Stepping in circles: how locomotor signals of rotation adapt over time.

KEY POINTS: While it has been well described that prolonged rotational stepping will adapt the podokinetic sense of rotation, the mechanisms involved are not clearly understood. By studying podokinetic after-rotations following conditioning rotations not previously reported we have shown that slower rotational velocities are more readily adapted than faster velocities and adaptation occurs more quickly than previously thought. We propose a dynamic feedback model of vestibular and podokinetic adaptation that can fit rotation trajectories across multiple conditions and data sets. Two adaptation processes were identified that may reflect central and peripheral processes and the discussion unifies prior findings in the podokinetic literature under this new framework. The findings show the technique is feasible for people with locomotor turning problems. ABSTRACT: After a prolonged period stepping in circles, people walk with a curved trajectory when attempting to walk in a straight line without vision. Podokinetic adaptation shows promise in clinical populations to improve locomotor turning; however, the adaptive mechanisms involved are poorly understood. The first phase of this study asks: how does the podokinetic conditioning velocity affect the response velocity and how quickly can adaptation occur? The second phase of the study asks: can a mathematical feedback model account for the rotation trajectories across different conditioning parameters and different datasets? Twelve healthy participants stepped in place on the axis of a rotating surface ranging from 4 to 20 deg s-1 for durations of 1-10 min, while using visual cues to maintain a constant heading direction. Afterward on solid ground, participants were blindfolded and attempted to step without rotating. Participants unknowingly stepped in circles opposite to the direction of the prior platform rotation for all conditions. The angular velocity of this response peaked within 1 min and the ratio of the stimulus-to-response peak velocity fitted a decreasing power function. The response then decayed exponentially. The feedback model of podokinetic and vestibular adaptive processes had a good fit with the data and suggested that podokinetic adaptation is explained by a short (141 s) and a long (27 min) time constant. The podokinetic system adapts more quickly than previously thought and subjects adapt more readily to slower rotation than to faster rotation. These findings will have implications for clinical applications of the technique.

Passive motion reduces vestibular balance and perceptual responses.

With the hypothesis that vestibular sensitivity is regulated to deal with a range of environmental motion conditions, we explored the effects of passive whole-body motion on vestibular perceptual and balance responses. In 10 subjects, vestibular responses were measured before and after a period of imposed passive motion. Vestibulospinal balance reflexes during standing evoked by galvanic vestibular stimulation (GVS) were measured as shear reaction forces. Perceptual tests measured thresholds for detecting angular motion, perceptions of suprathreshold rotation and perceptions of GVS-evoked illusory rotation. The imposed conditioning motion was 10 min of stochastic yaw rotation (0.5-2.5 Hz ≤ 300 deg s(-2) ) with subjects seated. This conditioning markedly reduced reflexive and perceptual responses. The medium latency galvanic reflex (300-350 ms) was halved in amplitude (48%; P = 0.011) but the short latency response was unaffected. Thresholds for detecting imposed rotation more than doubled (248%; P < 0.001) and remained elevated after 30 min. Over-estimation of whole-body rotation (30-180 deg every 5 s) before conditioning was significantly reduced (41.1 to 21.5%; P = 0.033). Conditioning reduced illusory vestibular sensations of rotation evoked by GVS (mean 113 deg for 10 s at 1 mA) by 44% (P < 0.01) and the effect persisted for at least 1 h (24% reduction; P < 0.05). We conclude that a system of vestibular sensory autoregulation exists and that this probably involves central and peripheral mechanisms, possibly through vestibular efferent regulation. We propose that failure of these regulatory mechanisms at different levels could lead to disorders of movement perception and balance control during standing.

Blood pressure and the contractility of a human leg muscle.

These studies investigate the relationships between perfusion pressure, force output and pressor responses for the contracting human tibialis anterior muscle. Eight healthy adults were studied. Changing the height of tibialis anterior relative to the heart was used to control local perfusion pressure. Electrically stimulated tetanic force output was highly sensitive to physiological variations in perfusion pressure showing a proportionate change in force output of 6.5% per 10 mmHg. This perfusion-dependent change in contractility begins within seconds and is reversible with a 53 s time constant, demonstrating a steady-state equilibrium between contractility and perfusion pressure. These stimulated contractions did not produce significant cardiovascular responses, indicating that the muscle pressor response does not play a major role in cardiovascular regulation at these workloads. Voluntary contractions at forces that would require constant motor drive if perfusion pressure had remained constant generated a central pressor response when perfusion pressure was lowered. This is consistent with a larger cortical drive being required to compensate for the lost contractility with lower perfusion pressure. The relationship between contractility and perfusion for this large postural muscle was not different from that of a small hand muscle (adductor pollicis) and it responded similarly to passive peripheral and active central changes in arterial pressure, but extended over a wider operating range of pressures. If we consider that, in a goal-oriented motor task, muscle contractility determines central motor output and the central pressor response, these results indicate that muscle would fatigue twice as fast without a pressor response. From its extent, timing and reversibility we propose a testable hypothesis that this change in contractility arises through contraction- and perfusion-dependent changes in interstitial K(+) concentration.

Perception of the postural vertical and falls in older people.

BACKGROUND: Research on the relationship between vestibular function and falls in older people is sparse. The perception of the postural vertical (PPV) provides an indicator measure of vestibular (otolith) function in the absence of visual input and diminished somatosensory feedback. OBJECTIVE: This study examined whether impaired PPV is associated with falls in this group. METHODS: One hundred and ninety-five people aged 70 plus years stood blindfolded on a motorised platform that could be tilted in the roll plane and attempted to adjust it so that their bodies were aligned to the vertical. Somatosensory feedback was minimised as the base and vertical support surfaces on the tilting platform were covered in thick soft foam rubber. PPV error from true vertical and PPV variability (°) were calculated. Participants also underwent an assessment of distal tactile sensitivity and the physiological profile assessment (PPA); fallers were defined as those who had one or more falls during a prospective 12-month follow-up period. RESULTS: Eighty-eight participants (45%) reported falling in the follow-up year. Increased PPV error and variability were correlated with increased lateral sway in a condition of absent visual input and reduced foot somatosensory feedback (eyes closed/foam; r range = 0.16-0.20, p < 0.05) and with composite PPA fall risk scores (r range = 0.22-0.26, p < 0.05). PPV variability was a significant and independent predictor of falls after adjusting for the composite PPA scores, age and gender [adjusted RR = 1.42 (1.01-1.98)]. CONCLUSIONS: Older people with increased PPV variability are at increased risk of falls. These findings indicate that assessment of PPV may augment fall risk assessments in older people.

The sense of self-motion, orientation and balance explored by vestibular stimulation.

The sense of orientation during locomotion is derived from our spatial relationship with the external environment, sensed predominantly by sight and sound, and from internal signals of motion, generated by the vestibular sense and the pattern of efferent and afferent signals to the muscles and joints. The sensory channels operate in different reference frames and have different time-dependent adaptive properties and yet the inputs are combined by the central nervous system to create an internal representation of self-motion. In normal circumstances vestibular, visual and proprioceptive cues provide congruent information on locomotor trajectory; however, in cases of sensory discord there must be a recalibration of sensory signals to provide a unitary representation. We develop a means of studying these fusion processes by perturbing each channel in isolation about a consistent behavioural axis. This review focuses on creating the vestibular perturbation of the orientation sense by transmastoidal galvanic stimulation, a technique generally used to evoke balance reflexes. Vector summation across the population of semicircular canal afferents creates a net signal that is interpreted by the brain as a vector of angular acceleration in a craniocentric reference frame. The signal feeds perceptual processes of orientation after transformation that resolves the 3-D signal onto the terrestrial or behavioural plane. Changing head posture changes the interpretation of the galvanic vestibular signal for balance and orientation responses. With appropriate head alignments during locomotion, the galvanic stimulus can be used to either steer trajectory over the terrestrial plane or perturb balance.

Adaptation of vestibular signals for self-motion perception.

A fundamental concern of the brain is to establish the spatial relationship between self and the world to allow purposeful action. Response adaptation to unvarying sensory stimuli is a common feature of neural processing, both peripherally and centrally. For the semicircular canals, peripheral adaptation of the canal-cupula system to constant angular-velocity stimuli dominates the picture and masks central adaptation. Here we ask whether galvanic vestibular stimulation circumvents peripheral adaptation and, if so, does it reveal central adaptive processes. Transmastoidal bipolar galvanic stimulation and platform rotation (20 deg s−1) were applied separately and held constant for 2 min while perceived rotation was measured by verbal report. During real rotation, the perception of turn decayed from the onset of constant velocity with a mean time constant of 15.8 s. During galvanic-evoked virtual rotation, the perception of rotation initially rose but then declined towards zero over a period of ∼100 s. For both stimuli, oppositely directed perceptions of similar amplitude were reported when stimulation ceased indicating signal adaptation at some level. From these data the time constants of three independent processes were estimated: (i) the peripheral canal-cupula adaptation with time constant 7.3 s, (ii) the central 'velocity-storage' process that extends the afferent signal with time constant 7.7 s, and (iii) a long-term adaptation with time constant 75.9 s. The first two agree with previous data based on constant-velocity stimuli. The third component decayed with the profile of a real constant angular acceleration stimulus, showing that the galvanic stimulus signal bypasses the peripheral transformation so that the brainstem sees the galvanic signal as angular acceleration. An adaptive process involving both peripheral and central processes is indicated. Signals evoked by most natural movements will decay peripherally before adaptation can exert an appreciable effect, making a specific vestibular behavioural role unlikely. This adaptation appears to be a general property of the internal coding of self-motion that receives information from multiple sensory sources and filters out the unvarying components regardless of their origin. In this instance of a pure vestibular sensation, it defines the afferent signal that represents the stationary or zero-rotation state.

The fusimotor and reafferent origin of the sense of force and weight.

Signals associated with the command the brain sends to muscles are thought to create the sensation of heaviness when we lift an object. Thus, as a muscle is weakened by fatigue or partial paralysis (neuromuscular blockade), the increase in the motor command needed to lift a weight is thought to explain the increasing subjective heaviness of the lifted object.With different fatiguing contractions we approximately halved the force output of the thumb flexor muscles, which were then used to lift an object. For two deafferented subjects the perceived heaviness of the lifted object approximately doubled, in keeping with the central-signal theory. However, for normal subjects this resulted in objects feeling the same or lighter, inconsistent with the central-signal theory but consistent with the expected effects of the conditioning contractions on the sensitivity of peripheral receptors. In separate experiments we subjected the forearm muscles to complete paralysis with a non-depolarising neuromuscular blocking agent and then allowed them to recover to approximately half-force output. This also resulted in objects feeling lighter when lifted by the semi-paralysed thumb, even though the motor command to the motoneurons must have been greater. This is readily explained by reduced lift-related reafference caused by the prolonged paralysis of muscle spindle intrafusal fibres.We conclude that peripheral signals, including a major contribution from muscle spindles, normally give rise to the sense of exerted force. In concept, however, reafference from peripheral receptors may also be considered a centrally generated signal that traverses efferent and then afferent pathways to feed perceptual centres rather than one confined entirely to the central nervous system. These results therefore challenge the distinction between central- and peripheral-based perception, and the concept that muscle spindles provide only information about limb position and movement.

The human semicircular canal model of galvanic vestibular stimulation.

A vector summation model of the action of galvanic stimuli on the semicircular canals has been shown to explain empirical balance and perceptual responses to binaural-bipolar stimuli. However, published data suggest binaural-monopolar stimuli evoke responses that are in the reverse direction of the model prediction. Here, we confirm this by measuring balance responses to binaural-monopolar stimulation as movements of the upper trunk. One explanation for the discrepancy is that the galvanic stimulus might evoke an oppositely directed balance response from the otolith organs that sums with and overrides the semicircular canal response. We tested this hypothesis by measuring sway responses across the full range of head pitch. The results showed some modulation of sway with pitch such that the maximal response occurred with the head in the primary position. However, the effect fell a long way short of that required to reverse the canal sway response. This indicates that the model is incomplete. Here, we examine alterations to the model that could explain both the bipolar and monopolar-evoked behavioural responses. An explanation was sought by remodelling the canal response with more recent data on the orientation of the individual canals. This improved matters but did not reverse the model prediction. However, the model response could be reversed by either rotating the entire labyrinth in the skull or by altering the gains of the individual canals. The most parsimonious solution was to use the more recent canal orientation data coupled with a small increase in posterior canal gain.

Lack of otolith involvement in balance responses evoked by mastoid electrical stimulation.

Passing current through mastoid electrodes (conventionally termed galvanic vestibular stimulation; GVS) evokes a balance response containing a short- and a medium-latency response. The origins of these two responses are debated. Here we test the hypotheses that they originate from net signals evoked by stimulation of otolith and semi-circular canal afferents, respectively. Based on anatomy and function, we predicted the directions of the stimulus-evoked net head rotation vector from the canals and the linear acceleration net vector from the otoliths. We tested these predictions in healthy adults by obtaining responses with the head in strategic postures to alter the relevance of the signals to the balance system. Cross-covariance between a stochastic waveform of stimulating current and motor output was used to assess the balance responses. Consistent with the canal hypothesis, with the head pitched down the medium-latency EMG response was abolished while the short-latency EMG response was maintained. The results, however, did not support the otolith hypothesis. The direction of the linear acceleration signal from the otoliths was predicted to change substantially when using monaural stimuli compared to binaural stimuli. In contrast, short-latency response direction measured from ground-reaction forces was not altered. It was always directed along the inter-aural axis irrespective of whether the stimulus was applied binaurally or monaurally, whether the head was turned in yaw through 90 deg, whether the head was pitched down through 90 deg, or combinations of these manipulations. We conclude that a net canal signal evoked by GVS contributes to the medium-latency response whilst a net otolith signal does not make a significant contribution to either the short- or medium-latency responses.

Non-linear vector summation of left and right vestibular signals for human balance.

The left and right vestibular organs always transduce the same signal of head movement, and with natural stimuli can only be activated simultaneously. To investigate how signals from the left and right vestibular organs are integrated to control human balance we electrically modulated the firing of vestibular afferents from each labyrinth independently and measured the resulting balance responses. Stimulation of one side at a time (monaural) showed that individual leg muscles receive equal inputs from the two labyrinths even though a single labyrinth appeared capable of signalling 3-D head motion. To deduce principles of left-right integration, balance responses to simultaneous stimulation of both sides (binaural) were compared with responses to monaural stimuli. The binaural whole-body response direction was compatible with vector summation of the left and right monaural responses. The binaural response magnitude, however, was only 64-74% that predicted by the monaural sum. This probably reflects a central non-linearity between vestibular input and motor output because stimulation of just one labyrinth revealed a power law relationship between stimulus current and response size with exponents 0.56 (force) and 0.51 (displacement). Thus, doubling total signal magnitude either by doubling monaural current or by binaural stimulation produced equivalent responses. We conclude that both labyrinths provide independent estimates of head motion that are summed vectorially and transformed non-linearly into motor output. The former process improves signal-to-noise and reduces artifactual common-mode changes, while the latter enhances responses to small signals, all critical for detecting the small head movements needed to control human balance.

Resolving head rotation for human bipedalism.

Alignment of the body to the gravitational vertical is considered to be the key to human bipedalism. However, changes to the semicircular canals during human evolution suggest that the sense of head rotation that they provide is important for modern human bipedal locomotion. When walking, the canals signal a mix of head rotations associated with path turns, balance perturbations, and other body movements. It is uncertain how the brain uses this information. Here, we show dual roles for the semicircular canals in balance control and navigation control. We electrically evoke a head-fixed virtual rotation signal from semicircular canal nerves as subjects walk in the dark with their head held in different orientations. Depending on head orientation, we can either steer walking by "remote control" or produce balance disturbances. This shows that the brain resolves the canal signal according to head posture into Earth-referenced orthogonal components and uses rotations in vertical planes to control balance and rotations in the horizontal plane to navigate. Because the semicircular canals are concerned with movement rather than detecting vertical alignment, this result shows the importance of movement control and agility rather than precise vertical alignment of the body for human bipedalism.

Physiological stress markers during breath-hold diving and SCUBA diving.

This study investigated the sources of physiological stress in diving by comparing SCUBA dives (stressors: hydrostatic pressure, cold, and hyperoxia), apneic dives (hydrostatic pressure, cold, physical activity, hypoxia), and dry static apnea (hypoxia only). We hypothesized that despite the hypoxia induces by a long static apnea, it would be less stressful than SCUBA dive or apneic dives since the latter combined high pressure, physical activity, and cold exposure. Blood samples were collected from 12SCUBA and 12 apnea divers before and after dives. On a different occasion, samples were collected from the apneic group before and after a maximal static dry apnea. We measured changes in levels of the stress hormones cortisol and copeptin in each situation. To identify localized effects of the stress, we measured levels of the cardiac injury markers troponin (cTnI) and brain natriuretic peptide (BNP), the muscular stress markers myoglobin and lactate), and the hypoxemia marker ischemia-modified albumin (IMA). Copeptin, cortisol, and IMA levels increased for the apneic dive and the static dry apnea, whereas they decreased for the SCUBA dive. Troponin, BNP, and myoglobin levels increased for the apneic dive, but were unchanged for the SCUBA dive and the static dry apnea. We conclude that hypoxia induced by apnea is the dominant trigger for the release of stress hormones and cardiac injury markers, whereas cold or and hyperbaric exposures play a minor role. These results indicate that subjects should be screened carefully for pre-existing cardiac diseases before undertaking significant apneic maneuvers.

Postural control at the human wrist.

In our movements and posture, we always act against a physical load. A key property of any load is its elastic stiffness (K), which describes how the force required to hold it must change with position. Here we examine how load stiffness affects the ability to maintain a stable posture at the wrist. Loads having positive (like a spring) and negative stiffness (like an inverted pendulum) were created by varying the position of weights on multiarm rigid pendulum. Subjects (n = 9) held 15 loads (K = +/- 0.04, +/- 0.01 and 0 N m deg(-1) at mean torques of 0.2, 0.4 and 0.6 N m) still for 60 s. Residual wrist movement (sway) increased with mean torque and increased as stiffness became more negative. Large effects of load stiffness were seen at low frequencies (< 1.5 Hz) but not at higher frequencies that reflect load resonance and reflex activity. Subjects accurately perceived their postural sway while holding the loads but measured psychophysical thresholds showed that load stiffness was not perceived. We conclude that load stiffness, independent of force levels, affects the ability to control a load and that the postural control process relies on perception and volitional tracking rather than more automatic reflex pathways. Despite an awareness of their postural errors, we see no evidence for adaptation of postural control processes to compensate for changes in load properties. This is unlike the adaptation of feedforward control processes that produce targeted volitional movements when load properties are altered. We propose that postural control and movement control are fundamentally different neural processes.

Effects of spatial and nonspatial memory tasks on choice stepping reaction time in older people.

BACKGROUND: Studies comparing the effects of spatial and nonspatial secondary tasks on balance have produced conflicting results. However, in most of these studies the difficulty levels of the secondary tasks have not been matched. In this study, we compared the effects of carefully matched visuospatial (VS) and nonspatial (NS) secondary tasks on choice stepping reaction time (CSRT). METHODS: Forty-one older people (mean age 78.8 years) completed a CSRT test under five conditions: (i) no secondary task; (ii) an easy NS counting backward task; (iii) a difficult NS counting back task; (iv) an easy VS memory task; and (v) a difficult VS memory task. Response times and secondary task errors were measured for each condition. Participants also gave difficulty ratings for each secondary task. RESULTS: The difficult tasks were rated significantly more difficult than the easy tasks in both VS and NS conditions, and cognitive task errors were moderately correlated with perceived difficulty. A repeated-measure analysis of variance with planned contrasts revealed a significant effect of task type, with the VS condition slowing CSRT more than the NS condition. There was also a significant task difficulty effect with the more difficult tasks increasing CSRT. CONCLUSIONS: The findings suggest that VS cognitive tasks affect CSRT more so than do NS tasks. The visuospatial sketchpad appears to be specifically utilized for carrying out motor tasks necessary for preserving balance. Practical implications are that tasks that require visuospatial attention and memory may adversely influence balance control in older people.

Choice stepping response and transfer times: effects of age, fall risk, and secondary tasks.

BACKGROUND: Deterioration with age of physiological components of balance control increases fall risk. Avoiding a fall can also require higher level cognitive processing to select correct motor and stepping responses. Here we investigate how a competing cognitive task and an obstacle to stepping affect the initiation and execution phases of choice stepping reaction times in young and older people. METHODS: Three groups were studied: young persons (YOUNG: 23-40 years, n = 20), older persons with a low risk of falls (OLR: 75-86 years, n = 18), and older persons with a high risk of falls (OHR: 78-88 years, n = 22). Four conditions were examined: choice stepping, choice stepping + obstacle, choice stepping + working memory task, and choice stepping + working memory task + obstacle. Step response and transfer times were measured for each condition, in addition to hesitant stepping, contacts with the obstacle and errors made in the memory test. RESULTS: Older participant groups had significantly longer response and transfer times than the young group had, and the OHR group had significantly longer response and transfer times than the OLR group had. There was a significant Group x Secondary task interaction for response time (F(2,215) = 12.6, p <.001). With the memory task, response time was minimally affected in the YOUNG (7% increase, p =.11) but was slowed significantly in the OLR fallers (42% increase, p <.001) and more so in the OHR fallers (48% increase, p <.001). The obstacle had a small but significant effect on response time (9.4%) and a larger effect on transfer time (43.3%), with no differences among the groups. Errors in stepping, performing the secondary task and contacting the obstacle increased with age and fall risk. CONCLUSIONS: Compared with young people, older people, and more so those at risk of falling, have an impaired ability to initiate and execute quick, accurate voluntary steps, particularly in situations where attention is divided.

A structural equation model relating impaired sensorimotor function, fear of falling and gait patterns in older people.

Many falls in older people occur while walking, however the mechanisms responsible for gait instability are poorly understood. Therefore, the aim of this study was to develop a plausible model describing the relationships between impaired sensorimotor function, fear of falling and gait patterns in older people. Temporo-spatial gait parameters and acceleration patterns of the head and pelvis were obtained from 100 community-dwelling older people aged between 75 and 93 years while walking on an irregular walkway. A theoretical model was developed to explain the relationships between these variables, assuming that head stability is a primary output of the postural control system when walking. This model was then tested using structural equation modeling, a statistical technique which enables the testing of a set of regression equations simultaneously. The structural equation model indicated that: (i) reduced step length has a significant direct and indirect association with reduced head stability; (ii) impaired sensorimotor function is significantly associated with reduced head stability, but this effect is largely indirect, mediated by reduced step length, and; (iii) fear of falling is significantly associated with reduced step length, but has little direct influence on head stability. These findings provide useful insights into the possible mechanisms underlying gait characteristics and risk of falling in older people. Particularly important is the indication that fear-related step length shortening may be maladaptive.

A tactile stimulus applied to the leg improves postural stability in young, old and neuropathic subjects.

The purpose of this study was to determine whether the application of passive tactile cues to the lower limb could improve postural stability in healthy young controls, older people and people with diabetic peripheral neuropathy. Antero-posterior sway was measured with eyes open and closed in 10 healthy young subjects (mean age 27 years, 5 male, 5 female), 10 older subjects without diabetic peripheral neuropathy (mean age 88 years, 2 male, 8 female) and 10 subjects with diabetic peripheral neuropathy (mean age 65 years, 6 male, 4 female) while a small piece of Velcro attached to a flexible mount was applied to three different sites on the leg (ankle, calf, and knee). Across all conditions, the mean sway of the neuropathic subjects was 93% greater than for the young subjects and 11% more than the older subjects. On average, subjects swayed 10% more with the eyes closed than with the eyes open. Each stimulus reduced sway, but the effect increased approximately in proportion to the height of the stimulus above the ankles (ankle 7.6%, calf 13.5%, knee 20.1% reduction compared to the no stimulus condition). This experiment demonstrates that a passive stimulus applied to the skin of the leg, which provides sensory information about body movement, significantly reduces body sway during standing. This applies to older subjects and subjects with peripheral neuropathy as well as healthy young subjects. These results have implications for novel approaches for improving stability in people with peripheral sensory loss.

Perceptions of Speed and Risk: Experimental Studies of Road Crossing by Older People.

Crossing a road safely is a complex task requiring good sensorimotor function and integration of information about traffic speed, distances and one's own speed. Poor judgement through age-related sensorimotor or cognitive impairment or a predisposition to take risks could lead to errors with serious consequences. On a simulated road, 85 participants (age ≥70 years) were asked to cross in front of an approaching car with a clearance as small as considered safe in two conditions; (1) with nothing else to attend to (free crossing) and (2) with an additional ball-gathering task while waiting to cross (task crossing). Participants were categorised according to their crossing outcome (failed to cross, 'hit', exact, safe, cautious). Participants also performed two sub-studies; (1) the perception of the time-to-arrival of moving objects and (2) the perception of own gait speed. Physical and cognitive function and everyday risk-taking behaviour were also assessed. In free crossing, clearances varied but no participants were "hit" by the car. In task crossing, participants allowed smaller clearances and 10% of participants would have been hit while 13% missed the opportunity to cross altogether. Across a wide range of physical and cognitive measures, including perceived and actual gait speed, a consistent pattern was observed in the task crossing condition. The exact group performed best, the 'hit', safe and cautious groups performed less well while those who missed the opportunity (fail) performed worst. The exact group reported taking the greatest risks in everyday life whereas the remaining groups reported being cautious. In conclusion, we found older people with poorer perceptual, physical and cognitive function made inappropriate and risky decisions in a divided attention road-crossing task despite self-reports of cautious behaviour in everyday life.

Activation of human inspiratory muscles in an upside-down posture.

During quiet breathing, activation of obligatory inspiratory muscles differs in timing and magnitude. To test the hypothesis that this coordinated activation can be modified, we determined the effect of the upside-down posture compared with standing and lying supine. Subjects (n=14) breathed through a pneumotachometer with calibrated inductance bands around the chest wall and abdomen. Surface electromyographic activity (EMG) was recorded from the scalene muscles. Crural diaphragmatic EMG and oesophageal and gastric pressures were measured in a subset of six subjects. Quiet breathing and standard lung function manoeuvres were performed. The upside-down posture reduced end-expiratory lung volume. During quiet breathing, for the same inspiratory airflow and tidal volume, ribcage contribution decreased, abdominal contribution increased and transdiaphragmatic pressure swing doubled in the upside-down posture compared to standing (p<0.05). Despite this, crural diaphragm EMG was unchanged, whereas scalene muscle EMG was reduced by ∼half (p<0.05). Thus, the mechanical effect of an upside-down posture differentially affects inspiratory muscle activation.