The Journal of Neurophysiology Vol. 79 No. 4 April 1998, pp. 1687-1701
Copyright ©1998 The American Physiological Society

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

Patricia Carlson-Kuhta, Tamara V. Trank, and Judith L. Smith

Department of Physiological Science, Laboratory of Neuromotor Control, University of California, Los Angeles, California 90095-1568

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.

This article has been cited by other articles:

				T. E. Higham and A. A. Biewener Integration within and between muscles during terrestrial locomotion: effects of incline and speed J. Exp. Biol., July 15, 2008; 211(14): 2303 - 2316. [Abstract] [Full Text] [PDF]

				D.A. McVea and K.G. Pearson Contextual learning and obstacle memory in the walking cat Integr. Comp. Biol., October 1, 2007; 47(4): 457 - 464. [Abstract] [Full Text] [PDF]

				T. J. Roberts, B. K. Higginson, F. E. Nelson, and A. M. Gabaldon Muscle strain is modulated more with running slope than speed in wild turkey knee and hip extensors J. Exp. Biol., July 15, 2007; 210(14): 2510 - 2517. [Abstract] [Full Text] [PDF]

				L. M. Day and B. C. Jayne Interspecific scaling of the morphology and posture of the limbs during the locomotion of cats (Felidae) J. Exp. Biol., February 15, 2007; 210(4): 642 - 654. [Abstract] [Full Text] [PDF]

				D. A. McVea and K. G. Pearson Long-Lasting, Context-Dependent Modification of Stepping in the Cat After Repeated Stumbling-Corrective Responses J Neurophysiol, January 1, 2007; 97(1): 659 - 669. [Abstract] [Full Text] [PDF]

				A. R. Lammers, K. D. Earls, and A. R. Biknevicius Locomotor kinetics and kinematics on inclines and declines in the gray short-tailed opossum Monodelphis domestica J. Exp. Biol., October 15, 2006; 209(20): 4154 - 4166. [Abstract] [Full Text] [PDF]

				R. J. Gregor, D. W. Smith, and B. I. Prilutsky Mechanics of Slope Walking in the Cat: Quantification of Muscle Load, Length Change, and Ankle Extensor EMG Patterns J Neurophysiol, March 1, 2006; 95(3): 1397 - 1409. [Abstract] [Full Text] [PDF]

				O. Ekeberg and K. Pearson Computer Simulation of Stepping in the Hind Legs of the Cat: An Examination of Mechanisms Regulating the Stance-to-Swing Transition J Neurophysiol, December 1, 2005; 94(6): 4256 - 4268. [Abstract] [Full Text] [PDF]

				J. M. Donelan and K. G. Pearson Contribution of Force Feedback to Ankle Extensor Activity in Decerebrate Walking Cats J Neurophysiol, October 1, 2004; 92(4): 2093 - 2104. [Abstract] [Full Text] [PDF]

				A. M. Gabaldon, F. E. Nelson, and T. J. Roberts Mechanical function of two ankle extensors in wild turkeys: shifts from energy production to energy absorption during incline versus decline running J. Exp. Biol., June 1, 2004; 207(13): 2277 - 2288. [Abstract] [Full Text] [PDF]

				L. C. Spezzano Jr and B. C. Jayne The effects of surface diameter and incline on the hindlimb kinematics of an arboreal lizard (Anolis sagrei) J. Exp. Biol., May 15, 2004; 207(12): 2115 - 2131. [Abstract] [Full Text] [PDF]

				M. A. Ashley-Ross and B. F. Bechtel Kinematics of the transition between aquatic and terrestrial locomotion in the newt Taricha torosa J. Exp. Biol., February 1, 2004; 207(3): 461 - 474. [Abstract] [Full Text] [PDF]

				T. E. Higham and B. C. Jayne Locomotion of lizards on inclines and perches: hindlimb kinematics of an arboreal specialist and a terrestrial generalist J. Exp. Biol., January 15, 2004; 207(2): 233 - 248. [Abstract] [Full Text] [PDF]

				T. E. Higham and B. C. Jayne In vivo muscle activity in the hindlimb of the arboreal lizard, Chamaeleo calyptratus: general patterns and the effects of incline J. Exp. Biol., January 15, 2004; 207(2): 249 - 261. [Abstract] [Full Text] [PDF]

				A. L. Ridgel, R. E. Ritzmann, and P. L. Schaefer Effects of aging on behavior and leg kinematics during locomotion in two species of cockroach J. Exp. Biol., December 15, 2003; 206(24): 4453 - 4465. [Abstract] [Full Text] [PDF]

				M. A. Daley and A. A. Biewener Muscle force-length dynamics during level versus incline locomotion: a comparison of in vivo performance of two guinea fowl ankle extensors J. Exp. Biol., September 1, 2003; 206(17): 2941 - 2958. [Abstract] [Full Text] [PDF]

				D. D. Sheriff Muscle pump function during locomotion: mechanical coupling of stride frequency and muscle blood flow Am J Physiol Heart Circ Physiol, June 1, 2003; 284(6): H2185 - H2191. [Abstract] [Full Text] [PDF]

				G. B. Gillis and A. A. Biewener Effects of surface grade on proximal hindlimb muscle strain and activation during rat locomotion J Appl Physiol, November 1, 2002; 93(5): 1731 - 1743. [Abstract] [Full Text] [PDF]

				J. Jing and K. R. Weiss Interneuronal Basis of the Generation of Related but Distinct Motor Programs in Aplysia: Implications for Current Neuronal Models of Vertebrate Intralimb Coordination J. Neurosci., July 15, 2002; 22(14): 6228 - 6238. [Abstract] [Full Text] [PDF]

				M. Y. C. Pang and J. F. Yang Sensory Gating for the Initiation of the Swing Phase in Different Directions of Human Infant Stepping J. Neurosci., July 1, 2002; 22(13): 5734 - 5740. [Abstract] [Full Text] [PDF]

				P. R. Murphy Tonic and Phasic Discharge Patterns in Toe Flexor gamma -Motoneurons During Locomotion in the Decerebrate Cat J Neurophysiol, January 1, 2002; 87(1): 286 - 294. [Abstract] [Full Text] [PDF]

				T. Lam and K. G. Pearson Proprioceptive Modulation of Hip Flexor Activity During the Swing Phase of Locomotion in Decerebrate Cats J Neurophysiol, September 1, 2001; 86(3): 1321 - 1332. [Abstract] [Full Text] [PDF]

				R. E. Burke, A. M. Degtyarenko, and E. S. Simon Patterns of Locomotor Drive to Motoneurons and Last-Order Interneurons: Clues to the Structure of the CPG J Neurophysiol, July 1, 2001; 86(1): 447 - 462. [Abstract] [Full Text] [PDF]

				K. Matsuyama and T. Drew Vestibulospinal and Reticulospinal Neuronal Activity During Locomotion in the Intact Cat. II. Walking on an Inclined Plane J Neurophysiol, November 1, 2000; 84(5): 2257 - 2276. [Abstract] [Full Text] [PDF]

				T. A. Abelew, M. D. Miller, T. C. Cope, and T. R. Nichols Local Loss of Proprioception Results in Disruption of Interjoint Coordination During Locomotion in the Cat J Neurophysiol, November 1, 2000; 84(5): 2709 - 2714. [Abstract] [Full Text] [PDF]

				L. H. Ting, S. A. Kautz, D. A. Brown, and F. E. Zajac Contralateral Movement and Extensor Force Generation Alter Flexion Phase Muscle Coordination in Pedaling J Neurophysiol, June 1, 2000; 83(6): 3351 - 3365. [Abstract] [Full Text] [PDF]

				J Quevedo, B Fedirchuk, S Gosgnach, and D A McCrea Group I disynaptic excitation of cat hindlimb flexor and bifunctional motoneurones during fictive locomotion J. Physiol., June 1, 2000; 525(2): 549 - 564. [Abstract] [Full Text] [PDF]

				T. Lamb and J. F. Yang Could Different Directions of Infant Stepping Be Controlled by the Same Locomotor Central Pattern Generator? J Neurophysiol, May 1, 2000; 83(5): 2814 - 2824. [Abstract] [Full Text] [PDF]

				R. Grasso, M. Zago, and F. Lacquaniti Interactions Between Posture and Locomotion: Motor Patterns in Humans Walking With Bent Posture Versus Erect Posture J Neurophysiol, January 1, 2000; 83(1): 288 - 300. [Abstract] [Full Text] [PDF]

				J. L. Smith, P. Carlson-Kuhta, and T. V. Trank Forms of Forward Quadrupedal Locomotion. III. A Comparison of Posture, Hindlimb Kinematics, and Motor Patterns for Downslope and Level Walking J Neurophysiol, April 1, 1998; 79(4): 1702 - 1716. [Abstract] [Full Text] [PDF]