Stabilization and mobility of the head, neck and trunk in horses during overground locomotion: comparisons with humans and other primates.
Abstract: Segmental kinematics were investigated in horses during overground locomotion and compared with published reports on humans and other primates to determine the impact of a large neck on rotational mobility (> 20 deg.) and stability (< or = 20 deg.) of the head and trunk. Three adult horses (Equus caballus) performing walks, trots and canters were videotaped in lateral view. Data analysis included locomotor velocity, segmental positions, pitch and linear displacements and velocities, and head displacement frequencies. Equine, human and monkey skulls and cervical spines were measured to estimate eye and vestibular arc length during head pitch displacements. Horses stabilized all three segments in all planes during all three gaits, unlike monkeys and humans who make large head pitch and yaw rotations during walks, and monkeys that make large trunk pitch rotations during gallops. Equine head angular displacements and velocities, with some exceptions during walks, were smaller than in humans and other primates. Nevertheless, owing to greater off-axis distances, orbital and vestibular arc lengths remained larger in horses, with the exception of head-neck axial pitch during trots, in which equine arc lengths were smaller than in running humans. Unlike monkeys and humans, equine head peak-frequency ranges fell within the estimated range in which inertia has a compensatory stabilizing effect. This inertial effect was typically over-ridden, however, by muscular or ligamentous intervention. Thus, equine head pitch was not consistently compensatory, as reported in humans. The equine neck isolated the head from the trunk enabling both segments to provide a spatial reference frame.
Publication Date: 2008-12-02 PubMed ID: 19043061PubMed Central: PMC2768006DOI: 10.1242/jeb.020578Google Scholar: Lookup
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- Comparative Study
- Journal Article
- Research Support
- N.I.H.
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- Research Support
- Non-U.S. Gov't
Summary
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This research investigates the motion and stability of the head, neck, and trunk of horses during different gaits and compares this to similar data from humans and other primates. The study offers meaningful insights into how the sizable neck of a horse affects its rotational mobility and stability.
Methodology
- The experiment involved three adult horses recorded in lateral view while walking, trotting, and cantering.
- Data collected include locomotor speed, positions in different segments, changes in angle (pitch) and linear motions and speeds, and the frequency of head displacements.
- Physical measurements were taken from horse, human, and monkey skulls and neck bones to predict the change in distance between the eyes and balance organ (vestibular arc length) during movements of the head.
Key Results
- Unlike humans and monkeys who have large head rotations when walking, and monkeys have large trunk rotations during gallops, the horses were able to maintain the stability of their head, neck, and trunk during all gaits.
- Though horses generally had smaller angular movements and speeds than humans and primates, the longer distances from the rotation axis resulted in more significant orbital and vestibular arc lengths. However, running humans showed greater arc lengths when compared with trotting horses.
- Unlike in monkeys and humans, the frequency range of head movement in horses was within the estimated range in which inertia helps stabilize the head, suggesting a different balance strategy.
Significance
- This research uncovers crucial knowledge about locomotion in horses and other primates. It shows that horses, thanks to their muscular or ligamentous intervention, manage to keep a stabilizing effect despite their large necks. Their large neck also allows their heads to work independently from their trunks, providing two separate reference frames for balance and spatial awareness.
Cite This Article
APA
Dunbar DC, Macpherson JM, Simmons RW, Zarcades A.
(2008).
Stabilization and mobility of the head, neck and trunk in horses during overground locomotion: comparisons with humans and other primates.
J Exp Biol, 211(Pt 24), 3889-3907.
https://doi.org/10.1242/jeb.020578 Publication
Researcher Affiliations
- Department of Anatomy and Neurobiology, and Caribbean Primate Research Center, University of Puerto Rico School of Medicine, PO Box 365067, San Juan, PR. ddunbar@rcm.upr.edu
MeSH Terms
- Animals
- Biomechanical Phenomena
- Body Weight
- Gait
- Head / physiology
- Horses / anatomy & histology
- Horses / physiology
- Humans
- Locomotion / physiology
- Male
- Neck / physiology
- Primates / physiology
- Videotape Recording
Grant Funding
- G12 MD007600 / NIMHD NIH HHS
- G12 RR003051 / NCRR NIH HHS
- P40 RR003640 / NCRR NIH HHS
- RR-03051 / NCRR NIH HHS
References
This article includes 102 references
- Alexander RM. Locomotion in Animals. Glasgow: Blackle; 1982.
- Angelakl DE, McHenry MQ, Dickman JD, Newlands SD, Hess BJM. Computation of inertial motion: neural strategies to resolve ambiguous ololith information. J Neurosci 1999;19:316–327.
- Angelakl DE, Shaikh AG, Green AM, Dickman JD. Neurons compute internal models of the physical laws of motion. Nature 2004;430:560–564.
- Ankel-Simons F. Primate Anatomy: An Introduction. 2. San Diego: Academic Press; 2000.
- Attenburrow DP. Time relationships between the respiratory cycle and the limb cycle in the horse. Equine Vet J 1982;14:69–72.
- Attenburrow DP, Goss VA. The mechanical coupling of lung ventilation to locomotion in the horse. Med Eng Phys 1994;16:188–192.
- Bramble DM. Axial-appendicular dynamics and the integration of breathing and gait in mammals. Am Zool 1989;29:171–186.
- Bramble DM, Carrier DR. Running and breathing in mammals. Science 1983;219:251–256.
- Bramble DM, Jenkins FA. Jr. Mammalian locomotor-respiratoty Integration: implications for diaphragmatic and pulmonary design. Science 1993;262:235–240.
- Cappozzo A. Low frequency self-generated vibration during ambulation in normal men. J Biomech 1982;8:599–609.
- Clayton HM. The Dynamic Horse. Mason, Ml: Sport Horse; 2004.
- Clayton HM, Townsend HGG. Kinematics of the cervical spine of the adult horse. Equine Vet J 1989;21:189–192.
- Crompton RH, Li Y, Alexander RM, Wang W, Gunther MM. Segmental inertial properties of primates: new techniques for laboratory and field studies of locomotion. Am J Phys Anthropol 1996;99:547–570.
- Cromwell R, Wellmon R. Sagittal plane head stabilization during level walking and ambulation on stairs. Physiother Res Int 2001;6:179–192.
- Cromwell RL, Aadland-Monahan TK, Nelson AT, Stem-Sylvestre SM, Seder B. Sagittal plane analysis of head, neck, and trunk kinematics and electromyographic activity during locomotion. J Orthop Sports Phys Ther 2001;31:255–262.
- Cromwell RL, Pldcoe PE, Griffin LA, Sotillo T, Ganninger D, Feagin M. Adaptations in horizontal head stabilization in response to altered vision and gaze during natural walking. J Vestib Res 2004;14:367–373.
- Das P, McCollum G. Invariant structure in locomotion. Neuroscience 1988;25:1023–1034.
- Day BL, Fltzpatrick RC. Virtual head rotation reveals a process of route reconstruction from human vestibular signals. J Physiol (Lond) 2005;567:591–597.
- DeBeer GR. How animals hold their heads. Proc Linn Soc Lond 1947;159:125–139.
- Dempster WT. Space requirements of the seated operator. Wright Air Development Center (WADC) Technical Report 55–159. Ohio: Wright-Patterson Air Force Base; 1955.
- Duke-Elder SS. System of Ophthalmology. Vol. 1, The Eye in Evolution. London: Henry Kimpton; 1958.
- Dunbar DC. Aerial maneuvers of leaping lemurs: the physics of whole-body rotations while airborne. Am J Primatol 1988;16:291–303.
- Dunbar DC. The influence of segmental movements and design on whole-body rotations during the airborne phase of primate leaps. Z Morphol Anthropol 1994;80:109–124.
- Dunbar DC. Stabilization and mobility of the head and trunk in vervel monkeys (Cereopithecus aethiops) during treadmill walks and gallops. J Exp Biol 2004;207:4427–4438.
- Dunbar DC, Macpherson JM. Activity of neuromuscular compartments in lateral gastrocnemius evoked by postural corrections during stance. J Neurophysiol 1993;70:2337–2349.
- Dunbar DC, Badam GL, Hallgrimsson B, Vlellledent S. Stabilization and mobility of the head and trunk in wild monkeys during terrestrial and flat-surface walks and gallops. J Exp Biol 2004;207:1027–1042.
- Dvorak J. Funktionelle Anatomie der oberen Halswirbelsäule unter besonderer Berückersichtigung des Bandapparates. In: Wolff HD, editor. Die Sonderstellung des Koplgelenkbereichs. Berlin: Springer; 1988. pp. 19–46.
- Edwards EH. The New Encyclopedia of the Horse. New York: Dorling Kindersley; 2001.
- Einstein A. Über das Relativitätsprinzip und die aus demselben gezogenen Folgerungen. Jahrb Radioakt 1908;4:411–462.
- Frohlich C. Do springboard divers violate angular momentum conservation?. Am J Phys 1979;47:583–592.
- Frohlich C. The physics of somersaulting and twisting. Sci Am 1980;242:154–164.
- Fuchs AF. Eye-head coordination. In: Towe AL, Luschei ES, editors. Handbook of Behavioral Neurobiology, Vol. 5: Motor Coordination. New York: Plenum Press; 1981. pp. 303–366.
- Gans C, Gaunt AS. Muscle architecture and control demands. Brain Behav Evol 1992;40:70–81.
- Gellman KS, Bertram JEA. The equine nuchal ligament 1, structural and material properties. Vet Comp Orthop Traumatol 2002a;15:1–6.
- Gellman KS, Bertram JEA. The equine nuchal ligament 2, passive dynamic energy exchange in locomotion. Vet Comp Orthop Traumatol 2002b;15:7–14.
- Gellman KS, Bertram JEA, Hermanson JW. Morphology, histoehemistry, and function of epaxial cervical musculature in the horse (Equus caballus). J Morphol 2002;251:182–194.
- Goody PC. Horse Anatomy: A Pictorial Approach to Equine Structure. London: J.A. Allen; 1983.
- Graf W, de Waele C, Vidal PP. Functional anatomy of the head-neck system of quadrupedal and bipedal mammals. J Anat 1995a;186:55–74.
- Graf W, de Waele C, Vidal PP, Wang DH, Evinger C. The orientation of the cervical vertebral column in unrestrained awake animals. II: movement strategies. Brain Behav Evol 1995b;45:209–231.
- Grossman GE, Leigh RJ, Abel LA, Lanska DJ, Thurston SE. Frequency and velocity of rotational head perturbation during locomolion. Exp Brain Res 1988;70:470–476.
- Harmon AM, Moore S, Hoskins R, Keller P. Horse vision and an explanation for the visual behaviour originally explained by the “ramp retina”. Equine Vet J 1999;31:384–390.
- Hess BJM. Sensorimotor transformations in spatial orientation relative to gravity. In: Mast FW, Jäncke L, editors. Spatial Processing in Navigation, Imagery and Perception. New York: Springer; 2007. pp. 281–300.
- Hildebrand M. Motions of the running cheetah and horse. J Mammal 1959;40:481–495.
- Hildebrand M. Analysis of symmetrical gaits of tetrapods. Folia Biotheoret 1966;13:9–22.
- Hildebrand M. Analysis of Vertebrate Structure. New York: John Wiley; 1974.
- Hildebrand M. Analysis of symmetrical gaits. J Mammal 1977;58:131–156.
- Hill AV. The dimensions of animals and their muscular dynamics. Sci Prog 1950;38:209–230.
- Hirasaki E, Kumakura H. Head movements during locomotion in a gibbon and Japanese macaques. NeuroReport 2004;15:643–647.
- Howell AB. Speed in Animals. Chicago: University of Chicago Press; 1944.
- Hughes A. The topography of vision in mammals of contrasting lifestyle: comparative optics and retinal organization. In: Crescitelli F, editor. Handbook of Sensory Physiology: the Visual System in Vertebrates. VII/5. Berlin: Springer Verlag; 1977. pp. 615–756.
- Imai T, Moore ST, Raphan T, Cohen B. Interaction of the body, head, and eyes during walking and turning. Exp Brain Res 2001;138:1–18.
- Jakobs T, Miller JAA, Schultz AB. Trunk position sense in the frontal plane. Exp Neurol 1985;90:129–138.
- Johnson RA, Wehrly T. Measures and models for angular correlation and angular-linear correlation. J R Stat Soc Ser B Methodol 1977;39:222–229.
- Jones GM, Spells KE. A theoretical and comparative study of functional dependence of the semicircular canal upon physical dimensions. Proc R Soc Lond, B, Biol Sci 1963;157:403–419.
- Kimura T, Okada M, Ishida H. Kinesiological characteristics of primate walking: its significance in human walking. In: Morbeck ME, Preuschoft H, Gomberg N, editors. Environment, Behavior, and Morphology: Dynamic Interactions in Primates. New York: Gustav Fischer; 1979. pp. 297–311.
- Kleine JF, Guan Y, Kiplani EL, Gionti L, Hoshi M, Büttner U. Trunk position influences vestibular responses of fastigial nucleus neurons in the alert monkey. J Neurophysiol 2004;91:2090–2100.
- Lanovaz JL, Clayton HM, Watson LG. In vitro attenuation of impact shock in equine digits. Equine Vet J Suppl 1998;26:96–102.
- Liao K, Walker MF, Joshi A, Reschke M, Leigh RJ. Vestibuloocular responses to vertical translation in normal human subjects. Exp Brain Res 2008;185:553–582.
- Lieberman DE, Raichien DA, Pontzer H, Bramble DM, Cutright-Smith E. The human gluteus maximus and its role in running. J Exp Biol 2006;209:2143–2155.
- Lindsay KW, Roberts TDM, Rosenberg JR. Asymmetrical tonic labyrinth reflexes and their interaction with neck reflexes in the decerebrate cat. J Physiol (Lond) 1976;261:583–601.
- Macpherson JM, Everaert DG, Stapley PJ, Ting LH. Bilateral vestibular loss in cats leads to active destabilization of balance during pitch and roll rotations of the support surface. J Neurophysiol 2007;97:4357–4367.
- Mardia KV. Linear-circular correlation coefficients and rhythmometry. Biometrika 1976;63:403–405.
- Merfeld DM, Zupan L, Peterka RJ. Humans use internal models to estimate gravity and linear acceleration. Nature 1999;398:615–618.
- Mergner T, Nardl GL, Becker W, Keeke L. The role of canal-neck interaction for the perception of horizontal trunk and head rotation. Exp Brain Res 1983;49:198–208.
- Mergner T, Siebold C, Schwelgart G, Becker W. Human perception of horizontal trunk and head rotation in space during vestibular and neck stimulation. Exp Brain Res 1991;85:389–404.
- Mergner T, Rottler G, Kimmig H, Becker W. Role of vestibular and neck inputs for the perception of object motion in space. Exp Brain Res 1992;89:655–668.
- Mittelstaedt H. The information processing structure of the subjective vertical: a cybernetic bridge between its psychophysics and its neurobiology. In: Marko H, Hauske G, Struppler A, editors. Processing Structures For Perception and Action. Weinheim: V. C. H. Verlagsgesellschaft; 1988. pp. 217–263.
- Mittelstaedt H. Evidence of somatic graviception from new and classical investigations. Acta Otolaryngol Suppl 1995;520:186–187.
- Mittelstaedt H. Somatic graviception. Biol Psychol 1996;42:53–74.
- Mittelstaedt H. Interaction of eye-, head-, and trunk-bound information in spatial perception and control. J Vestib Res 1997;7:283–302.
- Mittelstaedt H. Origin and processing of postural information. Neurosci Biobehav Rev 1998;22:473–478.
- Moore ST, Hirasaki E, Cohen B, Raphan T. Effect of viewing distance on the generation of vertical eye movements during locomotion. Exp Brain Res 1999;129:374–361.
- Paige GD. The influence of larget distance on eye movement responses during vertical linear motion. Exp Brain Res 1989;77:585–593.
- Patla AE. Understanding the control of human locomotion: a prologue. In: Patla AE, editor. Adaptability of Human Gait. Amsterdam: North-Holland; 1991. pp. 3–17.
- Patla AE. Understanding the roles of vision in the control of human locomotion. Gait Posture 1997;5:54–69.
- Peterka RJ. Sensorimotor integration in human posture control. J Neurophysiol 2002;88:1097–1118.
- Peterson BW, Boyle RD. Vestibulocollic reflexes. In: Highstein SM, Fay RR, Popper AN, editors. The Vestibular System. New York: Springer; 2004. pp. 343–374.
- Peterson BW, Goldberg J. Role of vestibular and neck reflexes in controlling eye and head position. In: Roucoux A, Crommelinck M, editors. Physiological and Pathological Aspects of Eye Movements. The Hague: Dr W. Junk; 1931. pp. 352–364.
- Pozzo T, Berthoz A, Lefort L. Head stabilization during various locomotor tasks in humans. I: Normal subjects. Exp Brain Res 1990;82:97–106.
- Roberts SM. Equine vision and optics. Vet Clin North Am Equine Pract 1992;8:451–457.
- Roberts TDM. Neurophysiology of Postural Mechanisms. 2. London: Butterworths; 1978.
- Rochon-Duvigneaud A. Les Yeaux et la Vision des Vértebrés. Paris: Mason; 1943.
- Russell NA, Horil A, Smith PF, Darlington CL, Bilkey DK. Long-term effects of permanent vestibular lesions on hippocampal spatial firing. J Neurosci 2003;23:6490–6498.
- Schmitt D. Insights into the evolution of human bipedalism from experimental studies of humans and other primates. J Exp Biol 2003;206:1437–1448.
- Schubert M, Bohner C, Berger WV, Sprundel M, Duysens JEJ. The role of vision in maintaining heading direction: effects of changing gaze and optic flow on human gait. Exp Brain Res 2003;150:163–173.
- Schwarz U, Miles FA. Ocular responses to translation and their dependence on viewing distance. I: Motion of the observer. J Neurophysiol 1991;66:851–864.
- Shumway-Cook A, Woollacott MH. Motor Control. 2. Philadelphia: Lipplncott Williams and Wilkins; 2001.
- Simpson GG. Horses. New York: Doubleday Anchor; 1961.
- Slijper EJ. Comparative biologic-anatomical investigations on the vertebral column and spinal musculature of mammals. Verh Kon Nederl Akad Wetensch Amsterdam II 1946;42:1–128.
- Stackman RW, Taube JS. Firing properties of head direction cells in the rat anterior thalamic nucleus: dependence on vestibular input. J Neurosci 1997;17:4349–4358.
- Stackman RW, Clark AS, Taube JS. Hippocampal spatial representations require vestibular input. Hippocampus 2002;12:291–303.
- Taylor JL, McCloskey DI. Proprioceptive sensation in rotation of the trunk. Exp Brain Res 1990;81:413–416.
- Timney B, Macuda T. Vision and hearing in horses. J Am Vet Med Assoc 2001;218:1567–1574.
- Townsend HG, Leach DH, Fretz PB. Kinematics of the equine thoracolumbar spine. Equine Vet J 1933;15:117–122.
- Ueda Y, Niki Y, Yoshida K, Masumitsu H. Force plate study of equine biomechanics. Floor reaction force of normal walking and trotting horses. Bull Equine Res Inst 1981;18:28–41.
- Vaitl D, Mittelstaedt H, Balsch F. Shifts in blood volume alter the perception of posture. Int J Psychophysiol 1997;27:99–105.
- Vaitl D, Mittelstaedt H, Saborowski R, Stark R, Baisch F. Shifts in blood volume alter the perception of posture: further evidence for somatic graviception. Int J Psychophysiol 2002;44:1–11.
- Vorstenbosch MATM, Buchner HHF, Savelberg HHCM, Schamhardt HC, Barneveld A. Modeling study of compensatory head movements in lame horses. Am J Vet Res 1997;58:713–718.
- Wake DB. The endoskeleton: the comparative anatomy of the vertebral column and ribs. In: Wake MH, editor. Hyman’s Comparative Vertebrate Anatomy. 3. Chicago: Univerisity of Chicago Press; 1979. pp. 192–237.
- White AA, Panjabi MM. Clinical Biomechanics of the Spine. 2. Philadelphia: Lippincolt; 1990.
- Wilson VJ, Peterson BW. Vestibulospinal and reticulospinal systems. In: Brooks VB, editor. Handbook of Physiology. Sec. 1, Vol. II: Motor Control, Part 1. Bethesda, MD: American Physiological Society; 1981. pp. 667–702.
- Xiang Y, Yakushin SB, Kunin M, Raphan T, Cohen B. Head stabilization by Vestibulocollic reflexes during quadrupedal locomotion in monkey. J Neurophysiol 2008;100:763–780.
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