Eur J Appl Physiol (1997) 76: 55 ± 61 Ó Springer-Verlag 1997 ORIGINAL ARTICLE Romuald Lepers á Andre X. Bigard á Jean-Pierre Diard Jean-FrancËois Gouteyron á Charles Y. Guezennec Posture control after prolonged exercise Accepted: 10 January 1997 Abstract The perturbations of equilibrium after prolonged exercise were investigated by dynamic posturography on nine well-trained subjects (four athletes and ®ve triathletes). A sensory organization test, where the platform and visual surround were either stable or referenced to the subject's sway with eyes open or closed, was performed before and after a 25-km run (average time 1h 44 min) by the nine subjects. In addition, the same test was performed on the ®ve triathletes only, before and after ergocycle exercise of identical duration (i.e. ergocycle time = running time). The results showed that the ability to maintain postural stability during con¯icting sensory conditions decreased after exercise, with some dierences depending on the kind of exercise. Sensory analysis revealed that the subjects made less eective use of vestibular inputs after running than after cycling (P < 0.05). Adaptation to prolonged stimulation of proprioceptive, vestibular and visual inputs had probably occurred in the integrating centres during exercise. This adaptation was maintained during the recovery period and could explain the postexercise balance disorders. Other mechanisms such as impairment of motor eerents or haemodynamic changes should not be excluded. Key words Equilibrium á Dynamic posturography á Sensory inputs á Prolonged exercise á Running Introduction The maintenance and control of posture and balance, whether under static or dynamic conditions, are essential requirements for physical and daily activities. The R. Lepers (&) á A.X. Bigard á C.Y. Guezennec DeÂpartement de Neurophysiologie, Institut de MeÂdecine AeÂrospatiale du Service de Sante des ArmeÂes, BP 73, F-91223 BreÂtigny-sur-Orge Cedex, France J.P. Diard á J.F. Gouteyron Centre Principal d'Expertise MeÂdicale du Personnel Navigant, Percy-Clamart, France maintenance of equilibrium is achieved by using orientation information derived from three independent sensory sources: somatosensory, vestibular and visual inputs. The aerent information is processed in the brainstem and cerebellum, and then motor commands are initiated. The three sensory systems all contribute to postural control, and damage to any of them, or to the brainstem or cerebellum, will in¯uence the overall output of the postural system. It has been shown that the contribution of individual sensory inputs to equilibrium can be quanti®ed by measuring the equilibrium adjustments of standing subjects deprived of vision and/or input from the support surface, the environment thus presenting the subjects with con¯icting visual, proprioceptive and vestibular stimuli (Nashner et al. 1982; Nashner 1983). In such an experiment, called ``dynamic posturography'', the various con¯icting sensory conditions are created by rotating the surface platform and/or the visual surround in proportion to the subject's postural sway, thus making these sources of sensory information inappropriate for the postural task. This method for assessing human balance and posture has been extensively used in clinical practice for assessing and dierentiating disturbances of vestibular, visual, and proprioceptive functions, as well as central co-ordination. Dynamic posturography has been found to be versatile in assessing equilibrium eects of anaesthesia (Gupta et al. 1991), acute and chronic eects of alcohol (Ledin and OÈdkvist 1991), and solvents (Ledin et al. 1991). Decreased postural stability has been studied in aged persons (Era and Heikkinen 1985; Straube et al. 1988; Teasdale et al. 1993), patients with vestibular de®cits (Nashner et al. 1982; Black and Nashner 1983), cerebral palsy (Nashner et al. 1983) or Parkinson's disease (Allum et al. 1988; Bloem 1992; Beckley et al. 1995) and many other diseases. Studies of human posture and equilibrium control in normal subjects and patients have provided much information on the capacity of the central nervous system to regulate posture. The decreased stability associated with postural sway can be related to 56 reduced peripheral sensibility in the visual, vestibular, or proprioceptive systems, and also to a defect or a slowing of the central integrative mechanisms responsible for con®guring the postural control system. From observations of the postural instability of runners after the end of a race, it would seem correct to assume that the equilibrium has been impaired by exhausting physical exercise. Sensory inputs are highly stimulated during a run: it has been demonstrated that muscle spindles, tendon organs, joint receptors and cutaneous aerents on the sole of the feet are activated at each stride (Merton 1964), the vestibular system is sensitive to each head acceleration (Kornhuber 1974), and the eyes are stimulated by the motion of the visual ®elds (Lestienne et al. 1977). Whereas the eects of long distance running on the cardiovascular and musculoskeletal systems are well known, the eects on the neurosensory system and therefore on the maintenance of equilibrium remain unknown. Nevertheless, there are some situations especially in sport where postural and balance requirements follow physical exercise (for example, cross-country skiing and shooting in the biathlon). To our knowledge, no study has yet quanti®ed the importance of postural instability after physical exercise. The present study was conducted in order to evaluate and quantify the equilibrium performance before and after long-distance running on well-trained subjects (triathletes and running specialists). To verify the importance of the movement of visual ®eld and the stimulation of face pressure receptors on the mechanisms of balance control, we also examined and quanti®ed the eects of a cycle ergometer exercise of similar duration and intensity. long-duration exercise and none had any history of vestibular or neurological disease. The experimental procedure was approved by the local Ethics Commission. Platform test procedures Procedures involved the use of an instrumented platform system (Equitest, Neurocom Int. Inc., Clackamas, Oregon, USA), which has been used in previous studies (Nashner et al. 1982; Ledin and OÈdkvist 1993; Beckley et al. 1995). The system included two movable support surfaces (one for each foot; each could independently move horizontally and rotate around an axis colinear to the ankle joints) and a movable visual surround (a 1-m square enclosure open at the back and bottom which rotated around an axis colinear to the ankle joints). Strain gauges within each platform measured the torsional forces and the total vertical force exerted by the foot resting upon its surface. Sensory organisation test Each subject stood quietly on the platform within the visual surround for three periods of 20 s (3-s rest inbetween) during six dierent test conditions (Fig. 1). During the ®rst three test conditions, the platform support surface remained ®xed with respect to the earth horizontal, but the visual surrounds were (1) ®xed with Methods Subjects Nine well-trained subjects whose physical characteristics are shown in Table 1, volunteered to participate in this study. Five of them were competitive triathletes, who trained on average 12±15 h a week, three times in each discipline: swimming, cycling and running. The four other subjects were competitive runners, who trained around 6±8 h a week. Every subject was familiarized with Table 1 Physical characteristics of the subjects Fig. 1 The six conditions of the sensory organisation test. During the ®rst three conditions 1, 2 and 3, the platform support surface was ®xed relative to earth horizontal. During the next three conditions 4, 5 and 6, the platform support surface was moved proportional to the subject's anteroposterior body sway. The three visual conditions were repeated in sequence, ®rst with the platform earth-®xed; second with the platform perceptually stabilized. The three visual conditions were repeated: 1 and 4 eyes open with a ®xed ®eld of vision 2 and 5 eyes closed, and 3 and 6 ®eld of vision moved in relation to body sway Subject Speciality Age (years) Height (cm) Body mass (kg) 25-km Time (h-min) Maximal O2 uptake cycling (ml á min)1 á kg)1) A B C D E F G H I Run Run Run Run Triathlon Triathlon Triathlon Triathlon Triathlon 44 27 32 29 41 23 40 27 44 172 178 175 180 183 174 176 183 171 60 66 61 59 73 55 66 74 60 1 1 1 1 1 1 1 1 1 ± ± ± ± 66 60 68 72 62 Mean SEM 34 3 176.8 1.4 63.8 2.2 1 h 44 0 h 03 h h h h h h h h h 43 32 51 50 41 57 34 40 51 65.6 2.1 57 respect to the earth vertical, (2) the subjects eyes were closed, and (3) in proportional to anteroposterior (AP) body sway (visual stabilization). It has been found that during this third condition, the visual enclosure sways in direct proportion to the changes in AP orientation, so that the relative orientation of the subject and his surround remains ®xed Nashner 1983. During the three other conditions (4), (5) and (6), the platform support surfaces rotated in proportional to AP body sway (support surface stabilization) and the ®rst three visual conditions were repeated. During these last three conditions, the relative orientation of the subject and the support surface was ®xed as has been described by Nashner (1971). During both support surface and visual stabilization procedures, the gain of surface motion relative to that of the body sway could be set at any level ranging from 1 (surface and body motions in direct proportion) to 2 (surface motions being double that of body motions). For each test an equilibrium score re¯ected the extent to which the subject's AP sway movements approached the limits of stability for the feet together stance, during each of the 20-s trials. A score between 0 (12.5° sway range, or falling) and 100 (no sway motion) was measured using the system of Nashner et al. (1990). The scores of the three successive 20-s trials of each test were averaged to give the test equilibrium score. The degree of ankle and hip movements were estimated by a strategy score (100: no horizontal AP forces, i.e. full ankle strategy 0: shear force of 110 N estimated to be the maximum possible, i.e. full hip strategy; Horak and Nashner 1986). Protocol The sensory organisation test (SOT) (Fig. 1) was performed on each subject, three times (runner specialists) or ®ve times (triathletes). A trial sequence was always performed a few days before the day of the test run, to familarize the subjects with the procedure and to reduce the learning eects. The other sequences were carried out just before (PRE) and immediately after (POST) the long-distance run and cycle exercise. Because disorientation and instability were possible during the assessment of fatigued subjects, sensory conditions were imposed to increase the diculty during the SOT. The ®rst three SOT quanti®ed the performance of each subject during three dierent visual conditions (normal, eyes closed and stabilized vision) when they were provided with a ®xed support surface. The three tests were repeated for the three dierent visual conditions, except that then the subjects stood with the support surface stabilized. Because the subjects showed good balance (preliminary test), stabilization gains during these trials were set at 2. Sensory analysis was accomplished by computing sensory ratios among the average equilibrium scores on speci®c pairs of sensory test conditions, as shown in Table 2. All the subjects ®rst performed the long distance run and the triathletes, at least 1 week later the cycle exercise. All the tests were performed in the morning. Long distance run The nine subjects were invited to run, at a speed close to their personal record, for 25 km outside on a generally ¯at route which had been measured. The subjects were followed during the entire run by Table 2 Sensory analysis. For de®nitions of conditions 1±6 see Fig. 1 a cyclist, who gave them information about distance and time. Water was provided on demand by the cyclist without restriction. Stationary cycle exercise Because the competitive runners were not trained cyclists, the second part of the experiment was conducted only with the triathletes. Triathletes were asked to cycle on an ergometer (Orion, Toulouse, France) at a power corresponding to approximately 65%±70% of their maximal oxygen uptake (determined during previous laboratory incremental tests) for a time equal to that of their 25-km run. Statistics Statistical dierences between PRE and POST conditions of the same exercise (run or cycle) and between POST conditions (run versus cycle) were determined by the nonparametric Mann Whitney test, for each sensory condition of SOT and for each sensory ratio of the sensory analysis. Results The subjects ran 25-km in times ranging from 1 h 32 min to 1 h 57 min with an average of 1 h 44 min (Table 1). Dynamic posturography after the 25-km run Figure 2a and b illustrates the average (and SEM) equilibrium scores and strategy scores during the six dierent sensory conditions, before and after the 25-km run for the nine subjects, respectively. The equilibrium scores, which re¯ected the subject's ability to maintain his balance, decreased after the run, except in condition 1. In the two other ®xed support surface conditions, stability performance was lower after the run than before. Strategy scores remained high during these three conditions, suggesting that the ankle strategy prevailed during ®xed support surface conditions. During the three stabilized support surface conditions, stability performance was lower and particularly altered in conditions 5 and 6 after the run. Equilibrium scores decreased signi®cantly postexercise (P < 0.05) in the stabilized surface condition 4. In condition 5, the average equilibrium score was reduced from 71 (3.6° body sway angle range) before the run to 52 (6° body sway angle range) after the run. In conditions 5 and 6 lower performance and greater variability resulted from the loss of balance (i.e. score 0) of one subject in 5 and two Ratio Name Ratio Pair Somatosensory Condition Condition Condition Condition Condition Condition Condition Condition Visual Vestibular Visual Preference Question 2 1 4 1 5 1 (3+6) (2+5) Does sway increase when visual cues are removed? Does sway increase when somatosensory cues are inaccurate? Does sway increase when visual cues are removed and somatosensory cues are inaccurate? Do inaccurate visual cues result in increased sway compared to no visual cues? 58 Fig. 2a Comparison of equilibrium scores before (PRE ) and after (POST ) a 25-km run (n=9). Signi®cance level *P < 0.05 b Comparison of strategy score before (PRE ) and after (POST ) a 25-km run (n = 9). Signi®cance level: *P < 0.05. For de®nitions of conditions 1±6 see Fig. 1 subjects in 6. Under the three stabilized support surface conditions, strategy scores were lower after exercise, re¯ecting a preferential use of hip strategy to control balance. This decrease was signi®cantly dierent in conditions 4 (P < 0.05) and 5 (P < 0.05) after the run. Comparison of dynamic posturography after ergometer exercise versus the run Figure 3a and b illustrates respectively the average and SEM equilibrium scores and strategy scores during the six dierent sensory conditions before and after the 25-km run and the ergocycle exercise for the ®ve triathletes. During ®xed support surface conditions, equilibrium scores were very similar either after the run or after cycling in 1, but present in 2 and 3 condition values which were lower postrun than after the ergometer exercise. Strategy scores did not show signi®cant dierences during the ®xed support surface conditions. During stabilized support surface conditions, the alterations of performances were similar with ®xed (4) or stabilized vision (6) either after the run or the cycle exercise. A dierence appeared nevertheless during the eyes closed condition when the scores tended to be substantially lower after running than after cycling, even though it was not statistically signi®cant. Strategy scores decreased similarly, being independent of the kind of exercise during stabilized support surface conditions. Sensory analysis Figure 4 illustrates the changes in sensory ratios. The sensory ratio of condition 2:1 remained unchanged 59 Fig. 3a Comparison of equilibrium scores before (PRE ) and after (POST ) a 25-km run and the same duration ergometer exercise (n = 5). Signi®cance levels *P < 0.05, **P < 0.01. b Comparison of strategy scores before (PRE ) and after (POST ) a 25-km run and the same duration ergometer exercise (n = 5). Signi®cance level *P < 0.05, **P < 0.01. For de®nitions of conditions 1±6 see Fig. 1 whereas the sensory ratios for both conditions 4:1 and 5:1 decreased postexercise. In the absence of a stable surface, the subjects did not make eective use of either vestibular or visual inputs after prolonged exercise. Vestibular input seems to have been more aected by the run than by the ergometer exercise (P < 0.05). Visual preference values remained unchanged postexercise. Discussion Fig. 4 Comparison of sensory ratios before (PRE ) and after (POST ) a 25-km run (n = 9) and the same duration ergometer exercise (n = 5). SOM somatosensory (condition 2/condition 1), VIS visual (condition 4/condition 1), VEST vestibular (condition 5/condition 1), VIS PREF visual preference (condition 3+6/condition 2+5). Signi®cantly dierent from PRE-exercise condition aP < 0.05, bP < 0.01. Signi®cantly dierent from POST-RUN condition cP<0.05 The results showed that the subjects' abilities to maintain their balance under con¯icting sensory conditions were altered by prolonged exercise with some dierences between the run and ergometer exercise. Estimation of human balancing ability by tradition has been limited to static test conditions, i.e. the person standing on a stable 60 support surface during the test. Static posturography limits the possibilities of the investigator to study external in¯uences. The eects of vision (Romberg test) may be tested easily, but the role of the concurrent systems were not studied. The SOT procedure represents a substantial improvement in the study of the equilibrium system as a control system. It must also be remembered that this method has drawbacks; it is not speci®c to any of the systems involved, it is also heavily in¯uenced by motivation and concentration. Whether a learning eect interfered with the equilibrium performances during the test may be questioned. Before the experimental measurements, one sequence was devoted to familarizing the subjects with the SOT procedure. However, no statistical dierences appeared between the equilibrium and strategy scores in the different sensory conditions of the prerun and the prebike situations. Moreover, the subjects performed their best scores during the second SOT procedure i.e. prerun test. It may thus be hypothesized that a learning eect did not signi®cantly aect the results. The post-tests were performed immediately after the end of the exercise, it corresponded then to a period of transition from exercise perturbation to a rest condition. The maintenance of equilibrium was appreciated here during the recovery time when the sensory stimulations of the exercise had stopped. However, it may be hypothesized that the mechanisms of postural regulation were impaired by the previous exercise. This observation raises the problem of the dierent mechanisms which could explain this impairment of balance control. In the sensory conditions 5 and 6 of SOT, when both the support surface and visual references were made simultaneously inaccurate, leaving vestibular input as the only potentially accurate orientation reference for postural control, the regulation of erect stance was more dicult after exercise, especially after the run. The increased sway under these conditions was typical of patients with reduced vestibular function (see Black and Nashner 1983). During the run, a continuous stimulation of the otholitic system, which is sensitive to linear head acceleration, would have occurred. In response to the prolonged stimulation during the run, the integrator centres of vestibular information could have decreased their sensibility threshold. Consequently, the vestibular omission resulting from an adaptation to running movement, which probably persisted during the beginning of the recovery, could in part explain the perturbations of postural control postexercise. In the same way, we cannot exclude an adaptation to the visual information by the integrator centres. Visually guided behaviour such as locomotion and postural stabilization have been shown to depend on visual information mediated by the peripheral visual inputs (Lestienne et al. 1977). During the run, in contrast to a stationary cycle exercise, the visual input was continuously stimulated by the moving ®eld of vision. Visual movement by itself could have given the perception of body sway and caused postural compensations, even when somatosensory and vestibular information did not signal the sway. Such a prolonged stimulation of the visual input, and the adaptation which resulted, could also have in¯uenced the maintenance of balance during the post running equilibrium test. The third input which was highly stimulated during such prolonged exercise was the somatosensation. Under normal circumstances, it has been demonstrated that an individual is more reliant on somatosensation than on visual inputs in correcting for body sway (Nashner and Berthoz 1978). During running, eccentric contractions alternate with concentric ones and muscle damage does occur. The tendons, the joint receptors and the cutaneous mechanoreceptors of the sole of the foot are then greatly and repetitively stimulated. All these receptors play an important role in the maintenance of balance. A loss of somatosensory information from the lower limbs, resulting from a disease or an experimental manipulation, has been shown to induce postural control abnormalities (Diener et al. 1984; Horak et al. 1990). Moreover, muscular fatigue associated with local glycogen depletion, metabolite accumulation and muscle damage probably occured in the ankle extensors and ¯exors during the run. A decrease of the eector system eciency due in part to muscle fatigue, and changes in the proprioceptive information and/or in their integration could probably impair the postural regulation loop. This hypothesis has been supported by previous observations showing that stretch-re¯ex sensitivity was reduced after exhausting exercise (Hortobagyi et al. 1991; Nicol et al. 1996). This phenomenon can be attributed to several factors associated with metabolic fatigue, neuromuscular spindle sensitivity or muscle damage. In our conditions, we did not make a direct measurement of muscle fatigue so that it was not possible to relate postural control changes with fatigue. Further studies are needed to obtain a reliable index of fatigue which could be compared to postural changes. The changes of postural strategies postexercise, especially the increase of hip movements to control postural stability, could be attributed to such an adaptative process. The dierences in equilibrium scores in some sensory conditions of SOT between the postrun and the postcycling situations would suggest that running exercise could have speci®c eects on equilibrium maintenance. The vestibular system, in particular the otholitic organs, were less stimulated during the ergometer exercise than during the run. It should nevertheless be pointed out that the sensitivity threshold of otholitic organs to linear accelerations was very low (5 cm á s)2), and the head oscillations during the ergometer exercise should be taken into consideration, but to a lesser extent. The ergometer exercise also suppressed the moving ®eld of vision thus reducing the stimulations of the visual input. Finally, the lower leg muscle recruitment and the stimulations of joint and tendon receptors, but also of cutaneous mechanoreceptors, were dierent, with smaller mechanical constraints. Such dierences in proprioceptive requirements between running and cy- 61 cling could explain the lower equilibrium score obtained after the run compared to cycling in condition 3 of SOT ± ®xed support surface, stabilized vision ± when the central system would have used essentially the proprioceptive information to regulate balance. The changes of sensory information integration postexercise are nevertheless not the only hypothesis to explain the equilibrium perturbations. During exercise, the peripheral blood volume can only be returned to the central circulation by continual activation of the skeletal muscle pump, as occurs with running or cycling. Therefore, the sudden stop in exercise would have immediately promoted peripheral blood pooling, which could have transiently perturbated vestibular vascularization in the ®rst minutes of the test. Thus, as has been suggested a perturbation in blood ¯ow parameters at the vestibular level could perhaps impair the sensory input even if the cerebellar blood ¯ow seems to be preserved during exercise (Jorgensen et al. 1992). To test this hypothesis, an audiogram with detection of auditory thresholds, for example, should be performed after exercise. Haemodynamic changes resulting from progressive dehydration, as often occurs during prolonged exercise, should also be taken into consideration. Conclusion The main result of this study was that the subjects' ability to maintain their balance in a dynamic environment was altered by prolonged exercise. Such alterations probably resulted from an adaptation by the integrator centres to the hyperstimulation of sensory inputs during exercise, which persisted during the recovery period. Nevertheless other parameters, such as impairment of motor eerents and haemodynamic changes, could also have been involved in the disturbance of equilibrium postexercise. The lesser stimulation of vestibular and visual inputs during the cycle ergometer exercise than during the run could explain the lesser alteration of postural control after the cycle ergometer exercise. Acknowledgements The authors wish to thank Mr. G. Porcheron for the English language revision of the manuscript. References Allum JHJ, Keshner EA, Honegger F, Wuetrich R (1988) Disturbance of posture in Parkinson's disease. In: Amblard B, Berthoz A, Clarac F (eds) Posture and gait. Development, adaptation and modulation. Elsevier, Amsterdam, pp 245±257 Black FO, Nashner LM (1983) Postural control in four classes of vestibular abnormalities. In: Igarashi, Black FO (eds) Vestibular and visual control on posture and locomotion equilibrium. 7th International Symposium International Society of Posturography, Houston, Texas. Karger, Basel, pp 271±281 Beckley DJ, Panzer VP, Rembler MP, Ilog LB, Bloem BR (1995) Clinical correlates of motor performance during paced postural tasks in Parkinon's disease. J Neurol Sci 132:133±138 Bloem BR (1992) Postural instability in Parkinon's disease. Clin Neurol Neurosurg [Suppl] 94:541±545 Diener HC, Dichgans J, Guschlbauer B, Man H (1984) The signi®cance of proprioception on postural stabilization as assessed by ischemia. Brain Res 296:103±109 Era P, Heikkinen E (1985) Postural sway during standing and unexpected disturbance of balance in random samples of men of dierent ages. J Gerontol Med Sci 40:287±295 Gupta A, Ledin T, Larsen LE, Lennmarken C, OÈdkvist LM (1991) Computerized dynamic posturography: a new method for the evaluation of postural stability following anesthesia. Br J Anaesth 66:667±672 Horak FB, Nashner LM (1986) Central programming of postural movements: adaptation to altered support surface con®gurations. J Neurophysiol 55:1369±1381 Horak FB, Nashner LM, Diener HC (1990) Postural strategies associated with somatosensory and vestibular loss. Exp Brain Res 82:167±177 Hortobagyi T, Lambert NL, Kroll WP (1991) Voluntary and re¯ex responses to fatigue with stretch-shortening exercise. Can J Sports Sci 6:142±150 Jorgensen LG, Perko M, Secher NM (1992) Regional cerebral artery mean velocity and blood ¯ow during dynamic exercise in humans. J Appl Physiol 73:1825±1830 Kornhuber HH (1974) Handbook of sensory physiology: the vestibular system, vol. VI. Springer, Berlin Heidelberg New York Ledin T, OÈdkvist LM (1991) Eect of alcohol measured by dynamic posturography. Acta Otolaryngol [Suppl] Stockh 481:576±581 Ledin T, OÈdkvist LM (1993) Eects of the increased inertial load in dynamic and randomized perturbed posturography. Acta Otolaryngol (Stockh) 113:249±252 Ledin T, Jansson E, MoÈller C, OÈdkvist LM (1991) Chronic toxic encephalopathy investigated by dynamic posturography. Am J Otolaryngol 12:96±100 Lestienne F, Berthoz A, Soetching J (1977) Postural readjustments by linear motion of visual scenes. Exp Brain Res 28:363±384 Merton PA (1964) Human position sense and sense of eort. Symp Soc Exp Biol 18:387±400 Nashner LM (1971) A model describing the vestibular detection of body sway motion. Acta Otolaryngol (Stockh) 72:429±436 Nashner LM (1983) Analysis of movement control in man using the movable platform. In: Desmedt JE (ed) Motor control mechanism in health and disease. Raven, New York, pp 607± 619 Nashner LM, Berthoz A (1978) Visual contribution to rapid motor responses during posture control. Brain Res 150:403±407 Nashner LM, Black FO, Wall C (1982) Adaptation to altered support and visual conditions during stance: patients with vestibular de®cits. J Neurosci 2:536±544 Nashner LM, Shumway-Cook A, Marin O (1983) Stance posture control in select groups of children with cerebral palsy: de®cits in sensory organisation and muscular coordination. Exp Brain Res 49:393±418 Nashner LM, Shuppert CL, Horak FB (1990) Organization of posture controls: an analysis of sensory and mechanical contraints. Prog Brain Res 80:411±418 Nicol C, Kmoi PV, Horita T, KyroÈlaÈinen H, Takala TES (1996) Reduced stretch-re¯ex sensitivity after exhausting stretchshortening cycle exercise. Eur J Appl Physiol 72:401±409 Straube A, BoÈtzel K, Hawken M, Paulus W, Brandt T (1988) Postural control in the elderly: dierential eects of visual, vestibular and somatosensory input. In: Amblard B, Berthoz A, Charac F (eds) Posture and gait. Development, adaptation and modulation. Elsevier, Amsterdam, pp 105±114 Teasdale N, Bard C, Larne J, Fleury M (1993) On the cognitive penetrability of posture control. Exp Aging Res 19:1±13
© Copyright 2026 Paperzz