Motions of the running horse and cheetah revisited: fundamental mechanics of the transverse and rotary gallop

Mammals use two distinct gallops referred to as the transverse (where landing and take-off are contralateral) and rotary (where landing and take-off are ipsilateral). These two gallops are used by a variety of mammals, but the transverse gallop is epitomized by the horse and the rotary gallop by the cheetah. In this paper, we argue that the fundamental difference between these gaits is determined by which set of limbs, fore or hind, initiates the transition of the centre of mass from a downward–forward to upward–forward trajectory that occurs between the main ballistic (non-contact) portions of the stride when the animal makes contact with the ground. The impulse-mediated directional transition is a key feature of locomotion on limbs and is one of the major sources of momentum and kinetic energy loss, and a main reason why active work must be added to maintain speed in locomotion. Our analysis shows that the equine gallop transition is initiated by a hindlimb contact and occurs in a manner in some ways analogous to the skipping of a stone on a water surface. By contrast, the cheetah gallop transition is initiated by a forelimb contact, and the mechanics appear to have much in common with the human bipedal run. Many mammals use both types of gallop, and the transition strategies that we describe form points on a continuum linked even to functionally symmetrical running gaits such as the tölt and amble.

[1]  É. Marey,et al.  Animal mechanism : a treatise on terrestrial and aerial locomotion , 2022 .

[2]  The Evolution of Locomotion in Mammals , 1936 .

[3]  K. Schmidt,et al.  Speed in animals : their specialization for running and leaping , 1945 .

[4]  M. Hildebrand Motions of the Running Cheetah and Horse , 1959 .

[5]  M. Hildebrand Analysis of Asymmetrical Gaits , 1977 .

[6]  G. Cavagna,et al.  Mechanical work in terrestrial locomotion: two basic mechanisms for minimizing energy expenditure. , 1977, The American journal of physiology.

[7]  R. Alexander,et al.  Mechanics of locomotion of dogs (Canis familiaris) and sheep (Ovis aries). , 2009, Journal of zoology.

[8]  A. J. Reynolds,et al.  Velocity distributions in plane turbulent channel flows , 1980, Journal of Fluid Mechanics.

[9]  D. F. Hoyt,et al.  Gait and the energetics of locomotion in horses , 1981, Nature.

[10]  R. Alexander,et al.  Forces exerted on the ground by galloping dogs (Canis familiaris) , 1987 .

[11]  M. Pandy,et al.  The dynamics of quadrupedal locomotion. , 1988, Journal of biomechanical engineering.

[12]  R. M. Alexander Why Mammals Gallop , 1988 .

[13]  N. Heglund,et al.  Speed, stride frequency and energy cost per stride: how do they change with body size and gait? , 1988, The Journal of experimental biology.

[14]  R. Blickhan The spring-mass model for running and hopping. , 1989, Journal of biomechanics.

[15]  M. Hildebrand The quadrupedal gaits of vertebrates , 1989 .

[16]  Tad McGeer,et al.  Passive Dynamic Walking , 1990, Int. J. Robotics Res..

[17]  H C Schamhardt,et al.  Ground reaction force patterns of Dutch Warmbloods at the canter. , 1993, American journal of veterinary research.

[18]  S. Knapp How Animals Move , 1995 .

[19]  M. Coleman,et al.  The simplest walking model: stability, complexity, and scaling. , 1998, Journal of biomechanical engineering.

[20]  G. Dalleau,et al.  The spring-mass model and the energy cost of treadmill running , 1998, European Journal of Applied Physiology and Occupational Physiology.

[21]  A. Minetti,et al.  The relationship between mechanical work and energy expenditure of locomotion in horses. , 1999, The Journal of experimental biology.

[22]  R. S. Simons Running, breathing and visceral motion in the domestic rabbit (Oryctolagus cuniculus): testing visceral displacement hypotheses. , 1999, The Journal of experimental biology.

[23]  M. Coleman,et al.  A point-mass model of gibbon locomotion. , 1999, The Journal of experimental biology.

[24]  J E Bertram,et al.  Segmental in vivo vertebral kinematics at the walk, trot and canter: a preliminary study. , 2001, Equine veterinary journal. Supplement.

[25]  Arthur D Kuo,et al.  Energetics of actively powered locomotion using the simplest walking model. , 2002, Journal of biomechanical engineering.

[26]  P. Aerts,et al.  Two distinct gait types in swimming frogs , 2002 .

[27]  A. Biewener,et al.  Estimates of circulation and gait change based on a three-dimensional kinematic analysis of flight in cockatiels (Nymphicus hollandicus) and ringed turtle-doves (Streptopelia risoria). , 2002, The Journal of experimental biology.

[28]  Rodger Kram,et al.  Simultaneous positive and negative external mechanical work in human walking. , 2002, Journal of biomechanics.

[29]  John E A Bertram,et al.  Understanding brachiation: insight from a collisional perspective , 2003, Journal of Experimental Biology.

[30]  Ronald F. Zernicke,et al.  Gait-related motor patterns and hindlimb kinetics for the cat trot and gallop , 1993, Experimental Brain Research.

[31]  J. Cech,et al.  Photophase and Illumination Effects on the Swimming Performance and Behavior of Five California Estuarine Fishes , 2004, Copeia.

[32]  C. Clanet,et al.  Skipping stones , 2005, Journal of Fluid Mechanics.

[33]  Andy Ruina,et al.  Energetic Consequences of Walking Like an Inverted Pendulum: Step-to-Step Transitions , 2005, Exercise and sport sciences reviews.

[34]  A. Ruina,et al.  A collisional model of the energetic cost of support work qualitatively explains leg sequencing in walking and galloping, pseudo-elastic leg behavior in running and the walk-to-run transition. , 2005, Journal of theoretical biology.

[35]  A. R. Biknevicius,et al.  Locomotor mechanics of the tölt in Icelandic horses. , 2006, American journal of veterinary research.

[36]  Alan M. Wilson,et al.  Centre of mass movement and mechanical energy fluctuation during gallop locomotion in the Thoroughbred racehorse , 2006, Journal of Experimental Biology.

[37]  Manoj Srinivasan,et al.  Computer optimization of a minimal biped model discovers walking and running , 2006, Nature.

[38]  Arthur D Kuo,et al.  The six determinants of gait and the inverted pendulum analogy: A dynamic walking perspective. , 2007, Human movement science.

[39]  Alan M. Wilson,et al.  Gait characterisation and classification in horses , 2007, Journal of Experimental Biology.

[40]  R. M. Walter,et al.  Ground forces applied by galloping dogs , 2007, Journal of Experimental Biology.

[41]  J. Hutchinson,et al.  The three-dimensional locomotor dynamics of African (Loxodonta africana) and Asian (Elephas maximus) elephants reveal a smooth gait transition at moderate speed , 2008, Journal of The Royal Society Interface.

[42]  R. Hackert,et al.  Steady locomotion in dogs: temporal and associated spatial coordination patterns and the effect of speed , 2008, Journal of Experimental Biology.

[43]  R. McN. Alexander,et al.  Fast locomotion of some African ungulates , 2009 .