Richard C. Fitzpatrick

Proprioceptive, visual and vestibular thresholds for the perception of sway during standing in humans.

1. Thresholds for the perception of postural sway induced by gentle perturbations were determined for five normal standing subjects. In this context we understand ‘perception’ to mean ‘able to give a subjective report’. The thresholds for the perception of movements that were equivalent to sway in velocity and amplitude were determined when the available sensory input was limited to only one, or a pair, of the vestibular, visual, and proprioceptive systems. To examine vestibular inputs alone, vision was excluded and the whole body was moved with the ankles in a fixed position. To examine visual inputs alone, the body was kept stationary and a ‘room’ was moved around the subjects to simulate the relative visual‐field movement that occurs during standing. To limit the available sensory input to proprioception from the legs, subjects were held stationary and balanced a load that was equivalent to their own body using their ankles. In this situation, perturbations were applied to the ‘equivalent body’ and these could only be perceived from the resulting ankle movements. Thresholds for perceiving ankle movements were also determined in the same posture, but with the leg muscles bearing no load. 2. The thresholds for the perception of sway during standing were very small, typically 0.003 rad at a velocity of 0.001 rad s‐1, and even smaller movements were perceived as the mean velocity of the sway increased up to 0.003 rad s‐1. No difference was found between the thresholds for perceiving forward sway and backward sway. Eye closure during standing did not affect the threshold for perceiving sway. 3. When sensory input was limited to proprioception from the legs, the thresholds for the perception of passive ankle movements were equivalent to the thresholds for the perception of sway during standing with all sensory inputs available. When the leg muscles were relaxed, the thresholds for perceiving ankle movements increased approximately twofold. 4. The visual thresholds for perceiving movement were higher than the proprioceptive thresholds at slower velocities of movement, but there was no difference at higher velocities. 5. Both the proprioceptive and visual thresholds were sufficiently small to allow perception of the sway that was recorded when the subjects stood normally in a relaxed manner. In contrast, the vestibular thresholds were an order of magnitude greater than the visual or proprioceptive thresholds and above the largest sway movements that were recorded during normal standing.(ABSTRACT TRUNCATED AT 400 WORDS)

Acceleration patterns of the head and pelvis when walking on level and irregular surfaces.

The aim of this study was to evaluate acceleration patterns at the head and pelvis while subjects walked on a level and an irregular walking surface, to develop an understanding of how the postural control system responds to challenging walking conditions. Thirty young, healthy subjects walked on a level corridor and on artificial grass underlain with foam and wooden blocks placed in an arbitrary manner. Temporo-spatial gait parameters and acceleration patterns at the head and pelvis were measured. The results revealed that when walking on the irregular surface, subjects were able to maintain their velocity, but adopted a slower and more variable cadence and a significantly longer stride length. The magnitude of pelvis accelerations increased, however head accelerations were not affected by the walking surface. When considered as an overall pattern of movement, these findings suggest that one of the primary objectives of the postural control system when walking on irregular surfaces is head control, and that subjects adapt their stepping pattern on irregular surfaces to ensure that the head remains stable.

The vestibular system.

Supported by grants from the MRC of Great Britain and the NHMRC of Australia. We thank P.B.C. Matthews for his description of the silent sense.

Task-dependent reflex responses and movement illusions evoked by galvanic vestibular stimulation in standing humans.

1. To identify the vestibular contribution to human standing, responses in leg muscles evoked by galvanic vestibular stimulation were studied. Step impulses of current were applied between the mastoid processes of normal subjects and the effects on the soleus and tibialis anterior electromyograms (EMGs), ankle torque, and body sway were identified by post‐stimulus averaging. The responses were measured when subjects stood on a stable platform or on an unstable platform and the effects of eye closure were also assessed. Responses were also recorded during voluntary contraction of the leg muscles and when subjects balanced a load equivalent to their own body in a situation where vestibular postural reflexes would not be useful. 2. At a mean post‐stimulus latency of 56 ms, there were reciprocal changes in soleus and tibialis anterior muscle activity followed, at 105 ms, by larger responses of opposite sign. These were termed the short‐ and middle‐latency responses, respectively. Both responses increased with stimulus intensity, but the short‐latency response had a higher threshold. The early response had a similar latency to EMG responses evoked by rapid postural perturbations. Both responses were larger when the eyes were closed, but eye closure was associated with increased sway and EMG activity, and the responses were of similar magnitude when scaled to background EMG level. 3. Both short‐ and middle‐latency EMG responses in soleus and tibialis anterior muscles produced small transient postural sways. The transient changes in EMG activity were followed by a larger prolonged sway which was not attributable to the activity in these muscles but rather to reflex or volitional adjustments to movements at other body segments. When subjects were prevented from swaying, the galvanic stimulus produced illusory movements in the opposite direction to the sway evoked when standing, and it is possible that the prolonged sway is a reaction to the illusion of sway. 4. The short‐ and middle‐latency responses were modified during different postural tasks according to the dependence on vestibular reflexes. When the support platform was unstable, the EMG responses to galvanic stimulation were larger. There were no vestibular‐evoked responses when seated subjects made voluntary contractions of the leg muscles or when they stood upright with the trunk supported, using the ankles to balance a body‐like load.(ABSTRACT TRUNCATED AT 400 WORDS)

Lateral Stability, Sensorimotor Function and Falls in Older People

AIMS: To design simple tests of lateral stability for assessing balance in older people and to determine whether poor performances in these tests are associated with impaired vision, lower limb sensation, quadriceps strength, simple reaction time, and falling in this group.

Stable human standing with lower‐limb muscle afferents providing the only sensory input.

1. This study investigated the sources of sensory information upon which normal subjects’ ability to stand depends. 2. An ‘equivalent body’ was used to simulate the physical properties of each subjects body during standing. The modulation of ankle torque required to support the equivalent body in an upright position was similar to that required to support the subjects own body when standing. However, when balancing the equivalent body, vestibular inputs were excluded from directing the appropriate changes in ankle torque. Thus, stability of stance could be studied with (normal stance) and without (balancing equivalent body) modulation by vestibular inputs. Vision could be excluded by closing the eyes. Sensory input from the feet and ankles could be removed by local anaesthesia from prolonged ischaemia, induced by occluding blood flow with inflated pneumatic cuffs just above the ankles. With vestibular, visual and peripheral sensory inputs negated, standing could rely only upon remaining sensory inputs, notably those from sensory receptors in the leg muscles. 3. Unlike the human body, the equivalent body used to negate vestibular inputs is not segmented. Therefore, the effects on stability of having a segmented body were determined by splinting subjects during standing so that only ankle movement was possible. This was done in the presence and absence of visual stabilization. 4. For each experimental task, either standing or balancing the equivalent body, sway was recorded while posture was unperturbed. Root mean square values of sway amplitude and power spectra were used to compare conditions. 5. Every subject could balance the equivalent body in a stable way when the eyes were closed, and when the feet were anaesthetized.(ABSTRACT TRUNCATED AT 250 WORDS)

Ankle stiffness of standing humans in response to imperceptible perturbation: reflex and task-dependent components.

1. It has been demonstrated that subjects can alter the reflex stiffness of the elbow and wrist in response to imperceptibly slow perturbations applied through a complaint coupling. We used this technique to measure ankle stiffness in standing subjects as a means of examining reflex activity. 2. During unperturbed stance, a linear relationship between ankle torque and ankle angle is expressed as a load stiffness. The load stiffness predicted from a subjects measured physical dimensions corresponds closely with the value measured by standing the subject on a force platform. 3. Slow perturbations were applied at waist level, through a spring, to standing subjects. The perturbations caused sway similar in magnitude and rate to the sway of normal stance. Ankle stiffness was measured during the period when the perturbations were unperceived. The contribution to ankle stiffness of reflexes that use visual information was assessed by eye closure. The ability of reflexes based on sensory information from the legs to maintain upright posture was assessed when subjects balanced a load equivalent to their own body, in a situation where neither visual nor vestibular information could assist. Ankle stiffness was measured while the load was perturbed. 4. The results show that a simple mechanical model of stance predicts the torque‐angle relationship at the ankle. This relationship determines the minimal ankle stiffness required to stand, and reflex muscle stiffness is a necessary component of this ankle stiffness. Visual, vestibular and lower limb sensorimotor reflexes each contribute to ankle stiffness; however, the local sensory reflexes alone are sufficient to stand. For responses to unperceived perturbations, standing subjects can alter their reflex ankle stiffness according to intentional set.

Effects of galvanic vestibular stimulation during human walking

1 To identify vestibular influences on human walking, galvanic vestibular stimulation was applied to normal adult subjects as they walked to a previously seen target. A transmastoidal step stimulus commenced as subjects started walking. With the eyes shut, the galvanic stimulus caused large turns towards the side with the anodal current. 2 Ability to perceive the trajectory of gait without visual cues was measured by guiding blindfolded subjects from one arbitrary point to another, either walking or seated in a wheelchair. On reaching a destination position and removing the blindfold, subjects pointed to indicate the starting position. Subjects made considerable errors in estimating the trajectory, but were equally accurate whether in the wheelchair or walking. 3 To determine the effects of vestibular stimulation on the perception of trajectory, the galvanic stimulus was applied to blindfolded subjects as they were guided from one point to another in the wheelchair. The vestibular stimulus produced an illusory shift in the trajectory travelled. This shift was towards the side with the cathode, i.e. in the opposite direction to the turn produced by the stimulus during walking. 4 We conclude that galvanic vestibular stimulation during walking causes subjects to turn from their planned trajectory. In part, this altered course may compensate for an altered perception of trajectory produced by the stimulus. However, altered perception of the vertical or the base of support, or direct vestibulo‐fugal influences on the leg muscles could contribute to the changes in gait.

Otolith and canal reflexes in human standing

We used galvanic vestibular stimulation (GVS) to identify human balance reflexes of the semicircular canals and otolith organs. The experiment used a model of vestibular signals arising from GVS modulation of the net signal from vestibular afferents. With the head upright, the model predicts that the GVS‐evoked canal signal indicates lateral head rotation while the otolith signal indicates lateral tilt or acceleration. Both signify body sway transverse to the head. With the head bent forward, the model predicts that the canal signal indicates body spin about a vertical axis but the otolith signal still signifies lateral body motion. Thus, we compared electromyograms (EMG) in the leg muscles and body sway evoked by GVS when subjects stood with the head upright or bent forward. With the head upright, GVS evoked a large sway in the direction of the anodal electrode. This response was abolished with the head bent forward leaving only small, oppositely directed, transient responses at the start and end of the stimulus. With the head upright, GVS evoked short‐latency (60–70 ms), followed by medium‐latency (120 ms) EMG responses, of opposite polarity. Bending the head forward abolished the medium‐latency but preserved the short‐latency response. This is compatible with GVS evoking separate otolithic and canal reflexes, indicating that balance is controlled by independent canal and otolith reflexes, probably through different pathways. We propose that the short‐latency reflex and small transient sway are driven by the otolith organs and the medium‐latency response and the large sway are driven by the semicircular canals.

Large–scale movement and reef fidelity of grey reef sharks

Despite an Indo-Pacific wide distribution, the movement patterns of grey reef sharks (Carcharhinus amblyrhynchos) and fidelity to individual reef platforms has gone largely unstudied. Their wide distribution implies that some individuals have dispersed throughout tropical waters of the Indo-Pacific, but data on large-scale movements do not exist. We present data from nine C. amblyrhynchos monitored within the Great Barrier Reef and Coral Sea off the coast of Australia. Shark presence and movements were monitored via an array of acoustic receivers for a period of six months in 2008. During the course of this monitoring few individuals showed fidelity to an individual reef suggesting that current protective areas have limited utility for this species. One individual undertook a large-scale movement (134 km) between the Coral Sea and Great Barrier Reef, providing the first evidence of direct linkage of C. amblyrhynchos populations between these two regions. Results indicate limited reef fidelity and evidence of large-scale movements within northern Australian waters.