Human skeletal changes due to bipedalism
The evolution of human bipedalism approximately four million years ago1 has led to morphological alterations to the human skeleton including changes to the arrangement and size of the bones of the foot, hip size and shape, knee size, leg length, and the shape and orientation of the vertebral column. The evolutionary factors that produced these changes have been the subject of several theories.
The human foot evolved to act as a platform to support the entire weight of the body, rather than acting as a grasping structure, as it did in early hominids. Humans therefore have smaller toes than their bipedal ancestors. This includes a non-opposable hallux, which is relocated in line with the other toes.2 Moreover, humans have a foot arch rather than flat feet.2 When non-human hominids walk upright, weight is transmitted from the heel, along the outside of the foot, and then through the middle toes while a human foot transmits weight from the heel, along the outside of the foot, across the ball of the foot and finally through the big toe. This transference of weight contributes to energy conservation during locomotion.13
Modern human hip joints are larger than in quadrupedal ancestral species to better support the greater amount of body weight passing through them,2 as well as having a shorter, broader shape. This alteration in shape brought the vertebral column closer to the hip joint, providing a stable base for support of the trunk while walking upright.4 Also, because bipedal walking requires humans to balance on a relatively unstable ball and socket joint, the placement of the vertebral column closer to the hip joint allows humans to invest less muscular effort in balancing.2 Change in the shape of the hip may have led to the decrease in the degree of hip extension, an energy efficient adaptation.15
Human knee joints are enlarged for the same reason as the hip – to better support an increased amount of body weight.2 The degree of knee extension (the angle between the thigh and shank in a walking cycle) has decreased. The changing pattern of the knee joint angle of humans shows a small extension peak, called the “double knee action,” in the midstance phase. Double knee action decreases energy lost by vertical movement of the center of gravity.1 Humans walk with their knees kept straight and the thighs bent inward so that the knees are almost directly under the body, rather than out to the side, as is the case in ancestral hominids. This type of gait also aids balance.2
An increase in leg length since the evolution of bipedalism changed how leg muscles functioned in upright gait. In humans the "push" for walking comes from the leg muscles acting at the ankle. A longer leg allows the use of the natural swing of the limb so that, when walking, humans do not need to use muscle to swing the other leg forward for the next step.2 As a consequence, since the human forelimbs are not needed for locomotion, they are instead optimized for carrying, holding, and manipulating objects with great precision.6 Having long hindlimbs and short forelimbs allows humans to walk upright, while orangutans and gibbons had the adaptation of longer arms to swing on branches.7 Apes can stand on their hindlimbs, but they cannot do so for long periods of time without getting tired. This is because they haven’t adapted their femur for bipedalism. Apes have vertical femurs, while humans have femurs that are slightly angled medially from the hip to the knee. This adaptation allows our knees to be closer together and under the body’s center of gravity. This permits humans to lock their knees and stand up straight for long periods of time without much effort from the muscles.8
The human skull is balanced on the vertebral column: The foramen magnum is located inferiorly under the skull, which puts much of the weight of the head behind the spine. Furthermore, the flat human face helps to maintain balance on the occipital condyles. Because of this, the erect position of the head is possible without the prominent supraorbital ridges and the strong muscular attachments found in, for example, apes. As a result, in humans the muscles of the forehead (the occipitofrontalis) are only used for facial expressions.6
The vertebral column of humans takes a forward bend in the lumbar (lower) region and a backward bend in the thoracic (upper) region. Without the lumbar curve, the vertebral column would always lean forward, a position that requires much more muscular effort for bipedal animals. With a forward bend, humans use less muscular effort to stand and walk upright.4 Together the lumbar and thoracic curves bring the body's center of gravity directly over the feet.2 Also, the degree of body erection (the angle of body incline to a vertical line in a walking cycle) is significantly smaller1 to conserve energy.
Even with much anatomical modification, some features of the human skeleton remain poorly adapted to bipedalism, leading to negative implications prevalent in humans today. The lower back and knee joints are plagued by osteological malfunction, lower back pain being a leading cause of lost working days,9 because the joints support more weight. Arthritis has been a problem since hominids became bipedal: scientists have discovered its traces in the vertebrae of prehistoric hunter-gatherers.9 Physical constraints have made it difficult to modify the joints for further stability while maintaining efficiency of locomotion.2
- Kondō, Shirō (1985). Primate morphophysiology, locomotor analyses, and human bipedalism. Tokyo: University of Tokyo Press. ISBN 4-13-066093-4.
- Aiello,Leslie and Christopher Dean (1990). An Introduction to Human Evolutionary Anatomy. Oxford: Elsevier Academic Press. ISBN 0-12-045591-9.
- Latimer, B., & Lovejoy, C.O. (1989). "The Calcaneus of Australopithecus afarensis and its implications for the Evolution of Bipedality". American Journal of Physical Anthropology 78 (3): 369–386. doi:10.1002/ajpa.1330780306. PMID 2929741.
- Wang, W.; Crompton, R.H.; Carey, T.S.; Günther, M.M.; Li, Y.; Savage, R.; Sellers, W.I. (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". Journal of Human Evolution 47 (6): 453–478. doi:10.1016/j.jhevol.2004.08.007. PMID 15566947. Retrieved 2008-04-02.
- Lovejoy, C. Owen (1988). "Evolution of Human walking". Scientific American 259 (5): 82–89. doi:10.1038/scientificamerican1188-118.
- Saladin, Kenneth S. (2003). 3rd, ed. Anatomy & Physiology: The Unity of Form and Function. McGraw-Hill. pp. 286–287. ISBN 0-07-110737-1.
- Thorpe, S. "Origin of Human Bipedalism As an Adaptation for Locomotion on Flexible Branches." AAAS Science. AAAS Science, 18 May 2007. Web. 08 Dec. 2010. <http://www.sciencemag.org/content/316/5829/1328.full>.
- Saladin, Kenneth S. "Chapter 8." Anatomy & Physiology: the Unity of Form and Function. 5th ed. Dubuque: McGraw-Hill, 2010. 281. Print.
- Jacob C. Koella; Stearns, Stephen K. (2008). Evolution in Health and Disease. Oxford University Press, USA. ISBN 0-19-920746-1.
- Grabowski MW, Polk JD, Roseman CC. (2011). "Divergent patterns of integration and reduced constraint in the human hip and the origins of bipedalism.". Evolution 65 (5): 1336–56. doi:10.1111/j.1558-5646.2011.01226.x. PMID 21521191.
- Crompton RH, Sellers WI, Thorpe SK. (2010). "Arboreality, terrestriality and bipedalism.". Philos Trans R Soc Lond B Biol Sci. 365 (1556): 3301–14. doi:10.1098/rstb.2010.0035. PMC 2981953. PMID 20855304.