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1. Hindlimb paw-shake responses were assessed before and after unilateral deafferentation (L3-S1) in chronic-spinal cats (n = 5), spinalized at the T12 level 1 yr earlier. Selected ankle flexor [tibialis anterior (TA)] and extensor [lateral gastrocnemius (LG)] and knee extensor [vastus lateralis (VL)] muscles were surgically implanted with chronic electromyographic (EMG) electrodes to determine mutable features of cycle characteristics and muscle synergies that are modulated by motion-dependent feedback as opposed to immutable features that are centrally programmed and not modulated by limb afference. 2. Paw-shake responses were difficult to elicit in the extensively deafferented hindlimb; this was true particularly during the first recovery weeks after deafferentation. By the end of the first month, however, brief responses of 1 or 2 cycles were commonly elicited in four of five cats, and responses of 3-7 cycles were common by the end of the second month in three of five cats. Initially, responses in the deafferented limb were elicited by stimuli applied to the dorsolateral thigh, an oval patch of skin innervated by intact S2 afferents. Over the 4-mo recovery period, however, the receptive field of the largely denervated skin expanded, and responses were also elicited by stimuli applied to the lateral aspect of the knee and shank, but usually not the paw. 3. In addition to fewer average cycles per response (5 vs. 10 cycles), paw shaking evoked in the deafferented hindlimb was characterized by longer-than-average cycle periods (124 vs. 98 ms), but the average cycle period varied widely among responses, ranging from 99 to 239 ms. Before deafferentation, the temporal organization of consecutive cycles was stereotypic; cycle periods increased linearly throughout a response. After deafferentation, however, there was no systematic relationship between cycle period and cycle number, and approximately 14% of the records with greater than or equal to 3 cycles were characterized by arhythmical sequences of EMG bursts. 4. At the ankle, LG burst duration was not altered by deafferentation, but TA onset and burst duration were affected. Before deafferentation, TA onset was invariant with respect to the beginning of the cycle, and burst duration increased linearly with cycle period. After deafferentation, however, TA onset was delayed, and the delay increased linearly with cycle period. Consequently, the TA burst duration was brief and unrelated to cycle period.(ABSTRACT TRUNCATED AT 400 WORDS)
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PMID:Mutable and immutable features of paw-shake responses after hindlimb deafferentation in the cat. 275 70

Neuromuscular patterns of scratching and the paw-shake response were studied in normal kittens from birth to postnatal day 60. Onset of both behaviors coincided with the development of secure weight-bearing posture and occurred on postnatal day 21 for scratching and postnatal day 26 for paw shaking. At onset, cycle periods for scratching (5-6 Hz) and paw shaking (8-10 Hz) were similar to that for adult cats, and EMG patterns were adult-like. The scratch cycle consisted of reciprocal flexor and extensor bursts of equal duration, while the shake cycle consisted of coactive knee extensor and ankle flexor bursts alternately active with ankle extensor bursts. The lack of scratching and paw shaking during the first 3 postnatal weeks and the adult-like EMG patterns at onset are consistent with the hypothesis that pattern-generating circuits within lumbosacral segments are available early in development but inhibited by the rostral neuraxis until postural control is sufficient to accommodate the response. To eliminate rostral inputs, including descending input critical for postural control, kittens were spinalized at the T12 level, and onset of paw shaking was accelerated. In kittens spinalized at birth, paw-shake onset occurred on postnatal day 14, while in kittens spinalized on postnatal day 14, onset occurred 48 h after spinalization. In all spinal kittens, however, knee extensor activity was disrupted and not normal by postnatal day 60. Mature neuromuscular patterns for scratching and paw shaking are available at onset of the behavior during normal development. Spinalization hastens the onset of paw shaking but the normal neuromuscular synergy is disrupted as well as the temporal structure of the multi-cycle response. Disruptions following spinalization may be due to altered development of spinal pattern generators or aberrant feedback from atypical hindlimb motions due to a retardation of hindlimb growth and an alteration of muscle contractile properties in spinal kittens.
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PMID:Neuromuscular patterns of stereotypic hindlimb behaviors in the first two postnatal months. III. Scratching and the paw-shake response in kittens. 334 32

The simultaneous control of the hindlimb paw-shake response and hindlimb walking at slow treadmill speeds (0.2-0.4 m/s) was examined in adult cats spinalized at the T12 level, 3-6 mo earlier. Paw shaking was elicited by either 1) application of adhesive tape or 2) water to the right hindpaw. To assess intralimb and interlimb coordination of the combined behaviors, activity from selected flexor and extensor muscles at the hip, knee, and ankle was recorded, and the kinematics of these joints were determined from high-speed cinefilm. When paw shaking was combined with hindlimb walking, the response in the stimulated limb was initiated during swing (F phase) of the step cycle. The onset of knee extensor activity provided the transition from the flexor synergy of swing to the mixed synergy of paw shake. At the end of the paw shake, an extensor synergy initiated the E-1 phase of swing, and the resultant joint motion was in-phase extension at the hip, knee, and ankle to lower the paw for contact with the treadmill belt. During the rapid (81 ms) paw-shake cycles, knee extensor and ankle flexor muscles exhibited single, coactive bursts that were reciprocal with coactive hip and ankle extensor bursts. This mixed synergy was reflected in the limb coordination, as knee flexion coincided with ankle extension and knee flexion coincided with ankle extension. Phasing of hip motions was variable, reflecting the role of the proximal in stabilization during paw shake (16). Although the number of paw-shake cycles combined during swing varied greatly from 2 to 14, average cycle periods, burst durations, and intralimb synergies were similar to those previously reported for spinal cats tested under conditions in which the trunk was suspended and hindlimbs were pendent (23, 27). For step cycles during which a long paw-shake response of 8-14 cycles occurred, swing duration of the shaking limb increased by 1 s, and during this prolonged interval, the contralateral hindlimb completed two support steps. Stance duration of the support steps was also prolonged. This adjustment maximized the duration of paw-contact and minimized any period of nonsupport by the contralateral hindlimb during paw shake. Completion of the paw-shake response was followed by either an alternating, or a nonalternating, gait pattern on the recovery steps. One spinal cat combined locomotion with short two-cycle paw-shake responses, and because the shortened response was limited primarily to the time ordinarily devoted to swing, interlimb adjustments were slight.(ABSTRACT TRUNCATED AT 400 WORDS)
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PMID:Simultaneous control of two rhythmical behaviors. II. Hindlimb walking with paw-shake response in spinal cat. 374 94

The present study was designed to examine the effects of an intraspinal transplantation of embryonic brainstem neurons on fictive motor patterns which can develop in hindlimb nerves of adult chronic spinal rats. Seventeen adult rats were spinalized at T8-9 level and, 8 days later, a suspension of embryonic cells obtained either from the raphe region (RR, n = 8) or from the locus coeruleus (LC, n = 9) was injected caudally (T12-13) to the cord transection. Eight control animals (control rats) were spinalized and injected with vehicle under the same conditions. One to three months later, the animals were decorticated and fictive motor patterns were recorded in representative hindlimb nerves. The data revealed that both control and grafted spinal rats could exhibit two distinctly different fictive motor patterns, one which could be associated with stepping and the other with hindlimb paw shaking. They further showed that following transplantation of embryonic RR or LC neurons the excitability of the spinal stepping generator was increased, whereas that of the spinal neural circuits which generate hindlimb paw shaking was not significantly affected. A histological analysis performed on the spinal cord segments below the transection revealed complete absence of serotonin and noradrenaline immunoreactivity in control spinal animals and, in both types of grafted rats, an extensive monoaminergic reinnervation with synaptic contacts between monoaminergic transplanted neurons and host interneurons and/or motoneurons. The possible mechanisms by which grafted monoaminergic neurons can influence the spinal motor networks are discussed.
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PMID:Fictive motor activities in adult chronic spinal rats transplanted with embryonic brainstem neurons. 854 78