The Olduvai Hominid 8 foot: Adult or subadult?

Olduvai Hominid 8 (OH 8), an articulating set of fossil hominin tarsal and metatarsal bones, is critical to interpretations of the evolution of hominin pedal morphology and bipedal locomotion. It has been suggested that OH 8 may represent the foot of a subadult and may be associated with the OH 7 mandible, the type specimen of Homo habilis. This assertion is based on the presence of what may be unfused distal metatarsal epiphyses. Accurately assessing the skeletal maturity of the OH 8 foot is important for interpretations of the functional morphology and locomotor behavior of Plio-Pleistocene hominins. In this study, we compare metatarsal fusion patterns and internal bone morphology of the lateral metatarsals among subadult hominines (85 modern humans, 48 Pan, and 25 Gorilla) to assess the likelihood that OH 8 belonged to either an adult or subadult hominin. Our results suggest that if OH 8 is indeed from a subadult, then it displays a metatarsal developmental pattern that is unobserved in our comparative sample. In OH 8, the fully fused base of the first metatarsal and the presence of trabecular bone at the distal ends of the second and third metatarsal shafts make it highly improbable that it belonged to a subadult, let alone a subadult that matches the developmental age of the OH 7 mandible. In total, the results of this study suggest that the OH 8 foot most likely belonged to an adult hominin.     

Metatarsal fusion pattern and developmental morphology of the Olduvai Hominid 8 foot: Evidence of adolescence

The morphology of the Olduvai Hominid (OH) 8 foot and the sequence of metatarsal epiphyseal fusion in modern humans and chimpanzees support the hypothesis that OH 8 belonged to an individual of approximately the same relative age as the OH 7 subadult, the holotype of Homo habilis. Modern humans and chimpanzees exhibit a variety of metatarsal epiphyseal fusion patterns, including one identical to that observed in OH 8 in which metatarsal 1 fuses before metatarsals 2-5. More than the metatarsal fusion sequence, however, the principal evidence of the youthful age of OH 8 lies in the morphology of metatarsals 1, 2, and 3. Because both OH 8 and OH 7 come from the same stratum at the FLK NN type site, the most parsimonious explanation of the OH 8 and OH 7 data is that this material belonged to the same individual, as originally proposed by Louis Leakey. The proposition that OH 8 belonged to an adult is unsupported by morphology, including radiographic evidence, and the fusion sequences in human and chimpanzee skeletal material reported here and in the literature.     

足籽骨的放射片研究

本文对山东枣庄市916人足放射片进行了足籽骨的观察与测量。发现每足有1-7个籽骨,其中二个者占77.8％,三个者占16.9％;籽骨位于跖骨头下方者占94.5％,位于趾间关节下方者占5.5％;各跖骨头籽骨出现率:Ⅰ占99.9％、Ⅱ占 2.2％、Ⅲ占0.3％、Ⅳ占0.5％、Ⅴ占6.4％;(足母)趾趾间关节籽骨占12.6％;二分及三分籽骨出现率占3.9％,明显低于欧美人。此外,本文认为籽骨是在先天籽骨原基基础上,加以后天运动的影响而形成。     

A complete second metatarsal (StW 89) from Sterkfontein Member 4, South Africa

The functional anatomy of the hominin foot has played a crucial role in studies of locomotor evolution in human ancestors and extinct relatives. However, foot fossils are rare, often isolated, and fragmentary. Here, we describe a complete hominin second metatarsal (StW 89) from the 2.0-2.6 million year old deposits of Member 4, Sterkfontein Cave, South Africa. Like many other fossil foot bones, it displays a mosaic of derived human-like features and primitive ape-like features. StW 89 possesses a domed metatarsal head with a prominent sulcus, indicating dorsiflexion at the metatarsophalangeal joint during bipedal walking. However, while the range of motion at the metatarsophalangeal joint is human-like in dorsiflexion, it is ape-like in plantarflexion. Furthermore, StW 89 possesses internal torsion of the head, an anatomy decidedly unlike that found in humans today. Unlike other hominin second metatarsals, StW 89 has a dorsoplantarly gracile base, perhaps suggesting more midfoot laxity. In these latter two anatomies, the StW 89 second metatarsal is quite similar to the recently described second metatarsal of the partial foot from Burtele, Ethiopia. We interpret this combination of anatomies as evidence for a low medial longitudinal arch in a foot engaged in both bipedal locomotion, but also some degree of pedal, and perhaps even hallucal, grasping. Additional fossil evidence will be required to determine if differences between this bone and other second metatarsals from Sterkfontein reflect normal variation in an evolving lineage, or taxonomic diversity.     

Cortical Structure of Hallucal Metatarsals and Locomotor Adaptations in Hominoids

Diaphyseal morphology of long bones, in part, reflects in vivo loads experienced during the lifetime of an individual. The first metatarsal, as a cornerstone structure of the foot, presumably expresses diaphyseal morphology that reflects loading history of the foot during stance phase of gait. Human feet differ substantially from those of other apes in terms of loading histories when comparing the path of the center of pressure during stance phase, which reflects different weight transfer mechanisms. Here we use a novel approach for quantifying continuous thickness and cross-sectional geometric properties of long bones in order to test explicit hypotheses about loading histories and diaphyseal structure of adult chimpanzee, gorilla, and human first metatarsals. For each hallucal metatarsal, 17 cross sections were extracted at regularly-spaced intervals (2.5% length) between 25% and 65% length. Cortical thickness in cross sections was measured in one degree radially-arranged increments, while second moments of area were measured about neutral axes also in one degree radially-arranged increments. Standardized thicknesses and second moments of area were visualized using false color maps, while penalized discriminant analyses were used to evaluate quantitative species differences. Humans systematically exhibit the thinnest diaphyseal cortices, yet the greatest diaphyseal rigidities, particularly in dorsoplantar regions. Shifts in orientation of maximum second moments of area along the diaphysis also distinguish human hallucal metatarsals from those of chimpanzees and gorillas. Diaphyseal structure reflects different loading regimes, often in predictable ways, with human versus non-human differences probably resulting both from the use of arboreal substrates by non-human apes and by differing spatial relationships between hallux position and orientation of the substrate reaction resultant during stance. The novel morphological approach employed in this study offers the potential for transformative insights into form-function relationships in additional long bones, including those of extinct organisms (e.g., fossils).     

Human foot bones from Klasies River main site, South Africa

The caves at Klasies River contain abundant archaeological evidence relating to human evolution in the late Pleistocene of southern Africa. Along with Middle Stone Age artifacts, animal bones, and other food waste, there are hominin cranial fragments, mandibles with teeth, and a few postcranial remains. Three foot bones can now be added to this inventory. An adult first metatarsal is similar in size and discrete anatomical features to those from Holocene burials in the Cape Province. A complete and well-preserved second metatarsal is especially long and heavy at midshaft in comparison to all Holocene and more recent South African homologues. A large fifth metatarsal is highly distinctive in its morphology. In overall size, these pedal elements resemble specimens from late Pleistocene sites in western Asia, but there are some differences in proportions. The fossils support earlier suggestions concerning a relatively high level of sexual dimorphism in the African Middle Stone Age population. Squatting facets on the two lateral metatarsals appear to indicate a high frequency of kneeling among members of this group. The new postcranial material also underlines the fact that the morphology of particular skeletal elements of some of the 100,000-year-old Klasies River individuals falls outside the range of modern variation.     

天津蓟县桃花园墓地明清时期缠足女性足骨的形态观察

本文对天津蓟县桃花园墓地明清时期101例缠足女性足骨形变情况进行了观察和分析。结果显示,不同个体的同名骨骼形变类型和程度不同,有的个体两侧足骨呈不对称状。跗骨在整体尺寸上缩小,且部分跗骨会产生形变。跖骨和近节趾骨因受缠足外力的影响在形态上会产生剧烈的变化,主要变现为跖骨和近节趾骨纤细、弓弯,关节面改变,其近、远端以及跖骨体和趾骨体的上下径和横径均产生形变。总体而言,跖骨和近节趾骨的形变程度较跗骨而言更大。本文总结了判断某个体是否缠足的依据,特别指出需要同时观察距骨和跟骨的形态改变。鉴于其他疾病(如高弓足、麻风病、风湿性关节炎)也可导致足骨的畸形样貌,在进行个体缠足判定时,需要进行综合成因分析。缠足由文化行为所导致,其足骨形变特征有别于因病理原因导致的足部畸形。功能压力分析能够有效地解释缠足个体足骨形变的成因及过程。     

Earliest complete hominin fifth metatarsal—Implications for the evolution of the lateral column of the foot

StW 114/115, from Sterkfontein, South Africa, is the earliest complete hominin fifth metatarsal. Comparisons of StW 114/115 to modern humans, extant apes, and partial hominin metatarsals AL 333‐13, AL 333‐78, SKX 33380, OH 8, and KNM‐ER 803f reveal a similar morphology in all six fossils consistent with habitual bipedality. Although StW 114/115 possesses some primitive characters, the proximal articular morphology and internal torsion of the head are very human‐like, suggesting a stable lateral column and the likely presence of lateral longitudinal and transverse tarsal arches. We conclude that, at least in the lateral component of the foot of the StW 114/115 individual, the biomechanical pattern is very similar to that of modern humans. This, however, may not have been the case in the medial column of the foot, as a mosaic pattern of hominin foot evolution and function has been suggested. The results of this study may support the hypothesis of an increased calcaneo‐cuboid stability having been an early evolutionary event in the history of terrestrial bipedalism. Am J Phys Anthropol 2009. © 2009 Wiley‐Liss, Inc.     

Brief communication: radiographic study of metatarsal one basal epiphyseal fusion: a note of caution on age determination

This study examines radiographs of first metatarsals of 131 individuals from age 17-88 years to determine whether internal basal epiphyseal lines may be visible past the age of metatarsal fusion, which usually occurs between 14 and 16 years of age (Scheuer and Black: The juvenile skeleton. San Diego: Elsevier Academic Press,2004). In 29% (38 out of 131) of the radiographed first metatarsals (MT1s) the basal epiphyseal scar is visible, including in one individual who was 80 years old. Statistically, there was no relationship between the loss of the epiphyseal scar and age. Thus, the presence of the epiphyseal scar does not necessarily indicate subadult age. These data suggest that OH 8's radiographically visible basal epiphyseal line has no bearing on whether it is a subadult or not.     

天津蓟州区桃花园墓地明清时期缠足女性的足骨形变

本文通过量化方法,对桃花园墓地明清时期101例女性足骨形变方式、程度、对称性等进行了系统分析。研究结果表明,双侧足骨形变总体上是对称的。缠足对跗骨的影响主要在于整体尺寸缩小和关节面改变。第1跖骨除整体尺寸缩小外,还存在诸多明显的骨体形态改变;第2、3跖骨头部和底部尺寸缩小,但形变不大。第3至第5近节趾骨骨体长度和高度的侧别差异明显,特别是第3近节趾骨双侧整体不对称。该群体至少在18岁时已经缠足,25岁以后足骨已发生形变,35岁之后形变较明显。部分个体足骨形变程度较轻,其生前可能仅束足纤直,未经裹弯。足骨形变程度与陪葬品多寡并无相关性,其形变差异很可能与身体疾病、劳作需要、缠足方法或缠足观念差异有关。     

Determination of adult stature from metatarsal length

The results of a study to determine the value of foot bones in reconstructing stature are presented. The data consist of length measurements taken on all ten metatarsals as well as on cadaver length from a sample of 130 adults of documented race, sex, stature, and, in most cases, age. Significant correlation coefficients (.58-.89) are shown between known stature and foot bone lengths. Simple and multiple regression equations computed from the length of each of these bones result in standard errors of estimated stature ranging from 40-76 mm. These errors are larger than those for stature calculated from complete long bones, but are approximately the same magnitude for stature calculated from metacarpals and fragmentary long bones. Given that metatarsals are more likely to be preserved unbroken than are long bones and given the ease with which they are accurately measured, the formulae presented here should prove useful in the study of historic and even prehistoric populations.     

Os intermetatarseum: A heritable accessory bone of the human foot

The os intermetatarseum is a relatively common accessory bone of the human foot located at the tarsometatarsal border between the first and second metatarsals. It can occur as an independent ossicle or as an osseous spur projecting from the proximal ends of the first two metatarsals, or the distal end of the first cuneiform. To determine the frequency of this congenital defect in native North American groups and East Asians from Japan, the skeletons of 846 Native Americans and 125 modern Japanese and Ainu were examined for the presence of the os intermetatarseum. The North American skeletons are from archaeological sites in various parts of Canada and the United States, including the Arctic coast, the Subarctic, the Aleutian Islands, the Northern Plains, the Illinois River area, and the Southwest. Overall frequencies ranged from no occurrence among the Arctic samples to 8% of individuals from Pecos Pueblo. Second metatarsal spurs occurred in particularly high frequencies among American Indians, whereas the East Asians were only found to have os intermetatarseums associated with the first cuneiform. Because pedigrees have shown the os intermetatarseum to be an inherited defect, its high frequency among some Native Americans may be due, in part, to a higher degree of genetic relatedness among the individuals in the North American samples than among the relatively modern East Asians. Am J Phys Anthropol 107:199–209, 1998. © 1998 Wiley-Liss, Inc.     

New foot remains from the Gran Dolina-TD6 Early Pleistocene site (Sierra de Atapuerca, Burgos, Spain)

This paper presents and describes new foot fossils from the species Homo antecessor, found in level TD6 of the site of Gran Dolina (Sierra de Atapuerca, Burgos, Spain). These new fossils consist of an almost complete left talus (ATD6-95) and the proximal three-quarters of a right fourth metatarsal (ATD6-124). The talus ATD6-95 is tentatively assigned to Hominin 10 of the TD6 sample, an adult male specimen with which the second metatarsal ATD6-70+107 (already published) is also tentatively associated. Analysis of these fossils and other postcranial remains has made possible to estimate a stature similar to those of the specimens from the Middle Pleistocene site of Sima de los Huesos (Sierra de Atapuerca, Burgos, Spain). The morphology of the TD6 metatarsals does not differ significantly from that of modern humans, Neanderthals and the specimens from Sima de los Huesos. Talus ATD6-95, however, differs from the rest of the comparative samples in being long and high, having a long and wide trochlea, and displaying a proportionally short neck.     

Hand osteoarthritis and bone loss: Is there an inverse relationship?

An inverse relationship between osteoarthritis (OA) and bone loss has been supported in clinical research, but there has been little research on bioarchaeological skeletal remains. The current study examines 115 adults from a prehistoric hunter–gatherer population to aid in determining whether hand OA and bone loss are negatively correlated. OA lipping is scored on a four-point scale on left and right trapezia, MC1s, and MC2s and then analyzed with regard to their relationships with sex, age, right MC2 cortical index, and left and right MC1 robusticity, midshaft circumference, and midshaft diameter values. With sexes and ages combined, higher OA scores are found in individuals with greater midshaft diameters. However, lower cortical indices were found in individuals with higher right MC2 OA scores. The data presented tenuously support that bone loss is lower in individuals with more severe osteoarthritis, but age-related changes in bone deposition may make cortical index and other external shaft dimensions an unsuitable variable to examine this relationship.     

Design of an implant for first metatarsophalangeal hemi-arthroplasty

This study was designed to examine the three-dimensional geometry of the head of the first metatarsal bone of the foot. Ninety-seven adult first metatarsal head (MTH1) bones were scanned using a laser scanner at 400 dpi. A best-fit ellipsoid was obtained from the articular surfaces of MTH1 for each size group using nonlinear unconstrained optimisation. Average root mean square errors between the articulating surfaces and the optimal fit surfaces of the bone specimens were between 0.29 and 0.42 mm. After classification based on sex and size groups, the profile provided a good fit to individual bones. Consideration of the thickness of cartilage overlying the metatarsal head (MTH) may further improve the fit. The proposed approach provides the basis for a design of an MTH hemi-arthroplasty that has good anatomical congruence with the native joint.     

Repair of bone defect of the lateral forefoot by double segment triangular fibula flap with vascular pedicle: A case report

Objective: To expound the clinical effect of a new operation by transplanting double segment triangular fibula flap with vascular pedicle to repair the forefoot with lateral bone defect, and to study how to improve the operation method in the following stage. Methods: The inclusion criteria: More than 2 phalangeal and metatarsal bones defects of the lateral forefoot, widespread skin and soft tissue defects on pelma and dorsal foot, and destruction of the anterior aspect of foot arch, which seriously affects the foot function. There was one case of clinical application in November 2014. The repairing method is as followed: the harvested vascularized free fibula was cut into 2 segments and then they were folded into a right angle. According to selected control points on the residual metatarsals, an optimal stereo triangular net was constructed. Meanwhile, according to flow-through mode, the free anterolateral thigh flap was incorporated to repair the forefoot and foot arch. Results: Postoperative bone flaps all survived. After a 17-month following up, it was found that the grafted fibular healed well, shape of the foot was good, weight-bearing walking was practical, a slight limp and discomfort with plantar pain existed, sensory recovery reached S3 level and functional recovery of weight-bearing walking by forefoot reached W3 level, comprehensive evaluation was good, and there were wear scar and ulcer on the plantar flap during long-time walking for patients, such results were excellent according to foot function scoring criteria. Conclusion: In this operation the grafted fibula was fold into a triangle according to actual need, which though not completely restores the tridimensional structure of the longitudinal, transverse arches of the lateral foot makes weight-bearing walking possible, besides, its appearance and function is satisfactory. Such an operation has overcome the shortage of non-tridimensional structure of the transverse arch etc. in traditional operations and it should be an ideal operation in repairing serious defects on the lateral forefoot through further improvement.     

The cross-sectional geometry of the hand and foot bones of the hominoidea and its relationship to locomotor behavior

Cheiridia are valuable indicators of positional behavior, as they directly contact the substrate, but systematic comparison of the structural properties of both metacarpals and metatarsals has never been carried out. Differences in locomotor behavior among the great apes (knuckle-walking vs. quadrumanous climbing) can produce biomechanical differences that may be elucidated by the parallel study of cross-sectional characteristics of metacarpals and metatarsals. The aim of this work is to study the cross-sectional geometric properties of these bones and their correlation with locomotor behavior in large-bodied hominoids. The comparisons between bending moments of metacarpals and metatarsals of the same ray furnished interesting results. Metacarpals III and especially IV of the knuckle-walking African apes were relatively stronger than those of humans and orangutans, and metatarsal V of humans was relatively stronger than those of the great apes. Interestingly, the relative robusticity of the metacarpal IV of the quadrumanous orangutan was between that of the African apes and that of humans. The main conclusions of the study are: 1) cross-sectional dimensions of metacarpals and metatarsals are influenced by locomotor modes in great apes and humans; 2) interlimb comparisons of cross-sectional properties of metacarpals and metatarsals are good indicators of locomotor modes in great apes and humans; and 3) the results of this study are in accord with those of previous analyses of plantar pressure and morphofunctional traits of the same bones, and with behavioral studies. These results provide a data base from which it will be possible to compare the morphology of the fossils in order to gain insight into the locomotor repertoires of extinct taxa.     

Kinematic model of tyrannosaurid (Dinosauria: Theropoda) arctometatarsus function

We present a hypothesis of tyrannosaurid foot function termed the "tensile keystone model," in which the triangular central metatarsal and elastic ligaments dynamically strengthened the foot. The tyrannosaurid arctometatarsus, in which the central metatarsal is proximally constricted, displays osteological correlates of distal intermetatarsal ligaments. The distal wedge-like imbrication of tyrannosaurid metatarsals indicates that rebounding ligaments drew the outer elements towards the middle digit early in the stance phase, unifying the arctometatarsus under high loadings. This suggests increased stability and resistance to dissociation and implies, but does not demonstrate, greater agility than in large theropods without an arctometatarsus.     

Midtarsal break variation in modern humans: Functional causes,skeletal correlates,and paleontological implications

The midtarsal break was once treated as a dichotomous, non-overlapping trait present in the foot of non-human primates and absent in humans. Recent work indicates that there is considerable variation in human midfoot dorsiflexion, with some overlap with the ape foot. These findings have called into question the uniqueness of the human lateral midfoot, and the use of osteological features in fossil hominins to characterize the midfoot of our extinct ancestors. Here, we present data on plantar pressure and pedal mechanics in a large sample of adults and children (n = 671) to test functional hypotheses concerning variation in midfoot flexibility. Lateral midfoot peak plantar pressure correlates with both sagittal plane flexion at the lateral tarsometatarsal joint, and dorsiflexion at the hallucal metatarsophalangeal joint. The latter finding suggests that midfoot laxity may compromise hallucal propulsion. Multiple regression statistics indicate that a low arch and pronation of the foot explain 40% of variation in midfoot peak plantar pressure, independent of age and BMI. MRI scans on a small subset of study participants (n = 19) reveals that curvature of the base of the 4th metatarsal correlates with lateral midfoot plantar pressure and that specific anatomies of foot bones do indeed reflect relative midfoot flexibility. However, while the shape of the base of the 4th metatarsal may reliably reflect midfoot mobility in individual hominins, given the wide range of overlapping variation in midfoot flexibility in both apes and humans, we caution against generalizing foot function in extinct hominin species until larger fossils samples are available. Am J Phys Anthropol 156:543–552, 2015. © 2015 Wiley Periodicals, Inc.