Forms of Forward Quadrupedal Locomotion. II. A Comparison of Posture, Hindlimb Kinematics, and Motor Patterns for Upslope and Level Walking

1998 ◽  
Vol 79 (4) ◽  
pp. 1687-1701 ◽  
Author(s):  
Patricia Carlson-Kuhta ◽  
Tamara V. Trank ◽  
Judith L. Smith
Keyword(s):  
Stance Phase ◽  
Emg Activity ◽  
Similar Data ◽  
Flexor Muscle ◽  
Motor Patterns ◽  
Quadrupedal Locomotion ◽  
Hindlimb Muscles ◽  
Level Walking ◽  
Ankle Joints ◽  
Hindlimb Kinematics

Carlson-Kuhta, Patricia, Tamara V. Trank, and Judith L. Smith. Forms of forward quadrupedal locomotion. II. A comparison of posture, hindlimb kinematics, and motor patterns for upslope and level walking. J. Neurophysiol. 79: 1687–1701, 1998. To gain insight into the neural mechanisms controlling different forms of quadrupedal walking of normal cats, data on postural orientation, hindlimb kinematics, and motor patterns of selected hindlimb muscles were assessed for four grades of upslope walking, from 25 to 100% (45° incline), and compared with similar data for level treadmill walking (0.6 m/s). Kinematic data for the hip, knee, ankle, and metatarsophalangeal joints were obtained from digitizing ciné film that was synchronized with electromyographic (EMG) records from 13 different hindlimb muscles. Cycle periods, the structure of the step cycle, and paw-contact sequences were similar at all grades and typical of lateral-sequence walking. Also, a few half-bound and transverse gallop steps were assessed from trials at the 100% grade; these steps had shorter cycle periods than the walking steps and less of the cycle (68 vs. 56%) was devoted to stance. Each cat assumed a crouched posture at the steeper grades of upslope walking and stride length decreased, whereas the overall position of the stride shifted caudally with respect to the hip joint. At the steeper grades, the range and duration of swing-related flexion increased at all joints, the stance-phase yield was absent at the knee and ankle joints, and the range of stance-phase extension at knee and ankle joints increased. Patterns of muscle activity for upslope and level walking were similar with some notable exceptions. At the steeper grades, the EMG activity of muscles with swing-related activity, such as the digit flexor muscle, the flexor digitorum longus (FDL), and the knee flexor muscle, the semitendinosus (ST), was prolonged and continued well into midswing. The EMG activity of stance-related muscles also increased in amplitude with grade, and three muscles not active during the stance phase of level walking had stance activity that increased in amplitude and duration at the steepest grades; these muscles were the ST, FDL, and extensor digitorum brevis. Overall the changes in posture, hindlimb kinematics, and the activity patterns of hindlimb muscles during upslope walking reflected the need to continually move the body mass forward and upward during stance and to ensure that the paw cleared the inclined slope during swing. The implications of these changes for the neural control of walking and expected changes in hindlimb kinetics for slope walking are discussed.

Forms of Forward Quadrupedal Locomotion. III. A Comparison of Posture, Hindlimb Kinematics, and Motor Patterns for Downslope and Level Walking

1998 ◽  
Vol 79 (4) ◽  
pp. 1702-1716 ◽  
Author(s):  
Judith L. Smith ◽  
Patricia Carlson-Kuhta ◽  
Tamara V. Trank
Keyword(s):  
Ankle Joint ◽  
Neural Control ◽  
Activity Patterns ◽  
Motor Patterns ◽  
Quadrupedal Locomotion ◽  
Hindlimb Muscles ◽  
Level Walking ◽  
Angular Displacements ◽  
External Forces ◽  
Hindlimb Kinematics

Smith, Judith L., Patricia Carlson-Kuhta, and Tamara V. Trank. Forms of forward quadrupedal locomotion. III. A comparison of posture, hindlimb kinematics, and motor patterns for downslope and level walking. J. Neurophysiol. 79: 1702–1716, 1998. To gain further insight into the neural mechanisms for different forms of quadrupedal walking, data on postural orientation, hindlimb kinematics, and motor patterns were assessed for four grades of downslope walking, from 25% (14° slope) to 100% (45°), and compared with data from level and downslope walking at five grades (5–25%) on the treadmill (0.6 m/s). Kinematic data were obtained by digitizing ciné film, and electromyograms (EMGs) synchronized with kinematic records were taken from 13 different hindlimb muscles. At grades from 25 to 75%, cycle periods were similar, but at the steepest grade the cycle was shorter because of a reduced stance phase. Paw-contact sequences at all grades were consistent with lateral-sequence walking, but pace walking often occurred at the steepest grades. The cats crouched at the steeper grades, and crouching was associated with changes in fore- and hindlimb orientation that were consistent with increasing braking forces and decreasing propulsive forces during stance. The average ranges of motion at the hindlimb joints, except at the hip, were often different at the two steepest slopes. During swing, the range of knee- and ankle-joint flexion decreased, and the range and duration of extension increased at the ankle joint to lower the paw downward for contact. During stance the range of flexion during yield increased at the ankle joint, and the range of extension decreased at the knee and metatarsophalangeal joints. Downslope walking was also associated with EMG changes for several muscles. The hip extensors were not active during stance; instead, hip flexors were active, presumably to slow the rate of hip extension. Although ankle extensors were active during stance, their burst durations were truncated and centered around paw contact. Ankle flexors were active after midstance at the steeper slopes before the need to initiate swing, whereas flexor and extensor digit muscles were coactive throughout stance. Overall the changes in posture, hindlimb kinematics, and activity patterns of hindlimb muscles during stance reflected a need to counteract external forces that would accelerate angular displacements at some joints. Implications of these changes are discussed by using current models for the neural control of walking.

COMMON AND DISTINCT MUSCLE SYNERGIES DURING LEVEL AND SLOPE LOCOMOTION IN THE CAT

2021 ◽  
Author(s):  
Alexander N Klishko ◽  
Adil Akyildiz ◽  
Ricky Mehta ◽  
Boris I. Prilutsky
Keyword(s):  
Stance Phase ◽  
Rectus Femoris ◽  
Emg Activity ◽  
Modular Organization ◽  
Muscle Synergies ◽  
Cycle Duration ◽  
Hindlimb Muscles ◽  
Level Walking ◽  
Adult Cats ◽  
Activity Cluster

Although it is well established that the motor control system is modular, the organization of muscle synergies during locomotion and their change with ground slope are not completely understood. For example, typical reciprocal flexor-extensor muscle synergies of level walking in cats break down in downslope: one-joint hip extensors are silent throughout the stride cycle, whereas hindlimb flexors demonstrate an additional stance phase-related EMG burst (Smith et al. 1998a). Here we investigated muscle synergies during Level, Upslope (27o) and Downslope (-27o) walking in adult cats to examine common and distinct features of modular organization of locomotor EMG activity. Cluster analysis of EMG burst onset-offset times of 12 hindlimb muscles revealed 5 flexor and extensor burst groups that were generally shared across slopes. Stance-related bursts of flexor muscles in downslope were placed in a burst group from Level and Upslope walking formed by the rectus femoris. Walking Upslope changed swing/stance phase durations of Level walking but not the cycle duration. Five muscle synergies computed using non-negative matrix factorization accounted for at least 95% of variance in EMG patterns in each slope. Five synergies were shared between Level and Upslope walking, whereas only 3 of those were shared with Downslope synergies; these synergies were active during the swing phase and phase transitions. Two stance-related synergies of downslope walking were distinct; they comprised a mixture of flexors and extensors. We suggest that the modular organization of muscle activity during Level and Slope walking results from interactions between motion-related sensory feedback, CPG, and supraspinal inputs.

Corrective responses to loss of ground support during walking. I. Intact cats

1994 ◽  
Vol 71 (2) ◽  
pp. 603-610 ◽  
Author(s):  
M. A. Gorassini ◽  
A. Prochazka ◽  
G. W. Hiebert ◽  
M. J. Gauthier
Keyword(s):  
Stance Phase ◽  
Afferent Input ◽  
Emg Activity ◽  
Step Cycle ◽  
Trap Door ◽  
Ground Support ◽  
Extensor Muscles ◽  
Activation Profile ◽  
Corrective Response ◽  
Hindlimb Kinematics

1. In the cat step cycle the electromyographic (EMG) activity in ankle extensor muscles commences approximately 70 ms before foot contact. There is a sharp peak between 10 and 25 ms after contact and the EMG then declines for the remainder of the stance phase. It has been posited that the abrupt transition in EMG after contact is the consequence of reflexes elicited by the large barrage of afferent input that signals foot touchdown. However, it is also possible that the basic profile might be generated within the CNS, with little modification by afferent input. 2. These ideas were tested in 11 normal cats. We compared EMG responses and hindlimb kinematics in steps with normal ground support and steps in which an actuator-controlled trap door unexpectedly opened, withdrawing ground support just before foot contact. 3. In the absence of ground support the transition in EMG activity was still present. The averaged EMG pattern was similar for at least 30 ms after the foot passed through the plane of the floor. We conclude that the basic extensor activation profile in this part of the cycle is generated centrally and is not substantially altered by afferent input. 4. Between 35 and 200 ms after contact the stance phase was aborted and the foot was lifted smartly out of the hole. This reaction varied both in latency and kinematic detail, suggesting a fairly complex corrective response.(ABSTRACT TRUNCATED AT 250 WORDS)

Adaptive control for backward quadrupedal walking. I. Posture and hindlimb kinematics

1990 ◽  
Vol 64 (3) ◽  
pp. 745-755 ◽  
Author(s):  
J. A. Buford ◽  
R. F. Zernicke ◽  
J. L. Smith
Keyword(s):  
Stride Length ◽  
Stance Phase ◽  
Horizontal Displacement ◽  
Knee Extension ◽  
Treadmill Walking ◽  
Cycle Period ◽  
Phase Duration ◽  
Ankle Joints ◽  
Hindlimb Kinematics ◽  
Hip Extension

1. To gain new perspectives on the neural control of different forms of quadruped locomotion, we studied adaptations in posture and hindlimb kinematics for backward (BWD) walking in normal cats. Data from four animals were obtained from high-speed (100 fr/s) cine film of BWD treadmill walking over a range of slow walking speeds (0.3-0.6 m/s) and forward (FWD) treadmill walking at 0.6 m/s. 2. Postural adaptations during BWD walking included flexion of the lumbar spine, compared to a relatively straight spine during FWD walking. The usual paw-contact sequence for FWD walking [right hindlimb (RH), right forelimb (RF), left hindlimb (LH), left forelimb (LF)] was typically reversed for BWD walking (RH, LF, LH, RF). The hindlimbs alternated consistently with a phase difference averaging 0.5 for both forms of walking, but the phasing of the forelimbs was variable during BWD walking. 3. As BWD walking speed increased from 0.3 to 0.6 m/s, average hindlimb cycle period decreased 21%, stance-phase duration decreased 29%, and stride length increased 38%. Compared to FWD walking at 0.6 m/s, stride length was 30% shorter, whereas cycle period and stance-phase duration were 17% shorter for BWD walking. For both directions, stance occupied 64 +/- 4% (mean +/- SD) of the step cycle. 4. During swing for both forms of walking, the hip, knee, and ankle joints had flexion (F) and extension (E1) phases; however, the F-E1 reversals occurred earlier at the hip and later at the knee for BWD than for FWD walking. At the ankle joint, the ranges of motion during the F and E1 phases were similar for both directions. During BWD walking, however, the knee flexed more and extended less, whereas the hip flexed less and extended more. Thus horizontal displacement of the limb resulted primarily from hip extension and knee flexion during BWD swing, but hip flexion and knee extension during FWD swing. 5. At the knee and ankle joints, there were yield (E2) and extension (E3) phases during stance for both forms of walking; however, yields at the knee and ankle joints were reduced during BWD walking. At the hip, angular motion was unidirectional, as the hip flexed during BWD stance but extended during FWD stance. Knee extension was the prime contributor to horizontal displacement of the body during BWD stance, but hip extension was the prime contributor to horizontal displacement during FWD stance. 6. Our kinematic data revealed two discriminators between BWD and FWD walking.(ABSTRACT TRUNCATED AT 400 WORDS)

Forms of forward quadrupedal locomotion. I. A comparison of posture, hindlimb kinematics, and motor patterns for normal and crouched walking

1996 ◽  
Vol 76 (4) ◽  
pp. 2316-2326 ◽  
Author(s):  
T. V. Trank ◽  
C. Chen ◽  
J. L. Smith
Keyword(s):  
Activity Patterns ◽  
Swing Phase ◽  
Average Height ◽  
Normal Walking ◽  
Step Cycle ◽  
Hindlimb Muscles ◽  
Ankle Joints ◽  
Hindlimb Kinematics ◽  
Treadmill Belt ◽  
Mtp Joints

1. Posture, hindlimb kinematics, and activity patterns of selected hindlimb muscles were compared for normal and crouched treadmill walking (0.5-0.6 m/s) for eight cats. To elicit crouched walking in which the trunk and head were lowered, cats were encouraged to walk under a light-weight Plexiglas ceiling suspended 17-20 cm above the treadmill belt. Kinematic data were obtained from high-speed cine film, and electromyograms (EMGs)-synchronized with the kinematic records-were taken from 11 hindlimb muscles. 2. The postures for the two forms of walking were distinctly different. During crouched walking, each cat lowered its entire body keeping its trunk horizontal to the treadmill belt. Also the head was lowered, with the top of the head in line with the dorsal surface of the trunk. Hip height, used as a measure for hindlimb crouch, was reduced by 30%, from an average height of 23 cm to an average height of 16 cm above the belt during the entire step cycle. 3. Average cycle periods (766 +/- 30 ms, mean +/- SD) and percentage of time devoted to swing (30%) and stance (70%) were similar for normal and crouched walking. The profiles of the hindlimb kinematics were also similar for the hip, knee, ankle, and metatarsophalangeal (MTP) joints during the step cycle, but the timing of some of the motion reversal, as well as the ranges of motion during various phases, were different at some joints for the two forms of walking. 4. During the swing phase, the transition between the flexion and extension (F-E1 reversal) occurred later in the normalized swing phase at the hip, knee, and ankle joints, and the range of flexion was increased at each joint. With greater flexion at these joints, the anatomic axis of the hindlimb (measured from hip joint to toe) was decreased and the hind paw advanced in the narrow space between the abdomen and treadmill belt. At contact, the position of the paw was less anterior to the perpendicular reference line (hip joint marker to belt) and all joints were more flexed for crouched than normal walking. 5. Throughout the stance phase, the knee and ankle joints remained significantly more flexed by 41-45 deg during crouched than normal walking. Although the hip and MTP joints started in a more flexed position at paw contact, both joints extended more during stance for crouched than normal walking, and at the time of peak extension (just before paw lift-off), the degree of extension at the hip and MTP joints was similar for both forms of walking. 6. Muscle patterns for crouched and normal walking were similar with some exceptions. The burst durations for three primary flexor muscles, the semitendinosus (knee flexor), extensor digitorum longus (EDL, ankle flexor), and flexor digitorum longus (digit flexor) were longer for crouched than normal walking, and this was consistent with the increased range and duration of flexion during the swing phase of crouched walking. Also, two muscles that normally showed mainly swing-related activity during normal walking, the EDL and the extensor digitorum brevis, had distinct stance-related bursts that occurred after midstance during crouched walking. 7. Crouched walking requires a postural change that typically occurs when cats stalk prey and when cats walk up and down sleep slopes. Postural set during walking appears to be determined by brain stem and diencephalic centers, and the postural orientation of the cat may require adjustments in the motor program provided by spinal centers for the cat to walk. The role of posture and locomotion and the adjustments in hindlimb kinematics and EMG activity patterns have been studied for forward and backward walking in the cat and now for crouched walking on the treadmill. These data will assist us in understanding the role of posture, especially crouched posture, during other walking behaviors.

Evaluation of an Experimental Testing System for Non-Powered Hand Tools

2000 ◽  
Vol 44 (22) ◽  
pp. 643-646 ◽  
Author(s):  
Tanja Niemelä ◽  
Markku Leppänen ◽  
Minna Päivinen ◽  
Markku Mattila
Keyword(s):  
Measurement System ◽  
Experimental Testing ◽  
Opening Angle ◽  
Emg Activity ◽  
Testing System ◽  
Strain Gages ◽  
Flexor Muscle ◽  
Hand Tools ◽  
Hand Tool ◽  
Forearm Flexor

During the Eurohandtool Project an experimental testing system for non-powered hand tools was developed for laboratory testing. With this system, it is possible to measure simultaneously (1) the EMG activity of two muscles, (2) the opening angle of hand tool blades by means of a potentiometer and, (3) by means of strain gages, the force transmitted to the handle. The first part of evaluation of the system was to determine its time of warming-up, reliability, linearity and repeatability. This paper concentrates on the second part, during which the aim was to test the measurement system by comparing the forces needed to cut wood of a certain diameter, and the actual force required, as measured by a material-testing system. The correlation between forearm flexor muscle activity and the compression force created by the user was investigated. The evaluation of experimental testing system for non-powered hand tools has shown that there are methods to measure force demand, opening angle and EMG-activity simultaneously. However, it is recommended to make some improvements before this measurement system can be taken into widespread use.

Brain stem integration of vocalization: role of the midbrain periaqueductal gray

1994 ◽  
Vol 72 (3) ◽  
pp. 1337-1356 ◽  
Author(s):  
S. P. Zhang ◽  
P. J. Davis ◽  
R. Bandler ◽  
P. Carrive
Keyword(s):  
Muscle Activity ◽  
Sound Production ◽  
Type A ◽  
Periaqueductal Gray ◽  
Emg Activity ◽  
Digastric Muscle ◽  
Type B ◽  
Motor Patterns ◽  
Homocysteic Acid ◽  
Lower Amplitude

1. The contribution of the midbrain periaqueductal gray (PAG) to the central regulation of vocalization was investigated by analyzing the electromyographic (EMG) changes in respiratory, laryngeal, and oral muscles evoked by microinjection of D,L-homocysteic acid (DLH) in the PAG of unanesthetized, precollicular decerebrate cats. Moderate to large (6-40 nmol) doses of DLH evoked natural-sounding vocalization as well as increases in inspiratory depth and respiratory rate. 2. Two basic types of vocalization were evoked, each associated with a distinct and characteristic pattern of respiratory, laryngeal and oral EMG changes. Type A vocalization (voiced sounds such as howl/mew/growl) was characterized by excitation of the cricothyroid (CT) and thyro-arytenoid (TA) muscles, and inhibition of the posterior crico-arytenoid (PCA) muscle, whereas type B vocalization (unvoiced hiss sounds) was characterized by excitation of the PCA and TA muscles and no significant activation of the CT muscle. In addition, stronger expiratory (external oblique, internal oblique, internal intercostal) EMG increases were associated with type A responses, and larger increases in genioglossus and digastric muscle activity were associated with type B responses. 3. Microinjections of small doses of DLH (300 pmol-3 nmol), also evoked patterned changes in muscle activity (usually without audible vocalization) that, although of lower amplitude, were identical to those evoked by injections of moderate to large DLH doses. In no such experiments (175 sites) were individual muscles activated by small dose injections of DLH into the PAG. Further, type A vocalization/muscle patterns were evoked from PAG sites caudal to those at which type B vocalization/muscle patterns were evoked. 4. Considered together these results indicate: that the PAG contains topographically separable groups of neurons that coordinate laryngeal, respiratory, and oral muscle patterns characteristic of two fundamental types of vocalization and that the underlying PAG organization takes the form of a representation of muscle patterns, rather than individual muscles. 5. The patterns of EMG activity evoked by excitation of PAG neurons were strikingly similar to previously reported patterns of EMG activity characteristic of major phonatory categories in higher species, including humans (e.g., vowel phonation, voiceless consonant phonation). These findings raise the possibility that the sound production circuitry of the PAG could well be utilized by cortical and subcortical "language structures" to coordinate basic respiratory and laryngeal motor patterns that are necessary for speech.

scholarly journals Major role for sensory feedback in soleus EMG activity in the stance phase of walking in man

2000 ◽  
Vol 523 (3) ◽  
pp. 817-827 ◽  
Author(s):  
T. Sinkjær ◽  
J. B. Andersen ◽  
M. Ladouceur ◽  
L. O. D. Christensen ◽  
J. B. Nielsen
Keyword(s):  
Sensory Feedback ◽  
Stance Phase ◽  
Emg Activity

scholarly journals Motor Patterns During Walking on a Slippery Walkway

2010 ◽  
Vol 103 (2) ◽  
pp. 746-760 ◽  
Author(s):  
Germana Cappellini ◽  
Yuri P. Ivanenko ◽  
Nadia Dominici ◽  
Richard E. Poppele ◽  
Francesco Lacquaniti
Keyword(s):  
Motor Output ◽  
Emg Activity ◽  
Head Orientation ◽  
Cycle Duration ◽  
Horizontal Shear ◽  
Motor Patterns ◽  
Spatiotemporal Organization ◽  
Surface Conditions ◽  
Gait Kinematics ◽  
Slippery Surface

Friction and gravity represent two basic physical constraints of terrestrial locomotion that affect both motor patterns and the biomechanics of bipedal gait. To provide insights into the spatiotemporal organization of the motor output in connection with ground contact forces, we studied adaptation of human gait to steady low-friction conditions. Subjects walked along a slippery walkway (7 m long; friction coefficient ≃ 0.06) or a normal, nonslippery floor at a natural speed. We recorded gait kinematics, ground reaction forces, and bilateral electromyographic (EMG) activity of 16 leg and trunk muscles and we mapped the recorded EMG patterns onto the spinal cord in approximate rostrocaudal locations of the motoneuron (MN) pools to characterize the spatiotemporal organization of the motor output. The results revealed several idiosyncratic features of walking on the slippery surface. The step length, cycle duration, and horizontal shear forces were significantly smaller, the head orientation tended to be stabilized in space, whereas arm movements, trunk rotations, and lateral trunk inclinations considerably increased and foot motion and gait kinematics resembled those of a nonplantigrade gait. Furthermore, walking on the slippery surface required stabilization of the hip and of the center-of-body mass in the frontal plane, which significantly improved with practice. Motor patterns were characterized by an enhanced (roughly twofold) level of MN activity, substantial decoupling of anatomical synergists, and the absence of systematic displacements of the center of MN activity in the lumbosacral enlargement. Overall, the results show that when subjects are confronted with unsteady surface conditions, like the slippery floor, they adopt a gait mode that tends to keep the COM centered over the supporting limbs and to increase limb stiffness. We suggest that this behavior may represent a distinct gait mode that is particularly suited to uncertain surface conditions in general.

Scratch responses in normal cats: hindlimb kinematics and muscle synergies

1990 ◽  
Vol 64 (6) ◽  
pp. 1653-1667 ◽  
Author(s):  
P. C. Kuhta ◽  
J. L. Smith
Keyword(s):  
Muscle Activity ◽  
Knee Flexion ◽  
Knee Extensor ◽  
Head Movements ◽  
Medial Gastrocnemius ◽  
Emg Activity ◽  
Circular Path ◽  
Contact Phase ◽  
Ankle Joints ◽  
Flexor Muscles

1. Scratch responses evoked by a tactile stimulus applied to the outer ear canal were characterized in nine adult cats. Chronic electromyographic (EMG) electrodes were surgically implanted in selected flexor and extensor muscles of the hip, knee, and ankle joints to determine patterns of muscle activity during scratching. In some trials EMG records were synchronized with kinematic data obtained by digitizing high-speed cine film, and in one cat, medial gastrocnemius (MG) tendon forces were recorded along with EMG. For analysis the response was divided into three components: the approach, cyclic, and return periods. Usually scratch responses were initiated with the cat in a sitting position, but in some trials the animal initiated the response from a standing or lying posture. 2. During the approach period the hindlimb ipsilateral to the stimulated ear was lifted diagonally toward the head by a combination of hip and ankle flexion with knee extension. Hindlimb motions during the approach period were associated with sustained EMG activity in hip-flexor, knee-extensor (occasionally), and ankle-flexor muscles. Initial hindlimb motions were typically preceded by head movements toward the hindpaw, and at the end of the approach period, the head was tilted downward with the stimulated pinna lower than the contralateral ear. During the return period movements were basically the reverse of the approach period, with the hindpaw returning to the ground and the head moving away from the hindlimb. 3. During the cyclic period the number of cycles per response varied widely from 1 to 60 cycles with an average of 13 cycles, and cycle frequency ranged from 4 to 8 cycles/s, with a mean of 5.6 cycles/s. During each cycle the paw trajectory followed a fairly circular path, and the cycle was defined by three phases: precontact, contact, and postcontact. On average the contact phase occupied approximately 50% of the cycle and was characterized by extensor muscle activity and extension at the hip, knee, and ankle joints. The hindpaw contacted the pinna or neck at the base of the pinna throughout the contact phase, and paw contact typically resulted in a rostral motion of the head as the hindlimb extended. 4. The postcontact phase constituted approximately 24% of scratch cycle and was usually initiated by the onset of knee flexion. Ankle and then hip flexion followed knee flexion, and flexor muscles were active during the postcontact phase as the paw was withdrawn from the head. The precontact phase constituted approximately 26% of scratch cycle and was initiated by knee joint extension and knee-extensor activity.(ABSTRACT TRUNCATED AT 400 WORDS)

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