Foot & Ankle International

The changes the human foot undergoes during its development are described as indications of structural and functional changes which might have occurred during the evolution of the vertebrate hind foot. The human foot, when compared to other vertebrate feet, appears to be a relatively minor modification of the plantigrade hind foot of the shrew-like insectivores that gave rise to all of the mammals. There is no developmental evidence to indicate that the first toe of the human foot was ever prehensile. Thus, the generally accepted evolutionary view, which depicts the feet of the early ancestors of modern human beings as having ape-like prehensile first toes, is seriously questioned.

The living primates are a highly diverse group of essentially arboreal animals whose feet are variously adapted for grasping, climbing, and leaping in trees. One of the most remarkable aspects of the anatomical variation in the feet of the extant primates is that this diversity can be arranged in a graduated sequence ranging from the primitive transtarsal-opposition type of grasping foot found in the lemurs, through the specialized transmeta-tarsal-adduction type that characterizes the higher primates, to'the unique nongrasping foot of humans. The comparative study of this graded series makes it possible, without recourse to the fossil record, to appreciate the adaptive and functional stages through which the human foot passed in its evolution. It is hypothesized that the initial stage of human erect posture was characterized by a foot which was adapted to both hallucial grasping and short distance bipedal walking. In many respects, the structure of the foot of this primitive human was probably similar to that of the living highland gorilla.

When compared with the feet of our closest living relatives, the African apes, the human foot is characterized by two major evolutionary specializations: the longitudinal and transverse arches, and the parallel first and second metatarsals. These two morphological adaptations, together with several associated specializations, are the structural basis of the bipedal human foot. Reconstructing the evolutionary and adaptive history of these specializations creates a better understanding of some of the more common clinical podiatric disorders. The modern human foot is structurally so well adapted to prolonged bipedal walking and standing that even slight deviations from its evolutionarily established pattern will produce debilitating clinical manifestations. In most cases, successful treatment of such disorders involves restoration of the foot's basic adaptive configuration.

The fossil evidence relating to the evolution of the primate foot is reviewed and evaluated. Many of the characteristic features of the primate foot had evolved by the early Tertiary over 40 million years ago. Probably the most significant of these developments was the progressive migration of the talus to a position over the calcaneum. These morphological features are followed through the Miocene hominoid genera from East Africa, Europe, and South Asia. While some features of Miocene hominoids, especially those relating to climbing abilities, are still evident in the predominantly bipedal earliest hominids of the Plio-Pleistocene, there is no evidence yet from the Miocene of the first stages in the evolution of that bipe-dalism.

The human foot serves a dual role during locomotion. It functions at times as a mobile structure and at times as a rigid lever. The human foot shows the hallmarks of an arboreal heritage wherein the foot was primarily a grasping organ. Over the course of the human career the human foot has evolved an elaborate plantar aponeurosis, strong plantar ligaments, longitudinal arches, an enlarged musculus flexor accessorius, an adducted (non-opposable) hallux, a remodeled calcaneocuboid joint, a long tarsus, and shortened toes (II to V). Comparisons of the chimpanzee and human foot allow us to reconstruct the pathway of foot evolution. Fossil foot bones of Homo habilis, dated at 1.76 million years, are remarkably like those of modern humans. Foot bones from Hadar, dated at around 3.5 million years, are remarkably chimpanzee-like, with only incipient human traits. The surprising chimpanzee-like qualities of the Hadar fossils strongly support the use of living apes as models of ancestral pongid-hominid morphotypes.

The fossil foot remains of the Neandertals (immediate predecessors of anatomically modern humans in Europe and western Asia between about 100,000 and 35,000 to 40,000 years ago) indicate an overall pedal configuration for these prehistoric humans that was largely indistinguishable from that of modern humans. They had fully adducted halluces, longitudinal and transverse pedal arches, compact posterior tarsal regions, slight valgus deviation of the hallux, and abbreviation of the lateral toes. Their pedal remains differed from those of modern humans only in their tendency to be more robust and their relatively short proximal hallucial phalanges and associated elongated distal hallucial phalanges. The former is related to their habitual levels of physical activity, which were greater than those of modern humans, and the latter is a pleiotropic effect of a biomechanical adaptation in the pollex for strength during manipulation.

Theories about the functions of the foot muscles have centered on their role in arch support. Previous anatomical and electromyographic studies (reviewed herein) have demonstrated that the arches are normally maintained by bones and ligaments. This study reports an electromyographic investigation of five foot muscles (flexor digito-rum longus, flexor digitorum brevis, flexor accessorius, abductor hallucis, and abductor digiti quinti) conducted on four humans. The three toe flexors act together to resist extension of the toes during the stance phase of locomotion. Despite the large flexor accessorius in humans, neither this muscle nor the flexor digitorum brevis are preferentially recruited over the flexor digitorum lon-gus for any normal posture or locomotion. The abductors affect the mediolateral distribution of pressure by positioning the forefoot. We suggest that the foot muscles play an important role in positioning of the forces on the foot in both posture and locomotion. Future electromyographic experiments on human and ape foot muscles in conjunction with detailed studies of early hominid fossils promise to elucidate the pathways of human locomotor evolution.