG modulation in anticipation of a possible trip during walking in young and older adults.

This study investigated whether changes in lower limb muscle activity occurred in anticipation of a possible perturbation in 11 young (mean age 27 years) and 11 older (mean age 68 years) adults. Altered muscle activity could affect tripping responses and consequently the ecological validity of experimental results of studies on tripping. It was hypothesized that anticipatory muscle activity would be present immediately after a trip, and decrease after several subsequent unperturbed (forewarned) walking trials. Electromyograms of lower limb muscles were measured in 3 conditions: during normal walking, during forewarned walking immediately after a trip, and during forewarned walking several trials after a trip had occurred. Small but statistically significant differences in averaged muscle activity over a stride were found among conditions. Young adults showed slightly increased activity immediately after tripping (co-contraction) in hamstrings, quadriceps and tibialis anterior muscles. This increased activity diminished after several unperturbed trials, although it did not return to the baseline activity levels during normal walking. In older adults, an increased muscle activity among conditions was only discerned in tibialis anterior and soleus muscles. This suggested that older adults prefer to avoid contact with the obstacle over joint stiffening. Yet, for both age-groups, the increases in muscle activity were very small when compared to tripping responses reported in the literature. Therefore, anticipatory effects are not expected to jeopardize the validity of experiments in which subjects are perturbed more than once.     

Slip-induced fall-risk assessment based on regular gait pattern in older adults

Aging-associated fall-risk assessment is crucial for fall prevention. Thus, this study aimed to develop a prognostic model to predict fall-risk following an unexpected over-ground slip perturbation based on normal gait pattern in healthy older adults. 112 healthy older adults who experienced a novel slip in a safe laboratory environment were included. Their slip trial and natural walking trial immediately prior to it were analyzed. To identify the best fall-risk predictive model, gait related variables including step length, segment angles, center of mass state, and ground reaction force (GRF) were determined and inputted into a stepwise logistic regression. The optimal slip-induced fall prediction model was based on the right thigh angle at slipping foot touchdown (TD), the maximum GRF of the slipping limb after TD, and the momentum change from TD to recovery foot liftoff (LO), with an overall prediction accuracy of 75.9%, predicting 74.5% of falls (sensitivity) and 77.2% of recoveries (specificity). Conversely, a model based on clinical and demographic measures predicted 78.2% of falls and 47.4% of recoveries, resulting in a much lower overall accuracy of 62.5%. The fall-risk model based on normal gait pattern which was developed for slip-induced perturbations in healthy older adults was able to provide a high predictive accuracy. This information could provide insight about the ideal normal gait measures which could be used to contribute towards development of therapeutic strategies related to dynamic balance and fall prevention to enhance preventive interventions in populations with high-risk for slip-induced falls.     

Disturbance type and gait speed affect fall direction and impact location

Since falling to the side and impacting on or near the hip increase hip fracture risk, we examined the fall direction and pelvis impact location resulting from four disturbances (faint, slip, step down, trip) at three gait speeds (fast, normal, slow) in 14 young adults instructed not to attempt recovery. We hypothesized that certain disturbances such as faints and slips and slow walking speed were more likely to result in an impact on the hip. For each trial, the fall direction, impact location and pelvis impact velocity were measured. The results showed that both disturbance type and gait speed significantly affected fall direction and impact location (analysis of covariance with repeated measures, p< or =0.0001) with a significant interaction (p<0.05). Trips and steps down usually resulted in forward falls, with frontal impacts regardless of gait speed. At fast gait speed, slips and faints also usually resulted in forward falls, with frontal impacts. As gait speed decreased, however, slips usually resulted in sideways or backward falls, with impact on the hip or buttocks, and faints resulted in a greater number of sideways falls, with impact near the hip. Therefore, compared to other disturbances and gait speeds, slipping or fainting while walking slowly was more likely to result in an impact on the hip, suggesting a greater risk for hip fracture. Furthermore, 56% of the impact velocities generated were within one standard deviation of the estimate of the mean impact velocity needed to fracture an elderly femur.     

Trip recovery strategies following perturbations of variable duration

Appropriately responding to mechanical perturbations during gait is critical to maintain balance and avoid falls. Tripping perturbation onset during swing phase is strongly related to the use of different recovery strategies; however, it is insufficient to fully explain how strategies are chosen. The dynamic interactions between the foot and the obstacle may further explain observed recovery strategies but the relationship between such contextual elements and strategy selection has not been explored. In this study, we investigated whether perturbation onset, duration and side could explain strategy selection for all of swing phase. We hypothesized that perturbations of longer duration would elicit lowering and delayed-lowering strategies earlier in swing phase than shorter perturbations. We developed a custom device to trip subjects multiple times while they walked on a treadmill. Seven young, healthy subjects were tripped on the left or right side at 10% to 80% of swing phase for 150 ms, 250 ms or 350 ms. Strategies were characterized by foot motion post-perturbation and identified by an automated algorithm. A multinomial logistic model was used to investigate the effect of perturbation onset, side, and the interaction between duration and onset on recovery strategy selection. Side perturbed did not affect strategy selection. Perturbation duration interacted with onset, limiting the use of elevating strategies to earlier in swing phase with longer perturbations. The choice between delayed-lowering and lowering strategies was not affected by perturbation duration. Although these variables did not fully explain strategy selection, they improved the prediction of strategy used in response to tripping perturbations throughout swing phase.     

Use of a cane for recovery from backward balance loss during treadmill walking

Purpose. To study whether a cane improved balance recovery after perturbation during walking.

Method. This study was a crossover comparison comparing the effect of walking with and without a cane for balance recovery after perturbation during treadmill walking. Five normal young volunteers participated. The velocity and acceleration of a marker sited on the seventh cerebral vertebra (C7) and vertical hand motion were measured by a motion analysis system.

Result. When using a cane, C7 backward velocity increased by approximately 15% (413 SD 95?mm/s with cane vs. 358 SD 88?mm/s without). In addition, C7 backward acceleration increased by approximately 23% (3.2 SD 0.7?m/s² with cane vs. 2.6 SD 0.8?m/s² without) and the vertical motion of the right hand decreased (187 SD 98?mm with cane vs. 372 SD 260?mm without). Additionally, no subject was able to use a cane to broaden their base of support.

Conclusions. The ability to limit trunk extension is crucial for preventing falls. Therefore, using a cane jeopardizes recovery from backward balance loss. The results encourage further research on the risk of a cane on balance recovery for the elderly population and habitual cane users.     

Two types of slip-induced falls among community dwelling older adults

Little is known about the landing behavior of the trailing (recovery) foot and ensuing types of falls following a forward slip in walking. The purposes of this study were to (1) determine if community-dwelling older adults experienced bilateral slips at the same rate as had been previously observed for young adults during over-ground walking; (2) determine if fall rate in older adults was dependent on slip type (unilateral vs. bilateral); and (3) identify differences in spatiotemporal variables of the trailing leg step between unilateral and bilateral slips. One-hundred-seventy-four participants experienced an unannounced, unrehearsed slip while walking on a 7-m walkway. Each trial was monitored with a motion capture system and bilateral ground reaction force plates. Although the experimental design, developed with original data from a young adult population, favored bilateral slips, more older adults (35%) than anticipated (10% previously observed in young, p<0.001) displayed a unilateral slip. The probability of fall was equal in the two types of slips. Eighty-two people recovered from the slip, while the remaining 92 (53%) fell. These 92 were classified into two exclusive categories based on the heel distance at the time of fall arrest using cluster analysis: those which resembled a fall into a "splits" position (n=47) or a feet-forward fall (n=45). All (100%) unilateral slips led to splits falls, as expected. Yet, not all bilateral slips (only 83%) resulted in feet-forward falls. A longer forward recovery step with a prolonged step time led to both feet slipping, nearly together, hence a feet-forward fall.     

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.     

Head position during various locomotor executions after prolonged microgravity exposure

Head orientation in the sagittal plane was assessed before and the second and sixth days after a prolonged microgravity exposure (6 months) in 7 astronauts, which were instructed to execute various locomotor movements (stair ascent, jump down and level walk). Crewmembers systematically flexed down the head when they executed the locomotion the second day post-flight compared to the pre-flight walk. Cephalic orientation was normally positioned when movements was performed the sixth day after Earth re-entry. This head flexion may reflect a strategy used by cosmonauts to increase the vestibular sensibility, and then improve the control of the dynamic equilibrium during locomotor movements. The high rate of re-adaptation indicated that cosmonauts rapidly recovered their normal walking capacities despite the prolonged microgravity exposure.     

Lower extremity corrective reactions to slip events.

A significant number of injuries in the workplace is attributed to slips and falls. Biomechanical responses to actual slip events determine whether the outcome of a slip will be recovery or a fall. The goal of this study was to examine lower extremity joint moments and postural adjustments for experimental evidence of corrective strategies evoked during slipping in an attempt to prevent falling. Sixteen subjects walked onto a possibly oily vinyl tile floor, while ground reaction forces and body motion were recorded at 350 Hz. The onset of corrective reactions by the body in an attempt to recover from slips became evident at about 25% of stance and continued until about 45% into stance, i.e. on average between 190 and 350 ms after heel contact. These reactions included increased flexion moment at the knee and extensor activity at the hip. The ankle, on the other hand, acted as a passive joint (no net moment) during fall trials. Joint kinematics showed increased knee flexion and forward rotation of the shank in an attempt to bring the foot back towards the body. Once again, the ankle kinematics appeared to play a less dominant role (compared to the knee) in recovery attempts. This study indicates that humans generate corrective reactions to slips that are different than previously reported responses to standing perturbations translating the supporting surface.     

Recording of short-latency vestibular evoked potentials induced by acceleration impulses in experimental animals: current status of the method and its applications

The short-latency vestibular evoked potential (VsEP) induced by angular acceleration impulses (maximal amplitude 30,000 deg/sec², rise time 2–3 msec) was recorded by skin electrodes in intact cats after various surgical and pharmacological procedures. The normal VsEP consists of 5–8 waves, several microvolts in amplitude, during the first 10 msec. The latency of the first wave (P1) is about 2 msec with respect to the start of head acceleration. The first and the second waves (P1 and P2) were shown to originate from the vestibular nerve and nucleus, respectively.The VsEP disappears permanently after bilateral labyrinthectomy, excision of the 8th nerves, or administration of large doses of gentamicin. Temporary disappearance is caused by anoxia induced for a brief period of time or injection of lidocaine (4%) into the vestibular nerve or into the inner ear after contralateral labyrinthectomy.The VsEPs in the intact cat are similar whether clockwise or counterclockwise stimuli are used and are not affected by changing the position of the head. Unilaterally labyrinthectomized animals, however, show asymmetric response whereby excitatory stimulation of any of the intact semicircular canals evokes prominent P1 and P2 waves which are absent with inhibitory stimulation.The rate and input-output intensity functions of the VsEP are described. The threshold of the VsEP was found to be 1000–1500 deg/sec².In addition to the neurogenic waves, 2 other potentials appear occasionally in the response: (1) large-amplitude and longer-duration waves with latencies of 8–20 msec, which are of myogenic origin, and (2) smaller waves with shorter latency which probably represent vestibular microphonics and generator potentials. Extracellular recordings of the responses of single second-order neurons in the vestibular nuclei to the same acceleration impulses confirmed that the kinetic vestibular neurons can respond to these stimuli with a latency as short as 3.5 msec.This method for inducing and recording VsEPs has proved to be a powerful tool for the evaluation of vestibular function in experimental animal models.     

Angular acceleration,compensatory head movements and the halteres of flies (Lucilia serricata)

Summary Tethered flies were subjected to accelerations about their vertical axes while flying or walking. These accelerations were applied either suddenly to stationary animals or continuously by oscillating the animal from side to side. Head and wing movements resulting from the imposed angular accelerations were photographed with a camera and a stroboscopic flash.Analysis of the photographs shows that the wing movements act to counter the imposed angular accelerations and that during sinusoidal oscillations about the vertical axis, head turns are in antiphase with angular acceleration.Head turns do not occur when the halteres are absent or present and not oscillating. When oscillating, the halteres detect high values of angular acceleration, outside the known capabilities of the visual movement detection system.     

Factors influencing codling moth larval response to α-farnesene

Behavioral responses of newly-emerged codling moth (Cydia pomonella L.) larvae to α-farnesene were compared for a laboratory-reared strain (‘lab’, 160 generations inbred) and a recently collected strain (‘wild’, 3 generations). Video recordings of single larvae placed 20 mm from Bond papers releasing a range of seven α-farnesene concentrations were scored for head turning, head lifting, head direction, and movement across a Petri dish. The laboratory strain was significantly less successful at finding the treated papers than the wild strain. The lab strain was also significantly less active, shown by more random orientation, slower walking speed, less head turning and less head lifting. Both strains showed lower response to α-farnesene concentrations less than 10⁻⁶ (w/v) α-farnesene. Orientation, walking speed, and head turning rate were significantly higher in the presence of α-farnesene concentrations >10⁻⁷ w/v, for both strains. Head lifting did not show any effect from α-farnesene. Head turning was associated with both head lifting and walking speed and this association improved with the presence of α-farnesene. Increased attraction to odour source was correlated with improved direction finding at concentrations above 10⁻⁷ w/v. Activity factors such as head turning and walking speed influenced time to locate odour source more than orientation factors. Odour-guided orientation to α-farnesene in codling moth larvae was composed of both locomotory, and to a lesser extent, orientation responses, which increased in a directed fashion to the stimulus.     

Role of stability and limb support in recovery against a fall following a novel slip induced in different daily activities

The purpose of this study was to determine whether stability and limb support play a similar role in governing slip outcome in gait-slip as in sit-to-stand-slip, and whether such prediction could also be derived based on measures of these variables during regular, unperturbed movements. Fifty-three and forty-one young subjects all took one recovery step following an unannounced, novel, forward slip induced in gait and in sit-to-stand, respectively. Logistic regression was used to predict recovery outcome based on preslip and reactive measures of stability and limb support across tasks. Following slip onset, all subjects in both tasks experienced rapid decay in stability and limb support (indicated by a hip descent), leading to some actual falls that could not have been predicted from regular, preslip walking. Immediately before recovery step touchdown, stability and limb support could together best predict 88.9% and 100% falls, respectively, for gait-slip and sit-to-stand-slip. Because of differences in the execution of the recovery step, stability became a better predictor of fallers in sit-to-stand-slip than in gait-slip after recovery limb touchdown. Recovery steps were highly effective in restoring stability, regardless of outcome and task. The predictive strength of stability diminished in gait-slip or reduced in sit-to-stand-slip after recovery touchdown, while limb support remained able to differentiate fallers from those who recovered in both tasks. When slip-induced instability was combined with inadequate limb support, falls were nearly inevitable in both tasks.     

Detection of tripping gait patterns in the elderly using autoregressive features and support vector machines

Elderly tripping falls cost billions annually in medical funds and result in high mortality rates often perpetrated by pulmonary embolism (internal bleeding) and infected fractures that do not heal well. In this paper, we propose an intelligent gait detection system (AR-SVM) for screening elderly individuals at risk of suffering tripping falls. The motivation of this system is to provide early detection of elderly gait reminiscent of tripping characteristics so that preventive measures could be administered. Our system is composed of two stages, a predictor model estimated by an autoregressive (AR) process and a support vector machine (SVM) classifier. The system input is a digital signal constructed from consecutive measurements of minimum toe clearance (MTC) representative of steady-state walking. The AR-SVM system was tested on 23 individuals (13 healthy and 10 having suffered at least one tripping fall in the past year) who each completed a minimum of 10 min of walking on a treadmill at a self-selected pace. In the first stage, a fourth order AR model required at least 64 MTC values to correctly detect all fallers and non-fallers. Detection was further improved to less than 1 min of walking when the model coefficients were used as input features to the SVM classifier. The system achieved a detection accuracy of 95.65% with the leave one out method using only 16 MTC samples, but was reduced to 69.57% when eight MTC samples were used. These results demonstrate a fast and efficient system requiring a small number of strides and only MTC measurements for accurate detection of tripping gait characteristics.     

The startle response during whiplash: a protective or harmful response?

Whiplash injuries are common following rear-end collisions. During such collisions, initially relaxed occupants exhibit brisk, stereotypical muscle responses consisting of postural and startle responses that may contribute to the injury. Using prestimulus inhibition, we sought to determine if the startle response elicited during a rear-end collision contributes to head stabilization or represents a potentially harmful overreaction of the body. Three experiments were performed. In the first two experiments, two groups of 14 subjects were exposed to loud tones (124 dB) preceded by prestimulus tones at either four interstimulus intervals (100-1,000 ms) or five prestimulus intensities (80-124 dB). On the basis of the results of the first two experiments, 20 subjects were exposed to a simulated rear-end collision (peak sled acceleration = 2 g; speed change = 0.75 m/s) preceded by one of the following: no prestimulus tone, a weak tone (85 dB), or a loud tone (105 dB). The prestimulus tones were presented 250 ms before sled acceleration onset. The loud prestimulus tone decreased the amplitude of the sternocleidomastoid (16%) and cervical paraspinal (29%) muscles, and key peak kinematics: head retraction (17%), horizontal head acceleration (23%), and head angular acceleration in extension (23%). No changes in muscle amplitude or kinematics occurred for the weak prestimulus. The reduced muscle and kinematic responses observed with loud tones suggest that the startle response represents an overreaction that increases the kinematics in a way that potentially increases the forces and strains in the neck tissues. We propose that minimizing this overreaction during a car collision may decrease the risk of whiplash injuries.     

Validation of a device for accurate timing of feeding events in marine animals

Two acceleration data loggers, each measuring surging and heaving acceleration, were attached to the head and mandible of three captive hooded seals, Cystophora cristata, for detection of underwater feeding events. Three sizes of prey: Atlantic herring (large), capelin (medium), and half a capelin (small) were tested. A highpass frequency-filtering method at 3 Hz provided more distinct prey ingestion signals for both head and mandible acceleration. The surge-axis signals from head acceleration suggested that the seals ingested their prey not only by biting, but also by thrusting. Moreover, prey ingestion movements showed higher surging acceleration from mandible than from head (mean ± SD from head: 5.37 ± 4.45 m s⁻², from mandible: 8.43 ± 5.15 m s⁻², n = 153), indicating that the data from the head is not required for precise identification of feeding events. Thus, our mandible acceleration device provides a practical method for the timing of underwater feeding events in seals.     

Evaluation of gait and slip parameters for adults with intellectual disability

Adults with intellectual disability (ID) experience more falls than their non-disabled peers. A gait analysis was conducted to quantify normal walking, and an additional slip trial was performed to measure slip response characteristics for adults with ID as well as a group of age- and gender-matched controls. Variables relating to gait pattern, slip propensity, and slip severity were assessed to compare the differences between groups. The ID group was found to have significantly slower walking speed, shorter step lengths, and increased knee flexion angles at heel contact. These gait characteristics are known to reduce the likelihood of slip initiation in adults without ID. Despite a more cautious gait pattern, however, the ID group exhibited greater slip distances indicating greater slip severity. This study suggests that falls in this population may be due to deficient slip detection or insufficient recovery response.