Diaphyseal Dimensions Research Articles

Variation in ontogenetic trajectories of limb dimensions in humans is attributable to both climatic effects and neutral evolution

Previous studies showed that there is variation in ontogenetic trajectories of human limb dimensions and proportions. However, little is known about the evolutionary significance of this variation. This study used a global sample of modern human immature long bone measurements and a multivariate linear mixed-effects model to study 1) whether the variation in ontogenetic trajectories of limb dimensions is consistent with ecogeographic predictions and 2) the effects of different evolutionary forces on the variation in ontogenetic trajectories. We found that genetic relatedness arising from neutral (nonselective) evolution, allometric variation associated with the change in size, and directional effects from climate all contributed to the variation in ontogenetic trajectories of all major long bone dimensions in modern humans. After accounting for the effects of neutral evolution and holding other effects considered in the current study constant, extreme temperatures have weak, positive associations with diaphyseal length and breadth measurements, while mean temperature shows negative associations with diaphyseal dimensions. The association with extreme temperatures fits the expectations of ecogeographic rules, while the association with mean temperature may explain the observed among-group variation in intralimb indices. The association with climate is present throughout ontogeny, suggesting an explanation of adaptation by natural selection as the most likely cause. On the other hand, genetic relatedness among groups, as structured by neutral evolutionary factors, is an important consideration when interpreting skeletal morphology, even for nonadult individuals.

Combining Variables to Improve Subadult Age Estimation

Anthropologists have reported that the combination of multiple variables and indicators generally increases precision and reduces bias in age estimates. However, endeavors specific to subadult age estimation have primarily focused on estimating age of the living and therefore on variables and indicators that are active later in ontogeny and easy to image. The current study aimed to determine if multivariable, single-indicator age-estimation models outperform single-variable age-estimation models throughout ontogeny using the three most common subadult age indicators: diaphyseal dimensions, epiphyseal fusion, and dental development. Data were collected from individuals from South Africa between birth and 12 years (N = 601) using Lodox Statscan radiographic images and from the United States between the ages of birth and 20 years (N = 1,277) using computed tomography images. Multivariate adaptive regression splines were used to build the multivariable, single-indicator, and single-variable models. Each subset used for model development had a unique training sample to build the model and testing sample to ensure that the results were generalizable. The multivariable models presented with increased precision and accuracy, reduced bias, and greater consistency across ontogeny compared to the single-variable models for both samples. Eighty percent of the independent test models (20/24) had ≥ 93% coverage, and 75% (18/24) of the independent tests models had ≥ 95% coverage. Besides providing more information to the resulting age estimate, multivariable models remove any a priori beliefs regarding variable importance and eliminate the requirement to contrive a final age estimate from multiple single-variable age-estimation models.

Testing the cross-applicability of juvenile sex estimation from diaphyseal dimensions

Sex estimation is a crucial component of the biological profile. Stull et al. (2017) have proposed a promising juvenile sex estimation method using long bone measurements taken from a South African sample, providing relatively high classification accuracies and made easy to use via the KidStats web-based app. In this study, we test the models developed by Stull et al. (2017) on an external historic population from Lisbon, Portugal, in order to determine whether the models can be reliably applied to archeological and forensic populations outside of the original population sample. The study sample consisted of 102 individuals (45 females and 57 males) aged under 13 years at death from the Lisbon identified skeletal collection. Measurements from these individuals were used to test the flexible discriminant analysis (FDA) models given by Stull et al. (2017). Allocation accuracies were calculated for boys and girls and children over and under 2 years separately and combined. Our findings show that the models developed by Stull et al. (2017) yield poor accuracy when applied to our external population and thus can potentially be misapplied on archeological skeletal remains or forensic remains of unknown origin. A number of statistical issues may explain why models fail to be transportable or even generalizable, namely multicollinearity, model overfitting and overly optimist bootstrapped cross-validation rates. It is also likely that population differences in size and sexual size dimorphism also affected the applicability of the models. We emphasize the importance of externally validating prediction models, particularly if they are intended to be applied across populations. Our study addresses Stull and co-worker’s request for further validation of the method on populations outside of South Africa, as the models cannot be confidently applied in the field until it has been externally validated.

The impact of terrain on lower limb bone structure.

Lower limb diaphyseal geometry is often used to evaluate mobility in past populations. Diaphyseal dimensions such as high shape (IX /IY ) indices generally thought to reflect high mobility may also result from walking over rough terrain. This study investigates the possible effects of terrain on lower limb diaphyseal cross-sectional geometric dimensions. The sample (N = 3,195) comprises adult skeletons from Europe, Africa, North America, and Asia, spanning from around 30,000 BP to mid-twentieth century. Femoral and tibial shape and bending/torsional strength dimensions were gathered either as part of a previous project or were generously provided by researchers. Local terrain for each site was quantified with ArcGIS mapping software using geographic coordinates and USGS elevation data, and characterized as flat, hilly, or mountainous. Analysis of variance shows significant differences (p < .05) in midshaft femoral and tibial shape ratio and relative bending/torsional strength among the three terrain categories, with more AP oriented diaphyseal shapes and greater relative strength in hilly and mountainous groups, even after correcting for the effect of subsistence. As expected, the impact of terrain is much more marked for hunter-gatherers and agriculturalists than for more mechanized recent populations. Interestingly, the effect of terrain is confounded in higher latitude individuals that exhibit increased ML bending strength, probably reflecting larger body breadth. This study underscores the mechanical significance of traveling over rough terrain and highlights the complex interactions of mobility, terrain, and body shape that contribute to shaping lower limb bone diaphyseal structure.

The Western (Buonfornello) necropolis (7th to 5th BC) of the Greek colony of Himera (Sicily, Italy): Site‐specific discriminant functions for sex determination in the common burials resulting from the battle of Himera (ca. 480 BC)

AbstractThe Western (Buonfornello) necropolis of Himera, dating from the 7th to 5th century BC, has yielded 10,096 burials, among which are six common burials (n = 2–22). On the basis of the archaeological and historical evidence, these are presumed to be the result of the Battle of Himera, fought in approximately 480 BC, or some other warfare episode taking place in the same period, and it is hypothesised that all the individuals are male. A reliable morphological sex determination is impossible in most cases due to the bad state of preservation of the pelves. The present paper aims to establish whether sex dimorphism in diaphyseal dimensions allows for the calculation of site‐specific discriminant functions (DFs) that are useful for sex determination and to verify the archaeological hypothesis that all individuals in the six common burials are male. DF computation was conducted for sex determination based on the diaphyseal dimensions recorded for a sample of 89 pelvis sexed individuals (57 males and 32 females). The DFs were selected by taking into account the large number of m/f samples (n &gt; 30) and the accuracy (&gt;90%).DF was tested on a sample of male/female Himeran pelvis sexed individuals not used for computing DFs (n = 17, 11 males and 6 females) and 27 skeletons from common burials. In the first case, pelvis and DF sex determination are concordant in 16 cases (94.1%). In the second case, three out of 27 individuals were discarded as DFs produced conflicting sex determinations. The remaining 24 individuals were sexed as males by means of DFs in 95.8% of cases. The archaeological hypothesis that all individuals in these common burials are male cannot be rejected as site‐specific DFs diagnose the male sex in a percentage that is higher than their accuracy in a sample of pelvis sexed individuals.

Lower limb articular scaling and body mass estimation in Pliocene and Pleistocene hominins

Previous attempts to estimate body mass in pre-Holocene hominins have relied on prediction equations derived from relatively limited extant samples. Here we derive new equations to predict body mass from femoral head breadth and proximal tibial plateau breadth based on a large and diverse sample of modern humans (avoiding the problems associated with using diaphyseal dimensions and/or cadaveric reference samples). In addition, an adjustment for the relatively small femoral heads of non-Homo taxa is developed based on observed differences in hip to knee joint scaling. Body mass is then estimated for 214 terminal Miocene through Pleistocene hominin specimens. Mean body masses for non-Homo taxa range between 39 and 49 kg (39–45 kg if sex-specific means are averaged), with no consistent temporal trend (6–1.85 Ma). Mean body mass increases in early Homo (2.04–1.77 Ma) to 55–59 kg, and then again dramatically in Homo erectus and later archaic middle Pleistocene Homo, to about 70 kg. The same average body mass is maintained in late Pleistocene archaic Homo and early anatomically modern humans through the early/middle Upper Paleolithic (0.024 Ma), only declining in the late Upper Paleolithic, with regional variation. Sexual dimorphism in body mass is greatest in Australopithecus afarensis (log[male/female] = 1.54), declines in Australopithecus africanus and Paranthropus robustus (log ratio 1.36), and then again in early Homo and middle and late Pleistocene archaic Homo (log ratio 1.20–1.27), although it remains somewhat elevated above that of living and middle/late Pleistocene anatomically modern humans (log ratio about 1.15).

Subadult sex estimation from diaphyseal dimensions.

Many studies on subadult sex estimation focus on elements that express sexually dimorphic features in adults. In contrast, diaphyseal dimensions have been shown to display sex-specific differences prior to adolescence. The current study evaluates the use of diaphyseal dimensions in subadult sex estimation. Eighteen postcranial measurements from six long bones were collected on Lodox Statscan radiographic images of 1,310 modern South African children between birth and 12 years of age. Linear (LDA) and flexible discriminant analysis (FDA) and logistic regression were employed with single and multiple variable models with age both included and excluded from the model. Bootstrapped cross-validation was employed because some of the multiple variable subsets had small sample sizes. Each of the bootstrapped accuracies has an associated 95% confidence interval demonstrating the ranges in classification. Classification methods utilizing multiple variables achieved the highest bootstrapped classification accuracies (70% to 93%). The inclusion of age in the models did not consistently increase or decrease the classification accuracies. Proximal and distal breadth measurements were consistently recognized as important measurements in model creation. FDA yielded the highest overall accuracies, but the logistic regression presented with overall smaller bootstrapped 95% confidence intervals. Quantifiable sex differences were discovered in the appendicular skeleton of children between birth and 12 years of age. The high classification accuracies were likely due to using numerous predictor variables from multiple skeletal elements, which were optimized for classification using FDA. To facilitate application, a graphical user interface, KidStats, was developed.

The Odocoileus virginianus Femur: Mechanical Behavior and Morphology.

Biomechanical research relies heavily on laboratory evaluation and testing with osseous animal structures. While many femora models are currently in use, including those of the European red deer (Cervus elaphus), the Odocoileus virginianus femur remains undocumented, despite its regional abundance in North America. The objective of this study was to compare biomechanical and morphological properties of the Odocoileus virginianus femur with those of the human and commonly used animal models. Sixteen pairs of fresh-frozen cervine femora (10 male, 6 female, aged 2.1 ± 0.9 years) were used for this study. Axial and torsional stiffnesses (whole bone) were calculated following compression and torsion to failure tests (at rates of 0.1 mm/sec and 0.2°/sec). Lengths, angles, femoral head diameter and position, periosteal and endosteal diaphyseal dimensions, and condylar dimensions were measured. The results show that the cervine femur is closer in length, axial and torsional stiffness, torsional strength, and overall morphology to the human femur than many other commonly used animal femora models; additional morphological measurements are comparable to many other species’ femora. The distal bicondylar width of 59.3mm suggests that cervine femora may be excellent models for use in total knee replacement simulations. Furthermore, the cervine femoral head is more ovoid than other commonly-used models for hip research, making it a more suitable model for studies of hip implants. Thus, with further, more application-specific investigations, the cervine femur could be a suitable model for biomechanical research, including the study of ballistic injuries and orthopaedic device development.

Using multivariate adaptive regression splines to estimate subadult age from diaphyseal dimensions.

Subadult age estimation is considered the most accurate parameter estimated in a subadult biological profile, even though the methods are deficient and the samples from which they are based are inappropriate. The current study addresses the problems that plague subadult age estimation and creates age estimation models from diaphyseal dimensions of modern children. The sample included 1,310 males and females between the ages of birth and 12 years. Eighteen diaphyseal length and breadth measurements were obtained from Lodox Statscan radiographic images generated at two institutions in Cape Town, South Africa, between 2007 and 2012. Univariate and multivariate age estimation models were created using multivariate adaptive regression splines. k-fold cross-validated 95% prediction intervals (PIs) were created for each model, and the precision of each model was assessed. The diaphyseal length models generated the narrowest PIs (2 months to 6 years) for all univariate models. The majority of multivariate models had PIs that ranged from 3 months to 5 and 6 years. Mean bias approximated 0 for each model, but most models lost precision after 10 years of age. Univariate diaphyseal length models are recommended for younger children, whereas multivariate models are recommended for older children where the inclusion of more variables minimized the size of the PIs. If diaphyseal lengths are not available, multivariate breadth models are recommended. The present study provides applicable age estimation formulae and explores the advantages and disadvantages of different subadult age estimation models using diaphyseal dimensions. Am J Phys Anthropol 154:376-386, 2014. © 2014 Wiley Periodicals, Inc.

Functional significance of genetic variation underlying limb bone diaphyseal structure

Limb bone diaphyseal structure is frequently used to infer hominin activity levels from skeletal remains, an approach based on the well-documented ability of bone to adjust to its loading environment during life. However, diaphyseal structure is also determined in part by genetic factors. This study investigates the possibility that genetic variation underlying diaphyseal structure is influenced by the activity levels of ancestral populations and might also have functional significance in an evolutionary context. We adopted an experimental evolution approach and tested for differences in femoral diaphyseal structure in 1-week-old mice from a line that had been artificially selected (45 generations) for high voluntary wheel running and non-selected controls. As adults, selected mice are significantly more active on wheels and in home cages, and have thicker diaphyses. Structural differences at 1 week can be assumed to primarily reflect the effects of selective breeding rather than direct mechanical stimuli, given that the onset of locomotion in mice is shortly after Day 7. We hypothesized that if genetically determined diaphyseal structure reflects the activity patterns of members of a lineage, then selected animals will have relatively larger diaphyseal dimensions at 1 week compared to controls. The results provide strong support for this hypothesis and suggest that limb bone cross sections may not always only reflect the activity levels of particular fossil individuals, but also convey an evolutionary signal providing information about hominin activity in the past.

Middle Paleolithic Human Remains from Bisitun Cave, Iran

Résumé: En 1949, C. S. Coon a identifié comme humains deux restes fossiles provenant des niveaux moustériens de la grotte de Bisitun, près de Kermanshah, en Iran. Le premier, une incisive d’un bovidé, doit donc être écarté de l’échantillon des fossiles humains. Le deuxième est une diaphyse proximale de radius droit humain. La comparaison des dimensions de cette diaphyse à celles des radius droits des Néandertaliens et des premiers hommes modernes du Paléolithique moyen la rend proche des Néandertaliens (et des hommes modernes du Paléolithique supérieur), mais elle la sépare des hommes fossiles connus du Paléolithique moyen de l’Asie du Sud-Ouest.

The case is unchanged and remains robust: Australopithecus afarensis exhibits only moderate skeletal dimorphism. A reply to

Previous attempts to estimate body mass in pre-Holocene hominins have relied on prediction equations derived from relatively limited extant samples. Here we derive new equations to predict body mass from femoral head breadth and proximal tibial plateau breadth based on a large and diverse sample of modern humans (avoiding the problems associated with using diaphyseal dimensions and/or cadaveric reference samples). In addition, an adjustment for the relatively small femoral heads of non-Homo taxa is developed based on observed differences in hip to knee joint scaling. Body mass is then estimated for 214 terminal Miocene through Pleistocene hominin specimens. Mean body masses for non-Homo taxa range between 39 and 49 kg (39–45 kg if sex-specific means are averaged), with no consistent temporal trend (6–1.85 Ma). Mean body mass increases in early Homo (2.04–1.77 Ma) to 55–59 kg, and then again dramatically in Homo erectus and later archaic middle Pleistocene Homo, to about 70 kg. The same average body mass is maintained in late Pleistocene archaic Homo and early anatomically modern humans through the early/middle Upper Paleolithic (0.024 Ma), only declining in the late Upper Paleolithic, with regional variation. Sexual dimorphism in body mass is greatest in Australopithecus afarensis (log[male/female] = 1.54), declines in Australopithecus africanus and Paranthropus robustus (log ratio 1.36), and then again in early Homo and middle and late Pleistocene archaic Homo (log ratio 1.20–1.27), although it remains somewhat elevated above that of living and middle/late Pleistocene anatomically modern humans (log ratio about 1.15).

Cross-population analysis of the growth of long bones and the os coxae of three Early Medieval Austrian populations

Inter-population variability in long-bone and pelvic-bone growth during the Early Medieval period is examined. The materials comprise four archaeological populations: two Slavonic (Gars-Thunau, Zwentendorf, Austria, 10th-century AD), one Avar (Zwölfaxing, Austria, 8th-century AD), and one Anglo-Saxon (Raunds, England, 10th-century AD). Bone measurements are analyzed against dental age estimates in order to assess inter-population differences in growth rates for long-bone and os coxae bone dimensions. Growth curves of the upper and lower extremities of additional archaeological populations and a modern North-American population are also assessed. The expectation was that the greatest differences in growth patterns would be found between the Anglo-Saxon and the Austrian samples, due to their distinct genetic and biocultural background. Minimal differences were expected between the two Slavonic populations, as these were approximately contemporaneous, recovered from geographically close locations, and shared relatively similar archaeological contexts. Growth curves were estimated for each bone dimension by fitting least-squares fourth-order polynomials (which allowed testing of population differences by analysis of covariance), and iteratively estimating Gompertz growth curves. The results showed differences between bones in the extent of inter-population variability, with diaphyseal long-bone growth showing equivalent patterns across the four populations, but significant differences between populations in the growth patterns of distal diaphyseal dimensions of the femur and humerus and the dimensions of the ilium. Varying growth patterns are therefore associated with inter-population differences in absolute dimensions in relation to age as well as variations in growth velocities. Inter-population variability in growth curves in the case of femoral and humeral dimensions were most pronounced during infancy (0-2 years). The most consistent differences in bone growth and related dimensions are between Zwölfaxing and the other samples. No significant differences in growth were detected between the Anglo-Saxon and the Austrian populations.

Gough's Cave 1 (Somerset, England): a study of the pelvis and lower limbs

The lower limb remains of Gough's Cave 1 retain most of the pelvis, both femora, one complete tibia and portions of the other, sections of both fibulae, two tarsals and three metatarsals. They are those of a largely average European Mesolithic young adult male. Overall diaphyseal robusticity is generally similar to that of other Mesolithic specimens, even though the fibula and third metatarsal appear gracile. Musculo-ligamentous attachment areas are generally weakly marked. The proximal femora and the femoral diaphyses exhibit a clear asymmetry, especially in their neck-shaft angles and diaphyseal dimensions, which is is accompanied in the pelvis by a greater degree of left iliac lateral flare. These aspects are associated with a pelvis that combines several distinctly male characteristics with an overall pelvic aperture shape which is female.

Dual-energy X-ray absorptiometry derived structural geometry for stress fracture prediction in male U.S. Marine Corps recruits.

A total of 626 U.S. male Marine Corps recruits underwent anthropometric measurements and dual-energy X-ray absorptiometry (DXA) scans of the femoral midshaft and the distal third of the tibia prior to a 12 week physical training program. Conventionally obtained frontal plane DXA scan data were used to measure the bone mineral density (BMD) as well as to derive the cross-sectional area, moment of inertia, section modulus, and bone width in the femur, tibia, and fibula. During training, 23 recruits (3.7%) presented with a total of 27 radiologically confirmed stress fractures in various locations in the lower extremity. After excluding 16 cases of shin splints, periostitis, and other stress reactions that did not meet fracture definition criteria, we compared anthropometric and bone structural geometry measurements between fracture cases and the remaining 587 normals. There was no significant difference in age (p = 0.8), femur length (p = 0.2), pelvic width (p = 0.08), and knee width at the femoral condyles (p = 0.06), but fracture cases were shorter (p = 0.01), lighter (p = 0.0006), and smaller in most anthropometric girth dimensions (p < 0.04). Fracture case bone cross-sectional areas (p < 0.001), moments of inertia (p < 0.001), section moduli (p < 0.001), and widths (p < 0.001) as well as BMD (p < 0.03) were significantly smaller in the tibia and femur. After correcting for body weight differences, the tibia cross-sectional area (p = 0.03), section modulus (p = 0.05), and width (p = 0.03) remained significantly smaller in fracture subjects. We conclude that both small body weight and small diaphyseal dimensions relative to body weight are factors predisposing to the development of stress fractures in this population. These results suggest that bone structural geometry measurements derived from DXA data may provide a simple noninvasive methodology for assessing the risk of stress fracture.

The Effect of Aging on the Shape of the Proximal Femur

The design of cementless femoral prostheses is based on the assumption that age and gender do not affect the shape of the proximal femur. To test this hypothesis, standard anteroposterior and lateral radiographs were prepared of 4 sets of 20 femora, obtained from young (range, 40-60 years) and elderly (range, 60-90 years) donors of both genders. The intracortical and extracortical borders of each femur were digitized electronically, and key parameters were measured to define the shape and dimensions of the medullary canal and the position of the femoral head. Systematic differences were observed between the size and shape of male and female femora. Extracortical dimensions were larger in the male femora by 14% to 19%, and endosteal dimensions by 11% to 24%. However, there were no significant differences between the canal shape of young male and young female femora in the coronal, sagittal, or transverse planes. The male femora displayed no significant differences in canal shape or endosteal width as a function of age. Profound differences were observed in the endosteal shape and diaphyseal dimensions of the young and old female femora. The older female femora had wider canals at the level of the isthmus, with a significant reduction in the canal flare index (the ratio between the canal width proximal to the lesser trochanter and at the isthmus). This study demonstrates that cementless femoral prostheses of 1 standard shape cannot provide a close fit to the endosteal contours of young and elderly women.

Postcranial robusticity in Homo. III: Ontogeny.

The influence of developmental factors on long-bone cross-sectional geometry and articular size in modern humans is investigated using two approaches: (1) an analysis of the effects of increased mechanical loading on long-bone structure when applied during different developmental periods, using data collected for a study of upper limb bone bilateral asymmetry in professional tennis players; and (2) an analysis of the relative timing of age changes in femoral dimensions among juveniles from the Pecos Pueblo Amerindian archaeological sample. Results of these analyses are used to interpret the femoral morphology of three pre-Recent Homo juveniles--the H. erectus KNM-WT 15000 and the archaic H. sapiens La Ferrassie 6 and Teshik-Tash 1--as well as observed differences in postcranial morphology between adult Recent and earlier Homo (Ruff et al., 1993). Our findings indicate the following: (1) There are age-related changes in long-bone diaphyseal envelope sensitivity to increased mechanical loading, with the periosteal envelope more responsive prior to mid-adolescence, and the endosteal envelope more responsive thereafter. The periosteal expansion and endosteal contraction of the diaphysis documented earlier for adult pre-Recent Homo relative to Recent humans (Ruff et al., 1993) is thus consistent with a developmental response to increased mechanical loading applied throughout life. The relatively large medullary cavity in the 11-12-year-old KNM-WT 15000 femur is also consistent with this model. However, the two archaic H. sapiens juveniles show relatively small medullary cavities, possibly indicating a modified developmental pattern in this group. (2) Articulations follow a growth pattern similar to that of long-bone length (and stature), while cross-sectional diaphyseal dimensions (cortical area, second moments of area) show a contrasting growth pattern, with slower initial growth from childhood through mid-adolescence, followed by a "catch-up" period that continues through early adulthood. This latter pattern is more similar to the growth curve for body weight, and may in fact partially reflect adaptation of the diaphysis to increased weight bearing. Because of these different growth patterns, articulations appear relatively large, and diaphyseal breadths relatively small during late childhood to mid-adolescence (i.e., about 9-13 years), when compared to adults from the same population. KNM-WT 15000 shows this same proportional difference from adult early Homo specimens, which is therefore interpreted as simply a developmental consequence of his age at death.(ABSTRACT TRUNCATED AT 400 WORDS)