Evolution of plantigrade gait: is there a neuronal correlate?
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Biomedical subjects
Publications and source records attributed to H Forssberg.
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The authors examined the force regulation during prehension of 12 children with cerebral palsy (CP) and an age-matched control group. The control group appropriately scaled the isometric force output in one force-rate pulse toward the weight of the previously lifted object, but children with CP exhibited stepwise increases in force in which the force amplitudes were not influenced by the object's weight. When the object was held in the air, most children with CP were able to modify their force output according to the object's weight, using sensory feedback, but their forces were more variable than those of the control group and less influenced by the object's weight. Although children with CP have disturbed sensory feedback--likely to lead to an indistinct internal representation of an object's physical properties--their main disability stems from impaired central co-ordination of motor activity.
1. Anticipatory postural adjustments were studied in children (6-14 yr of age) walking on a treadmill while pulling a handle. Electromyographs (EMGs) and movements were recorded from the left arm and leg. 2. Postural activity in the leg muscles preceded voluntary arm muscle activity in all age groups, including the youngest children (6 yr of age). The latency to both leg and arm muscle activity, from a triggering audio signal, decreased with age. 3. In older children the latency to both voluntary and postural activity was influenced by the phase of the step cycle. The shortest latency to the first activated postural muscle occurred during single support phase in combination with a long latency to arm muscle activity. 4. In the youngest children, there was no phase-dependent modulation of the latency to the activation of the postural muscles. The voluntary activity was delayed during the beginning of the support phase resulting in a long delay between leg and arm muscle activity. 5. The postural muscle activation pattern was modified in a phase-dependent manner in all children. Lateral gastrocnemius (LG) and hamstring muscles (HAM) were activated during the early support phase, whereas tibialis anterior (TA) and quadriceps (Q) muscles were activated during the late support phase and during the swing phase. However, in the 6-yr-old children, LG was also activated in the swing phase. LG was activated before the HAM activity in the youngest children but after HAM in 14-yr-old children and adults. 6. The occurrence of LG activity in postural responses before heel strike suggests an immature (nonplantigrade) gating of postural activity.(ABSTRACT TRUNCATED AT 250 WORDS)
Neuronal multiplication occurs mainly from the 10th to the 20th gestational weeks, after which probably no new nerve cells are formed, though neuronal arborisation and the formation and re-organisation of synapses continues until adulthood. An intriguing question is how the blueprint for the formation of about 100 billion nerve cells and their dendrites and synapses can be contained in the human genome. Environmental factors are probably crucially involved in the development of the brain, particularly from the later stages of gestation onwards. Somatosensory functions are developed at an early stage. The pain threshold is assumed to be lower in the fetus than in the adult. In man, general fetal movements appear from the 8th gestational week, and more complex movements such as sucking, swallowing and breathing during the 10th-12th gestational weeks. These movements are generated by neuronal networks, and seem to occur spontaneously without any sensory stimulation. At birth there is a general excitation of the infant, and the neonate is awake and aroused. Studies in rats have shown a multifold increase in noradrenaline turnover to occur in the brain stem at birth, which is assumed to be related to the arousal of the newborn. Recent studies have shown that there is a switch-on of various excitatory neuropeptide genes at birth, and an increase in noradrenaline in the locus coeruleus which is assumed to be the arousal centre.
The coordination of manipulative forces was examined while children and adults repeatedly lifted a small object between the thumb and index finger. Grip force, load force (vertical lifting force), grip force rate and the vertical position of the test object were continuously measured. In adults, the force generation was highly automatized and was nearly invariant between trials. After a preload phase in which the grip was established, the grip and load forces increased in parallel under isometric conditions until the load force overcame the force of gravity and the object started to move. During this loading phase, the force rate profiles were essentially bell shaped and single peaked, suggesting that the force increases were programmed as one coordinated event. Children below the age of two exhibited a prolonged preload phase and a loading phase during which the grip and load forces did not increase in parallel. A major increase in grip force preceded the increase in load force, and at the start of the loading phase, the grip force was usually several Newtons (N). The force rate profiles were multi peaked with stepwise force increases most likely allowing peripheral feedback to play an important role in the control of the forces. After the age of two, the grip force increased less during the preload phase. The loading phase was more regularly characterized by a parallel increase of the grip force and load force and the duration of the various phases decreased. The older children programmed the forces in one force rate pulse indicating the emergence of an anticipatory strategy. Yet, the mature coordination of forces was not fully developed until several years later.(ABSTRACT TRUNCATED AT 250 WORDS)
A size-weight illusion was used to examine the role of visual cues in the programming of manipulative forces during the lifting of test objects utilizing the precision grip. Three different boxes of equal weight and unequal size were lifted. These were equipped with an instrumented grip handle to measure the employed grip force, load force (vertical lifting force), force rates and vertical movement. All fifteen subjects participating in the study reported that the smallest box was the heaviest, which is consistent with size-weight illusion predictions. However, the rate of increase of the isometric grip and load forces initially during the lift, the peaks of the grip and load force and the vertical acceleration were all found to increase with the box size. Thus, despite the conscious perception indicating a heavier weight for the small object, the motor program was scaled for a lighter weight. Yet, no differences were found in grip force during the static phase of the lift, where weight related information was apparently available via sensory feedback. Previous studies have reported that the programming of the precision grip is based on somatosensory information gained during previous lifts (Johansson and Westling 1984, 1988a, b). The present study suggests that visual cues are integrated in the programming of manipulative forces during precision grip.
Recent evidence for the use of visual cues in the programming of the precision grip has been given by Gordon et al. (1991). Visually invoked size-related information influenced the physical forces used to produce a lift, even when it was not consistent with other sensory information. In the present study, blind-folded subjects were required to feel the size of an object by haptic exploration prior to lifting it. Two boxes of equal weight and unequal size were used for the lift objects and were attached to an instrumented (grip) handle. Grip force and load force, their rates, and the vertical movement of the object were measured. Most subjects reported that the small box was heavier, which is consistent with size-weight illusion predictions. However, peak grip force, grip force rate, peak load force, and load force rate were greater for the large box when the boxes were randomly presented, but not when the same boxes were lifted consecutively. If subjects did not feel the box prior to a lift, these parameters were scaled in between those normally employed for the large and small box. Most subjects apparently programmed the parallel increase of the grip and load force during the loading phase as one force rate pulse. This represented a "target strategy" in which an internal neural representation of the objects weight determined the actual target parameter (i.e. just enough force required to overcome gravity). The other subjects exhibited a slower stepwise increase in grip and load force rate. The subjects choosing this "probing strategy" did not scale the force parameters differently for the two boxes.(ABSTRACT TRUNCATED AT 250 WORDS)
Evidence has recently been given by Gordon et al. (1991a, b) for the use of visually and haptically acquired information in the programming of lifts with the precision grip. The size-related information influences the development of manipulative forces prior to the lift-off, and the force output for larger objects is adjusted for a heavier weight even if the weight of the objects is kept the same. However, the size influences on the force output were small compared to the relative effects of the expected weight in previous trials (Johansson and Westling 1988). In the present study, both the size and weight of objects were changed between consecutive lifts to more fully determine the strength of visual size cues. During most trials, the size and weight covaried (i.e. the weight was proportional to the volume). However, in some trials, only the size was switched while the weight was kept the same to create a mismatch between the size and weight. The forces were still appropriately scaled towards an expected weight proportional to the volume of the object. It was concluded that visual size cues are highly purposeful. The effects were much larger than previously reported and were similar in magnitude to the effects based upon the expected weight. Thus, the small effects reported in the previous experiments may have been a result of conflicting "size-weight" information.
Neonatal neuronal exuberance and its retention following neonatal brain damage have been demonstrated in a number of species but not in humans. The purpose of the present ongoing study is to determine if there is any evidence of neonatal neuronal exuberance and its retention following damage to the CNS in the human. Of equal concern is the determination of the neurological mechanisms underlying abnormal movement and reflex development in children with cerebral palsy.
Electromyographic and kinematic data were collected during treadmill locomotion by normal infants and infants with cerebral palsy. Locomotor patterns of the infants with cerebral palsy were similar to those of normal infants during the stage of supported locomotion, but as they matured, some of the characteristics of the infant stepping pattern, such as synchronous muscle activity with excessive muscular co-contraction and short-latency reflexes at foot contact, were retained. The normal plantigrade features of adult gait did not develop in these children.
The coordination of manipulatory forces during prehension was studied in 12 children with cerebral palsy (CP) and compared with that of controls. The results indicated that coupling of grip force and load force does not develop in children with CP. These children's force development increased in stages, with an early onset of excessive grip force. They did not use anticipatory control of the isometric force development during the load phase. Prolonged delays between successive phases indicated inefficient sensory feedback during the movement. The early onset of grip force and the over-all high force employment may compensate for the lack of anticipatory control and inefficient sensorimotor integration.
1. The ability of the CNS to coordinate several motor tasks was studied in humans walking on a treadmill while pulling on a handle. Subjects were instructed to respond to an audio signal that was presented in different phases of the step cycle. Electromyograph (EMG) and movements were recorded from the left arm and leg. 2. The activity of the arm muscle was preceded by postural activity in the leg muscles. The pattern of the anticipatory postural activity differed in the various phases of the step cycle. Lateral gastrocnemius and hamstring muscles were activated during responses occurring in the early support phase whereas tibialis anterior and quadriceps muscles were activated when the pull was exerted during the late support phase and during the swing phase. In the middle of the support phase the combination of both muscle activity was present. 3. The temporal sequencing and the spatial distribution of the anticipatory muscle activity changed gradually. Early during the support phase the hamstring muscles were activated before the gastrocnemius muscle, whereas the order was reversed during midstance. The EMG amplitude of the hamstring and gastrocnemius muscles was largest in the beginning of the support phase and then gradually decreased, whereas the amplitude of the tibialis anterior and quadriceps muscles increased during the later parts of the support phase. 4. The anticipatory responses to pulls exerted during the first part of the support phase reduced the ankle flexion during the single support phase.(ABSTRACT TRUNCATED AT 250 WORDS)
Neuronal multiplication occurs mainly from the 10th to the 20th gestational weeks, after which probably no new nerve cells are formed, though neuronal arborization and the formation and re-organisation of synapses continues until adulthood. An intriguing question is how the blueprint for the formation of about 100 billion nerve cells and their dendrites and synapses can be contained in the human genome. Environmental factors are probably crucially involved in the development of the brain, particularly from the later stages of gestation onwards. Somatosensory functions are developed at an early stage. The pain threshold is assumed to be lower in the fetus than in the adult. In man, general fetal movements appear from the 8th gestational week, and more complex movements such as sucking, swallowing and breathing during the 10th-12th gestational weeks. These movements are generated by neuronal networks, and seem to occur spontaneously without any sensory stimulation. At birth there is a general excitation of the infant, and the neonate is awake and aroused. Studies in rats have shown a multifold increase in noradrenaline turnover to occur in the brain stem at birth, which is assumed to be related to the arousal of the newborn. Recent studies have shown that there is a switch-on of an excitatory neuropeptide genes at birth, and an increase in noradrenaline in the locus coeruleus which is assumed to be the arousal centre.
Experiments were performed to determine whether spinal and supraspinal components of reciprocal inhibition (a neural mechanism responsible for the prevention of muscular co-ordination during voluntary movement) were present in groups of non-disabled children and children with cerebral palsy. Changes in the gastrocnemius-soleus H reflex were examined during voluntary dorsiflexion and plantarflexion of the ankle and during a vibration applied to the anterior tibial tendon. The results indicate that children with cerebral palsy have impairments in reciprocal inhibition, both before and during voluntary movement. These deficits, which involve damage to supraspinal centres, contribute to their inability to perform smooth, co-ordinated movements.
The kinematics of the pelvis and thorax are important in gait studies since their movement patterns are closely related to gait efficiency and 'smoothness' of locomotion. The purpose of this study was to identify features of normal gait patterns for later comparisons with pathological and developmental gait patterns. A two camera SELSPOT system interfaced with an HP1000 minicomputer was used to obtain three-dimensional kinematic/temporal data for the pelvis and thorax. Data from treadmill walking of eight adults were used for within subject (at different speeds) analyses. The analyses revealed a very complex pattern with a set of breakpoints which was consistent over all subjects. Some features were invariant over a range of walking speeds although the total range of motion changed considerably.
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This study examines the interactions between anteroposterior postural responses and the control of walking in human subjects. In the experimental paradigm, subjects walked upon a treadmill, gripping a rigid handle with one hand. Postural responses at different phases of stepping were elicited by rapid arm pulls or pushes against the handle. During arm movements, EMG's recorded the activity of representative arm, ankle, and thigh segment muscles. Strain gauges in the handle measured the force of the arm movement. A Selspot II system measured kinematics of the stepping movements. The duration of support and swing phases were marked by heel and toe switches in the soles of the subjects' shoes. In the first experiment, subjects were instructed to pull on the handle at their own pace. In these trials all subjects preferred to initiate pulls near heel strikes. Next, when instructed to pull as rapidly as possible in response to tone stimuli, reaction times were similar for all phases of the step cycle. Leg muscle responses associated with arm pulls and pushes, referred to as "postural activations," were directionally specific and preceded arm muscle activity. The temporal order and spatial distribution of postural activations in the muscles of the support leg were similar when arm pull movements occurred while the subject was standing in place and after heel strike while walking. Activations began in the ankle and radiated proximally to the thigh and then the arm. Activations of swing leg muscles were also directionally specific and involved flexion and forward or backward thrust of the limb. When arm movements were initiated during transitions from support by one leg to the other, patterns of postural activations were altered. Alterations usually occurred 10-20 ms before hell strikes and involved changes in the timing and sometimes the spatial structure of postural activations. Postural activation patterns are similar during in-place standing and during the support phase of locomotion. Walking and posture control appear to be separately organized but interrelated activities. Our results also suggest that the stepping generators, not peripheral feedback time locked to heel strikes, modulate postural activation patterns.