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The role of plantigrady and heel-strike in the mechanics and energetics of human walking with implications for the evolution of the human foot

Webber, James T., Raichlen, David A.·2016·Journal of Experimental Biology

Abstract

Human bipedal locomotion is characterized by a habitual heel-strike (HS) plantigrade gait, yet the significance of walking foot-posture is not well understood. To date, researchers have not fully investigated the costs of non-heel-strike (NHS) walking. Therefore, we examined walking speed, walk-to-run transition speed, estimated locomotor costs (lower limb muscle volume activated during walking), impact transient (rapid increase in ground force at touchdown) and effective limb length (ELL) in subjects (n=14) who walked at self-selected speeds using HS and NHS gaits. HS walking increases ELL compared with NHS walking since the center of pressure translates anteriorly from heel touchdown to toe-off. NHS gaits led to decreased absolute walking speeds (P=0.012) and walk-to-run transition speeds (P=0.0025), and increased estimated locomotor energy costs (P<0.0001) compared with HS gaits. These differences lost significance after using the dynamic similarity hypothesis to account for the effects of foot landing posture on ELL. Thus, reduced locomotor costs and increased maximum walking speeds in HS gaits are linked to the increased ELL compared with NHS gaits. However, HS walking significantly increases impact transient values at all speeds (P<0.0001). These trade-offs may be key to understanding the functional benefits of HS walking. Given the current debate over the locomotor mechanics of early hominins and the range of foot landing postures used by nonhuman apes, we suggest the consistent use of HS gaits provides key locomotor advantages to striding bipeds and may have appeared early in hominin evolution.

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  • The use of a flatter foot contact and absence of a clear heel strike may affect walking economy by reducing the total distance travelled by the body's centre of mass during a given step and/or increasing the amount of mechanical work needed to accelerate the body's centre of mass between steps.

    “The use of a flatter foot contact and absence of a clear heel strike may affect walking economy, as it may reduce the total distance travelled by the body's centre of mass during a given step (Webber & Raichlen, 2016) and/or increase the amount of mechanical work needed to accelerate the body's centre of mass between steps (Adamczyk & Kuo, 2013; Holowka & Lieberman, 2018).”
  • The data presented here and in previous studies suggest that some of the kinematic features that are believed to give humans their edge in walking economy, such as use of extended legs and heel strikes, are compromised on irregular natural terrains.

    “However, the data presented here and in previous studies suggest that some of the kinematic features that are believed to give humans their edge in walking economy, such as use of extended legs and heel strikes (Pontzer & Kamilar, 2009; Webber & Raichlen, 2016), are compromised on irregular natural terrains.”

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Research works printed but not linked: 72

  1. Alexander, 1976, Mechanics of bipedal locomotion, Perspect. Exp. Biol., 1, 493, 10.1016/b978-0-08-018767-9.50047-0 · DOI
  2. Alexander, 1989, Optimization and gaits in the locomotion of vertebrates, Physiol. Rev., 69, 1199, 10.1152/physrev.1989.69.4.1199 · DOI · Paper page
  3. Alexander, 1983, A dynamic similarity hypothesis for the gaits of quadrupedal mammals, J. Zool., 201, 135, 10.1111/j.1469-7998.1983.tb04266.x · DOI
  4. Biewener, 2004, Muscle mechanical advantage of human walking and running: implications for energy cost, J. Appl. Physiol., 97, 2266, 10.1152/japplphysiol.00003.2004 · DOI · Paper page
  5. Bramble, 2004, Endurance running and the evolution of Homo, Nature, 432, 345, 10.1038/nature03052 · DOI
  6. Carey, 2005, The metabolic costs of “bent-hip, bent-knee” walking in humans, J. Hum. Evol., 48, 25, 10.1016/j.jhevol.2004.10.001 · DOI · Paper page
  7. Carrier, 1984, The energetic paradox of human running and hominid evolution, Curr. Anthropol., 25, 483, 10.1086/203165 · DOI
  8. Cunningham, 2010, The influence of foot posture on the cost of transport in humans, J. Exp. Biol., 213, 790, 10.1242/jeb.038984 · DOI
  9. Desilva, 2012, The primitive aspects of the foot and ankle of Australopithecus sediba, Am. J. Phys. Anthropol., 147
  10. Dirks, 2010, Geological setting and age of Australopithecus sediba from southern africa, Science, 328, 205, 10.1126/science.1184950 · DOI
  11. Divert, 2005, Mechanical comparison of barefoot and shod running, Int. J. Sports Med., 26, 593, 10.1055/s-2004-821327 · DOI
  12. Divert, 2008, Barefoot-shod running differences: shoe or mass effect?, Int. J. Sports Med., 29, 512, 10.1055/s-2007-989233 · DOI
  13. Elftman, 1935, Chimpanzee and human feet in bipedal walking, Am. J. Phys. Anthropol., 20, 69, 10.1002/ajpa.1330200109 · DOI
  14. Farinatti, 2010, Walk-run transition in young and older adults: with special reference to the cardio-respiratory responses, Eur. J. Appl. Physiol., 109, 379, 10.1007/s00421-010-1366-1 · DOI
  15. Foster, 2013, Muscle force production during bent-knee, bent-hip walking in humans, J. Hum. Evol., 65, 294, 10.1016/j.jhevol.2013.06.012 · DOI · Paper page
  16. Franz, 2012, Metabolic cost of running barefoot versus shod, Med. Sci. Sports Exerc., 44, 1519, 10.1249/MSS.0b013e3182514a88 · DOI
  17. Gebo, 1992, Plantigrady and foot adaptation in African apes: implications for hominid origins, Am. J. Phys. Anthropol., 89, 29, 10.1002/ajpa.1330890105 · DOI
  18. Gray, 1944, Studies in the mechanics of the tetrapod skeleton, J. Exp. Biol., 20, 88, 10.1242/jeb.20.2.88 · DOI
  19. Griffin, 2004, Biomechanical and energetic determinants of the walk-trot transition in horses, J. Exp. Biol., 207, 4215, 10.1242/jeb.01277 · DOI
  20. Hildebrand, 1998, Analysis of Vertebrate Structure
  21. Holliday, 1997, Postcranial evidence of cold adaptation in European Neandertals, Am. J. Phys. Anthropol., 104, 245, 10.1002/(SICI)1096-8644(199710)104:2<245::AID-AJPA10>3.0.CO;2-# · DOI
  22. Hreljac, 2008, The relationship between joint kinetic factors and the walk-run gait transition speed during human locomotion, J. Appl. Biomech., 24, 149, 10.1123/jab.24.2.149 · DOI
  23. Jungers, 1988, Lucy's length: stature reconstruction in Australopithecus afarensis (A.L.288-1) with implications for other small-bodied hominids, Am. J. Phys. Anthropol., 76, 227, 10.1002/ajpa.1330760211 · DOI
  24. Jungers, 2009, The foot of Homo floresiensis, Nature, 459, 81, 10.1038/nature07989 · DOI
  25. Jungers, 2009, Descriptions of the lower limb skeleton of Homo floresiensis, J. Hum. Evol., 57, 538, 10.1016/j.jhevol.2008.08.014 · DOI
  26. Kram, 2012, Is barefoot running more economical?, Int. J. Sports Med., 33, 10.1055/s-0032-1301923 · DOI
  27. Kram, 1990, Energetics of running: a new perspective, Nature, 346, 265, 10.1038/346265a0 · DOI · Paper page
  28. Kram, 1997, Effect of reduced gravity on the preferred walk-run transition speed, J. Exp. Biol., 200, 821, 10.1242/jeb.200.4.821 · DOI
  29. Kuo, 2001, A simple model of bipedal walking predicts the preferred speed-step length relationship, J. Biomech. Eng., 123, 264, 10.1115/1.1372322 · DOI
  30. Latimer, 1989, The calcaneus of australopithecus-afarensis and its implications for the evolution of bipedality, Am. J. Phys. Anthropol., 78, 369, 10.1002/ajpa.1330780306 · DOI
  31. Leakey, 1981, Tracks and tools, Philos. Trans. R. Soc. Lond. B Biol. Sci., 292, 95, 10.1098/rstb.1981.0017 · DOI
  32. Leakey, 1979, Pliocene footprints in the Laetolil Beds at Laetoli, northern Tanzania, Nature, 278, 317, 10.1038/278317a0 · DOI
  33. Lieberman, 2012, What we can learn about running from barefoot running: an evolutionary medical perspective, Exerc. Sport Sci. Rev., 40, 63, 10.1097/JES.0b013e31824ab210 · DOI
  34. Lieberman, 2010, Foot strike patterns and collision forces in habitually barefoot versus shod runners, Nature, 463, 531, 10.1038/nature08723 · DOI
  35. Miller, 1998, Locomotor advantages of Neandertal skeletal morphology at the knee and ankle, J. Biomech., 31, 355, 10.1016/S0021-9290(98)00031-1 · DOI
  36. Morwood, 2004, Archaeology and age of a new hominin from Flores in eastern Indonesia, Nature, 431, 1087, 10.1038/nature02956 · DOI
  37. Németh, 1985, In vivo moment arm lengths for hip extensor muscles at different angles of hip flexion, J. Biomech., 18, 129, 10.1016/0021-9290(85)90005-3 · DOI
  38. Pontzer, 2005, A new model predicting locomotor cost from limb length via force production, J. Exp. Biol., 208, 1513, 10.1242/jeb.01549 · DOI
  39. Pontzer, 2007, Effective limb length and the scaling of locomotor cost in terrestrial animals, J. Exp. Biol., 210, 1752, 10.1242/jeb.002246 · DOI
  40. Pontzer, 2007, Predicting the energy cost of terrestrial locomotion: a test of the LiMb model in humans and quadrupeds, J. Exp. Biol., 210, 484, 10.1242/jeb.02662 · DOI
  41. Pontzer, 2009, The metabolic cost of walking in humans, chimpanzees, and early hominins, J. Hum. Evol., 56, 43, 10.1016/j.jhevol.2008.09.001 · DOI
  42. Pontzer, 2014, Bipedal and quadrupedal locomotion in chimpanzees, J. Hum. Evol., 66, 64, 10.1016/j.jhevol.2013.10.002 · DOI
  43. Raichlen, 2008, The effects of gravity on human walking: a new test of the dynamic similarity hypothesis using a predictive model, J. Exp. Biol., 211, 2767, 10.1242/jeb.020073 · DOI · Paper page
  44. Raichlen, 2010, Laetoli footprints preserve earliest direct evidence of human-like bipedal biomechanics, PLoS ONE, 5, e9769, 10.1371/journal.pone.0009769 · DOI
  45. Raichlen, 2011, Calcaneus length determines running economy: Implications for endurance running performance in modern humans and Neandertals, J. Hum. Evol., 60, 299, 10.1016/j.jhevol.2010.11.002 · DOI
  46. Ren, 2010, Integration of biomechanical compliance, leverage, and power in elephant limbs, Proc. Natl. Acad. Sci., 107, 7078, 10.1073/pnas.0911396107 · DOI
  47. Roberts, 1998, Energetics of bipedal running. II. Limb design and running mechanics, J. Exp. Biol., 201, 2753, 10.1242/jeb.201.19.2753 · DOI
  48. Roberts, 1998, Energetics of bipedal running. I. Metabolic cost of generating force, J. Exp. Biol., 201, 2745, 10.1242/jeb.201.19.2745 · DOI
  49. Rolian, 2009, Walking, running and the evolution of short toes in humans, J. Exp. Biol., 212, 713, 10.1242/jeb.019885 · DOI
  50. Rugg, 1990, In vivo moment arm calculations at the ankle using magnetic resonance imaging (MRI), J. Biomech., 23, 495, 10.1016/0021-9290(90)90305-M · DOI
  51. Schmitt, 1998, Neandertal's calcaneal variability. Comparison with modern humans, Bull. Mém. Soc. Anthropol. Paris, 10, 273, 10.3406/bmsap.1998.2518 · DOI
  52. Schmitt, 1995, Heel contact as a function of substrate type and speed in primates, Am. J. Phys. Anthropol., 96, 39, 10.1002/ajpa.1330960105 · DOI
  53. Sellers, 2005, Stride lengths, speed and energy costs in walking of Australopithecus afarensis: using evolutionary robotics to predict locomotion of early human ancestors, J. R. Soc. Interface, 2, 431, 10.1098/rsif.2005.0060 · DOI
  54. Sockol, 2007, Chimpanzee locomotor energetics and the origin of human bipedalism, Proc. Natl. Acad. Sci. USA, 104, 12265, 10.1073/pnas.0703267104 · DOI · Paper page
  55. Spoor, 1994, Implications of early hominid labyrinthine morphology for evolution of human bipedal locomotion, Nature, 369, 645, 10.1038/369645a0 · DOI
  56. Steudel, 1995, Does limb length predict the relative energetic cost of locomotion in mammals?, J. Zool., 235, 501, 10.1111/j.1469-7998.1995.tb01765.x · DOI
  57. Taylor, 1994, Relating mechanics and energetics during exercise, Adv. Vet. Sci. Comp. Med., 38A, 181
  58. Thorstensson, 1987, Adaptations to changing speed in human locomotion-speed of transition between walking and running, Acta Physiol. Scand., 131, 211, 10.1111/j.1748-1716.1987.tb08228.x · DOI
  59. Trinkaus, 1975, A Functional Analysis of the Neanderthal Foot
  60. Trinkaus, 1981, Neanderthal limb proportios and cold adaptation, 187
  61. Trinkaus, 1983, Functional-aspects of neandertal pedal remains, Foot Ankle Int., 3, 377, 10.1177/107110078300300606 · DOI
  62. Trinkaus, 1996, Neandertal pedal proximal phalanges: diaphyseal loading patterns, J. Hum. Evol., 30, 399, 10.1006/jhev.1996.0035 · DOI
  63. Usherwood, 2005, Why not walk faster?, Biol. Lett., 1, 338, 10.1098/rsbl.2005.0312 · DOI
  64. Usherwood, 2012, The human foot and heel–sole–toe walking strategy: a mechanism enabling an inverted pendular gait with low isometric muscle force?, J. R. Soc. Interface, 9, 2396, 10.1098/rsif.2012.0179 · DOI
  65. Vereecke, 2003, Dynamic plantar pressure distribution during terrestrial locomotion of bonobos (Pan paniscus), Am. J. Phys. Anthropol., 120, 373, 10.1002/ajpa.10163 · DOI
  66. Vereecke, 2005, Functional analysis of the gibbon foot during terrestrial bipedal walking: plantar pressure distributions and three-dimensional ground reaction forces, Am. J. Phys. Anthropol., 128, 659, 10.1002/ajpa.20158 · DOI
  67. Visser, 1990, Length and moment arm of human leg muscles as a function of knee and hip-joint angles, Eur. J. Appl. Physiol. Occup. Physiol., 61, 453, 10.1007/BF00236067 · DOI
  68. Wang, 2004, Comparison of inverse-dynamics musculo-skeletal models of AL 288-1 Australopithecus afarensis and KNM-WT 15000 Homo ergaster to modern humans, with implications for the evolution of bipedalism RID A-6670-2010, J. Hum. Evol., 47, 453, 10.1016/j.jhevol.2004.08.007 · DOI
  69. White, 1987, Hominid footprints at laetoli-facts and interpretations, Am. J. Phys. Anthropol., 72, 485, 10.1002/ajpa.1330720409 · DOI
  70. Winter, 2009, Biomechanics and Motor Control of Human Movement, 10.1002/9780470549148 · DOI
  71. Wright, 2001, The application of ground force explains the energetic cost of running backward and forward, J. Exp. Biol., 204, 1805, 10.1242/jeb.204.10.1805 · DOI
  72. Zipfel, 2011, The foot and ankle of Australopithecus sediba, Science, 333, 1417, 10.1126/science.1202703 · DOI

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Metadata

Authors
Webber, J. T., & Raichlen, D. A.
Affiliations
School of Anthropology, University of Arizona, Tucson, AZ 85721, USA
Venue
Journal of Experimental Biology
Published
2016-12-01
Volume
219
Issue
23
Pages
3729-3737
ISSN
  • 1477-9145
  • 0022-0949
Cited by
37
References
0
Language
en
Funders
University of Arizona

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