Adaptations to humeral torsion in medieval Britain

Adaptations to the humeral torsion angle have been identified in the professional throwing athlete. This movement pattern increases the humeral torsion angle, and also increases the extent of external rotation movements in the dominant, throwing limb when compared with the nondominant limb. The purpose of this paper is to test the hypothesis that the humeral torsion angle is an adaptation to upper limb use. This project examines the humeral torsion angle in a number of medieval British populations, as well as a modern cadaver-based sample. The results identify significant differences in the humeral torsion angle both between and within male (P < 0.001, ANOVA) and female (P < 0.014, ANOVA) populations, although the results are not consistent with expected behavior patterns. Statistically significant differences between males and females within the same site were identified in 2 of the 5 samples examined. The mean level of bilateral asymmetry does not approach that reported for the professional throwing athlete. However, a number of individuals have high levels of asymmetry in excess of that identified in the professional throwing athlete. This analysis demonstrates the need for individual rather than population-based analyses, as the heterogeneity within population samples obscures individual variation in activity patterns. The diversity within British medieval society and a lack of specific known behaviors prevent further identification of the functional significance of the humeral torsion angle within the archaeological record examined here.     

Patterns of humeral asymmetry among Late Pleistocene humans

Human humeral diaphyseal asymmetry in midshaft and mid-distal rigidity is assessed through the Late Pleistocene in samples of late archaic (Neandertal) and early modern humans. It is considered with respect to directionality (handedness), levels of asymmetry, body size and sexual differences. The overall Late Pleistocene sample indicates a right-handed preference in frequencies (right: 74.8%, left: 15.0%, ambiguous: 10.3%), which are similar to those of recent human samples. Average levels of humeral asymmetry are elevated relative to Holocene samples through all but the small Middle Paleolithic modern human and eastern Eurasian late Upper Paleolithic samples. Humeral asymmetry is especially high among the males relative to the females, and the possibility of a division of labor between uni-manual tasks (mostly male) and bi-manual tasks (mostly female) is considered. At the same time, there is a general pattern of increased asymmetry with larger body size, but it remains unclear to what extent it reflects body size versus sexual effects on bilateral humeral loading. There do not appear to have been substantial changes in humeral asymmetry through time, indicating a continuity of similar manual behavioral patterns through the Late Pleistocene, despite considerable changes in technology through the Late Pleistocene.     

The ontogeny of Holocene and Late Pleistocene human postcranial strength

While a wide variety of studies have focused on population variation in adult cross‐sectional properties, relatively little is known about population variation in postcranial robusticity in immature individuals. Furthermore, the age at which the population differences readily detected in adults manifest during growth is also unknown. This research addresses these gaps in our current understanding through the analysis of immature humeral and femoral long bone strength. Cross‐sectional geometry was used to compare the developmental trajectories of diaphyseal strength in Late Pleistocene Neandertal and modern human subadults to a sample of immature humans from seven geographically diverse Holocene populations. Population differences in size‐standardized cross‐sectional properties appear to be systemic and develop very early in ontogeny in the Holocene sample. In many cases, these differences are present before one year of age. In general, the Late Pleistocene fossil samples fit within the range of recent human variation in long bone strength. Population differences detected here are likely related to a combination of factors including activity patterns, genetic propensities, and nutritional status. These results highlight the complex mosaic of processes that result in adult postcranial robusticity, and suggest that further exploration of the developmental interplay between intrinsic and extrinsic influences on skeletal robusticity will likely enhance our understanding of adult postcranial morphology. Am J Phys Anthropol 2010. © 2009 Wiley‐Liss, Inc.     

Sex differences in humeral bilateral asymmetry in two hunter‐gatherer populations: California Amerinds and British Columbian Amerinds

This study uses two prehistoric Amerindian populations of hunter‐gatherer subsistence patterns to determine whether levels of sexual dimorphism in humeral bilateral cross‐sectional asymmetry are related to sex‐specific differences in activities among these populations. Results confirmed that males of the California Amerind population who engaged in the more unimanual activities of spear hunting and warfare were more asymmetrical than were their female counterparts who engaged in the more bimanual activities of grinding acorns. California Amerind males were also more asymmetrical than British Columbian Amerind males who rowed (using both arms) extensively. Sex differences within British Columbian Amerinds were not statistically significant, nor were female differences between populations. In general, levels of humeral asymmetry appear to be more dependent on sex and population‐specific behaviors rather than broad subsistence patterns. Am J Phys Anthropol 2009. © 2009 Wiley‐Liss, Inc.     

Postcranial adaptations for climbing in Loridae (Primates)

The Loridae are an arboreal family of small primates that are specialized for slow and quiet climbing. This paper examines the relationship between lorid locomotory behaviour and postcranial skeletal morphology. Lorid humeral and femoral diaphyseal geometric cross-sectional properties, articular surface areas, and lengths are compared to those properties in other small primates with less specialized locomotory behaviour. The comparative sample includes both closely related prosimians and more distantly related platyrrhines.
Results indicate that lorids have greater humeral and femoral diaphyseal rigidity than other quadrupedal primates of similar body size, suggesting that lorid limbs are subjected to greater forces. Lorids also have relatively larger humeral and femoral articulations, corresponding to field and laboratory observations which indicate that lorid joints are highly mobilc. In addition, lorids have long humeri relative to femoral length, and compared to humeral length in less specialized prosimians of similar body mass. Long humeral length relative to femoral length is interpreted as a climbing adaptation because similar limb proportions are also seen in many non-primate climbers. Altogether, humeral and femoral diaphyseal cross-sectional properties, articular surface areas, and lengths comprise a suite of characters which have potential for identifying climbing specialists in the fossil record.     

Postcranial robusticity in Homo. II: Humeral bilateral asymmetry and bone plasticity

The analysis of humeral asymmetry in Recent human skeletal samples and an extant tennis-player sample documents minimal asymmetry in bone length, little asymmetry in distal humeral articular breadth, but pronounced and variable asymmetry in mid- and distal diaphyseal crosssectional geometric parameters. More specifically, skeletal samples of normal modern Euroamericans, prehistoric and early historic Amerindians, and prehistoric Japanese show moderate (ca. 5–14%) median asymmetry in diaphyseal cross-sectional areas and polar second moments of area, whereas the tennis-player sample, with pronounced unilateral physical activity, exhibits median asymmetries of 28–57% in the same parameters. A sample of Neandertals with nonpathological upper limbs exhibits similarly low articular asymmetry but pronounced diaphyseal asymmetries, averaging 24–57%. In addition, three Neandertals with actual or possible post-traumatic upper limb alterations have the same low articular asymmetry but extremely high diaphyseal asymmetries, averaging 112–215%. These data support those from experimental work on animals, exercise programs of humans, and human clinical contexts in establishing the high degree of diaphyseal plasticity possible for humans, past and present, under changing biomechanical loading conditions. This lends support to activity-related functional interpretations of changing human diaphyseal morphology and robusticity during the Pleistocene. © 1994 Wiley-Liss, Inc.     

Limb bone bilateral asymmetry: variability and commonality among modern humans

Humans demonstrate species-wide bilateral asymmetry in long bone dimensions. Previous studies have documented greater right-biases in upper limb bone dimensions--especially in length and diaphyseal breadth--as well as more asymmetry in the upper limb when compared with the lower limb. Some studies have reported left-bias in lower limb bone dimensions, which, combined with the contralateral asymmetry in upper limbs, has been termed "crossed symmetry." The examination of sexual dimorphism and population variation in asymmetry has been limited. This study re-examines these topics in a large, geographically and temporally diverse sample of 780 Holocene adult humans. Fourteen bilateral measures were taken, including maximum lengths, articular and peri-articular breadths, and diaphyseal breadths of the femur, tibia, humerus, and radius. Dimensions were converted into percentage directional (%DA) and absolute (%AA) asymmetries. Results reveal that average diaphyseal breadths in both the upper and lower limbs have the greatest absolute and directional asymmetry among all populations, with lower asymmetry evident in maximum lengths or articular dimensions. Upper limb bones demonstrate a systematic right-bias in all dimensions, while lower limb elements have biases closer to zero %DA, but with slight left-bias in diaphyseal breadths and femoral length. Crossed symmetry exists within individuals between similar dimensions of the upper and lower limbs. Females have more asymmetric and right-biased upper limb maximum lengths, while males have greater humeral diaphyseal and head breadth %DAs. The lower limb demonstrates little sexual dimorphism in asymmetry. Industrial groups exhibit relatively less asymmetry than pre-industrial humans and less dimorphism in asymmetry. A mixture of influences from both genetic and behavioral factors is implicated as the source of these patterns.     

Diet and diversity at later medieval Fishergate: the isotopic evidence

We present the results of stable carbon and nitrogen isotope analysis of bone collagen for 155 individuals buried at the Later Medieval (13th to early 16th century AD) Gilbertine priory of St. Andrew, Fishergate in the city of York (UK). The data show significant variation in the consumption of marine foods between males and females as well as between individuals buried in different areas of the priory. Specifically, individuals from the crossing of the church and the cloister garth had consumed significantly less marine protein than those from other locations. Isotope data for four individuals diagnosed with diffuse idiopathic skeletal hyperostosis (DISH) are consistent with a diet rich in animal protein. We also observe that isotopic signals of individuals with perimortem sharp force trauma are unusual in the context of the Fishergate dataset. We discuss possible explanations for these patterns and suggest that there may have been a specialist hospital or a local tradition of burying victims of violent conflict at the priory. The results demonstrate how the integration of archaeological, osteological, and isotopic data can provide novel information about Medieval burial and society.     

Femoral/humeral strength in early African Homo erectus

Lower-to-upper limb-bone proportions give valuable clues to locomotor behavior in fossil taxa. However, to date only external linear dimensions have been included in such analyses of early hominins. In this study, cross-sectional measures of femoral and humeral diaphyseal strength are determined for the two most complete early Homo erectus (or ergaster) associated skeletons--the juvenile KNM-WT 15000 and the adult KNM-ER 1808. Modern comparative samples include an adult human skeletal sample representative of diverse body shapes, a human longitudinal growth series, and an adult chimpanzee sample. When compared to appropriately age-matched samples, both H. erectus specimens fall very close to modern human mean proportions and far from chimpanzee proportions (which do not overlap with those of humans). This implies very similar mechanical load-sharing between the lower and upper limbs, and by implication, similar locomotor behavior in early H. erectus and modern humans. Thus, by the earliest Pleistocene (1.7 Ma), completely modern patterns of bipedal behavior were fully established in at least one early hominin taxon.     

Does skeletal anatomy reflect adaptation to locomotor patterns? cortical and trabecular architecture in human and nonhuman anthropoids

Although the correspondence between habitual activity and diaphyseal cortical bone morphology has been demonstrated for the fore- and hind-limb long bones of primates, the relationship between trabecular bone architecture and locomotor behavior is less certain. If sub-articular trabecular and diaphyseal cortical bone morphology reflects locomotor patterns, this correspondence would be a valuable tool with which to interpret morphological variation in the skeletal and fossil record. To assess this relationship, high-resolution computed tomography images from both the humeral and femoral head and midshaft of 112 individuals from eight anthropoid genera (Alouatta, Homo, Macaca, Pan, Papio, Pongo, Trachypithecus, and Symphalangus) were analyzed. Within-bone (sub-articular trabeculae vs. mid-diaphysis), between-bone (forelimb vs. hind limb), and among-taxa relative distributions (femoral:humeral) were compared. Three conclusions are evident: (1) Correlations exists between humeral head sub-articular trabecular bone architecture and mid-humerus diaphyseal bone properties; this was not the case in the femur. (2) In contrast to comparisons of inter-limb diaphyseal bone robusticity, among all species femoral head trabecular bone architecture is significantly more substantial (i.e., higher values for mechanically relevant trabecular bone architectural features) than humeral head trabecular bone architecture. (3) Interspecific comparisons of femoral morphology relative to humeral morphology reveal an osteological "locomotor signal" indicative of differential use of the forelimb and hind limb within mid-diaphysis cortical bone geometry, but not within sub-articular trabecular bone architecture.     

Role of Nonbehavioral Factors in Adjusting Long Bone Diaphyseal Structure in Free-ranging <Emphasis Type="Italic">Pan troglodytes</Emphasis>

Limb bones deform during locomotion and can resist the deformations by adjusting their shapes. For example, a tubular-shaped diaphysis best resists variably-oriented deformations. As behavioral profiles change during adulthood, patterns of bone deformation may exhibit age trends. Habitat characteristics, e.g., annual rainfall, tree density, and elevation changes, may influence bone deformations by eliciting individual components of behavioral repertoires and suppressing others, or by influencing movements during particular components. Habituated chimpanzee communities provide a unique opportunity to examine these factors because of the availability of morphological data and behavioral observations from known-age individuals inhabiting natural habitats. We evaluated adult femora and humeri of 18 female and 10 male free-ranging chimpanzees (Pan troglodytes) from communities in Gombe (Tanzania), Mahale Mountains (Tanzania), and Taï Forest (Côte d’Ivoire) National Parks. We compare cross sections at several locations (35%, 50%, 65% diaphyseal lengths). Community comparisons highlight different diaphyseal shapes of Taï females relative to Mahale and Gombe females, particularly in humeral diaphyses. Age trends in diaphyseal shapes are consistent with reduced activity levels in general, not only reduced arboreal activity. Age-related bone loss is apparent among community females, but is less striking among males. Community trends in diaphyseal shape are qualitatively consistent with ranked annual rainfall at localities, tree density, and elevation change or ruggedness of terrain. Habitat characteristics may contribute to variation in diaphyseal shape among chimpanzee communities, much like among modern human groups, but verification awaits further rigorous experimental and comparative analyses.     

内蒙古和林格尔土城子农业人群与林西井沟子游牧人群股骨中部的生物力学对比

肢骨的形态结构可以反映人类进化、古代人群的生存适应性活动和生存环境等重要信息。基于“骨骼功能适应”和“杠杆原理”,有学者对不同生计方式的古代人群下肢股骨开展了大量的研究工作,但是,国内外尚未有关于农业人群和游牧人群股骨之间差异性研究的报道。本文选取两个具有代表性的古代人群,即内蒙古和林格尔土城子戍边农业人群和内蒙古林西井沟子游牧人群进行对比研究。通过对股骨骨干中部横断面生物力学分析发现,农业人群股骨粗壮度与游牧人群之间具有显著差异。前者的平均粗壮度较大,后者女性组下肢骨的活动强度明显较小,这可能与游牧人群经常从事骑马活动而下肢骨活动强度相对较少有关。农业人群股骨指数的变异范围均大于游牧人群,这可能与前者男性的士兵身份有关;同时,也提示土城子男性组股骨所反映的行为活动信息并不代表真正意义上的纯农业人群下肢骨行为模式,而是一种农业和士兵行为的混合模式。在性别分工上,井沟子组的男女性均从事骑马活动,两侧股骨受力较为一致,在两侧不对称性程度和骨干横断面形状上的男女差异不大;男性股骨的粗壮度要明显大于女性,这与井沟子组男性还从事一定的狩猎行为有关。与游牧人群女性较为纤细的股骨不同,土城子组女性作为典型的农业人群代表,其下肢骨整体的活动强度较大,几乎与同组的男性和井沟子组男性相当,组内的性别差异相对较小;骨干横断面形状的显著性差异说明,土城子组内部男性和女性的行为活动方式存在明显的性别分工。本文研究结果说明农业人群女性的下肢骨活动强度较大,在行为活动方式上,戍边农业人群具有更为明显的性别分工。     

Stable isotope evidence for 1500 years of human diet at the city of York, UK

We present here the results of a large-scale diachronic palaeodietary (carbon and nitrogen isotopic measurements of bone collagen) study of humans and animals from a single site, the city of York (U.K.), dating from the Roman period to the early 19th century. The human sample comprises 313 burials from the cemeteries of Trentholme Drive and Blossom Street (Roman), Belle Vue House (Anglo-Saxon), Fishergate (High and Later Medieval), and All Saints, Pavement (Later and Post-Medieval). In addition, 145 samples of mammal, fish and bird bone from the sites of Tanner Row and Fishergate were analyzed. The isotope data suggest dietary variation between all archaeological periods, although the most significant change was the introduction of significant quantities of marine foods in the Medieval periods. These are first evident in the diet of a small group of individuals from the High Medieval cemetery at Fishergate, although they were consumed almost universally in the following periods. The human isotope values are also remarkable due to unusually elevated delta(15)N ratios that are not sufficiently explained by the comparably small enrichment in (13)C that accompanies them. We discuss the possible reasons behind this and the archaeological significance of the data set.     

Activity patterns in the Sahara Desert: an interpretation based on cross-sectional geometric properties

The Garamantian civilization flourished in modern Fezzan, Libya, between 900 BC and 500 AD, during which the aridification of the Sahara was well established. Study of the archaeological remains suggests a population successful at coping with a harsh environment of high and fluctuating temperatures and reduced water and food resources. This study explores the activity patterns of the Garamantes by means of cross-sectional geometric properties. Long bone diaphyseal shape and rigidity are compared between the Garamantes and populations from Egypt and Sudan, namely from the sites of Kerma, el-Badari, and Jebel Moya, to determine whether the Garamantian daily activities were more strenuous than those of other North African populations. Moreover, sexual dimorphism and bilateral asymmetry are assessed at an intra- and inter-population level. The inter-population comparisons showed the Garamantes not to be more robust than the comparative populations, suggesting that the daily Garamantian activities necessary for survival in the Sahara Desert did not generally impose greater loads than those of other North African populations. Sexual dimorphism and bilateral asymmetry in almost all geometric properties of the long limbs were comparatively low among the Garamantes. Only the lower limbs were significantly stronger among males than females, possibly due to higher levels of mobility associated with herding. The lack of systematic bilateral asymmetry in cross-sectional geometric properties may relate to the involvement of the population in bilaterally intensive activities or the lack of regular repetition of unilateral activities.     

Humeral torsion revisited: a functional and ontogenetic model for populational variation

Anthropological interest in humeral torsion has a long history, and several functional explanations for observed variation in the orientation of the humeral head have been proposed. Recent clinical studies have revived this topic by linking patterns of humeral torsion to habitual activities such as overhand throwing. However, the precise functional implications and ontogenetic history of humeral torsion remain unclear. This study examines the ontogeny of humeral torsion in a large sample of primarily immature remains from six different skeletal collections (n = 407). The results of this research confirm that humeral torsion displays consistent developmental variation within all populations of growing children; neonates display relatively posteriorly oriented humeral heads, and the level of torsion declines steadily into adulthood. As in adults, variation in the angle of humeral torsion in immature individuals varies by population, and these differences arise early in development. However, when examined in the context of the developing muscles of the shoulder complex, it becomes apparent that variation in the angle of humeral torsion is not necessarily related to specific habitual activities. Variability in this feature is more likely caused by a generalized functional imbalance between muscles of medial and lateral rotation that can be produced by a wide variety of upper limb activity patterns during growth.     

The paleobiology of two adult skeletons from Baousso da Torre (Bausu da Ture) (Liguria,Italy): Implications for Gravettian lifestyle

This article presents the main skeletal characteristics of the two largely unpublished Gravettian adult skeletons from Baousso da Torre (Liguria, Italy). BT1 and BT2 were two adult tall males, who died aged respectively between 20 and 50 years, and between 20 and 30 years. Their body proportions fall within the range of variation known for the middle Upper Paleolithic, and their skeletal remains are characterized by considerable osteometric values, which fall consistently in the upper part of the Upper Paleolithic male sample variation. They show a high degree of upper limb lateralization, implying that they were likely involved in strenuous and/or repetitive unimanual tasks. They also exhibit very robust lower limbs, likely related to repeated long-distance travels in mountainous terrains. These results are in total agreement with previous hypotheses on Late Pleistocene population behavioral patterns.     

Sexual dimorphism in human lower limb bone structure: relationship to subsistence strategy and sexual division of labor

Cross-sectional geometric properties of the human femur and tibia are compared in males and females in a number of recent and archaeological population samples extending back to the Middle Paleolithic. There is a consistent decline in sexual dimorphism from hunting-gathering to agricultural to industrial subsistence strategy levels in properties which measure relative anteroposterior bending strength of the femur and tibia in the region about the knee. This trend parallels and is indicative of reductions in the sexual division of labor, in particular differences in the relative mobility of males and females. Sexual dimorphism in mediolateral bending strength near the hip shows no consistent temporal trend, probably reflecting relatively constant sex differences in pelvic structure related to the requirements of childbirth. Upper and Middle Paleolithic samples are indistinguishable in terms of sexual dimorphism from modern huntergatherers, suggesting a similar sexual division of labor. The results illustrate the utility of cross-sectional geometric parameters of long bone diaphyses in reconstructing behavioral differences within and between past populations. Some variations in the accuracy of sexing techniques based on diaphyseal measurements of the lower limb long bones may also be explained by these behavioral and structural factors.     

Ontogenetic adaptation to bipedalism: age changes in femoral to humeral length and strength proportions in humans, with a comparison to baboons

The increase in lower/upper limb bone length and strength proportions in adult humans compared to most other anthropoid primates is commonly viewed as an adaptation to bipedalism. The ontogenetic development of femoral to humeral proportions is examined here using a longitudinal sample of 20 individuals measured radiographically at semiannual or annual intervals from 6 months of age to late adolescence (a subset of the Denver Growth Study sample). Anthropometric data (body weights, muscle breadths) were also available at each examination age. Results show that while femoral/humeral length proportions close to those of adults are already present in human infants, characteristically human femoral/humeral diaphyseal strength proportions only develop after the adoption of bipedalism at about 1 year of age. A rapid increase in femoral/humeral strength occurs between 1 and 3 years, followed by a slow increase until mid-late adolescence, when adult proportions are reached. When age changes in material properties are factored in, femoral strength shows an almost constant relationship to body size (body mass.bone length) after 5 years of age, while humeral strength shows a progressive decline relative to body size. Femoral/humeral length proportions increase slightly throughout growth, with no apparent change in growth trajectory at the initiation of walking, and with a small decline in late adolescence due to later growth in length of the humerus. A sex difference in femoral/humeral strength proportions (females greater) but not length proportions, develops early in childhood. Thus, growth trajectories in length and strength proportions are largely independent, with strength proportions more responsive to actual changes in mechanical loading. A cross-sectional ontogenetic sample of baboons (n=30) illustrates contrasting patterns of growth, with much smaller age changes in proportions, particularly strength proportions, although there is some indication of an adaptation to altered limb loadings early in baboon development.     

Perforation of the coronoid-olecranon septum. Humero-ulnar relationships in Netherlands and African populations

The mechanical interaction of the humerus and ulna during articulation has been examined in skeletal material from African and European populations. It was found that the presence or absence of perforation of the humeral septum and the thickness of the septum when unperforated can be predicted with considerable accuracy when two factors are known: the length of the coronoid and olecranon processes of the ulna relative to the minimum diameter of the trochlea humeri, and the total angle of movement at the elbow from full extension to full flexion. The influences which control either or both of these factors also determine the thickness of the septum of the humerus. These influences are not well known and there is opportunity for both genetic and environmental effects. The coefficients of correlation between septum thickness, flexion angle, humeral robusticity, minimum circumference of the humerus, trochlea diameter and brachial index are given. Septum thickness is correlated more highly with trochlea diameter than with robusticity or minimum circumference of the humerus. The higher frequency of perforation of the septum in the African sample (47%) compared with the European (6%) is associated with a greater range of movement at the elbow in the Africans. Bilateral, sexual and population differences in the frequency of perforation of the humeral septum may be associated with differences in the extent of movement at the elbow joint. Variation in this feature, in turn, may be associated with muscularity and somatotype.     

Muscle markers revisited: activity pattern reconstruction with controls in a central California Amerind population

Anthropologists frequently use musculoskeletal stress markers to reconstruct past activity patterns. Yet, researchers have called into question the reliability of muscle marker measurements in part because body size and age affect muscle marker scores. In this study, the author examined an aggregate upper limb muscle marker to determine if after controlling for the effects of body size and age, one could reconstruct activity patterns of a prehistoric Amerind population. Analyses were made of a sample of 102 (43 males, 59 females) prehistoric central California Amerinds. Muscle markers were measured using two-point observer rating scales; body size was measured by humeral articular surfaces; age and sex were determined previously through standard procedures. Using sex separated rankings and partial correlations, disaggregated muscle markers were examined for correlations with age and size to determine if specific muscle markers may be useful in pinpointing to activity patterns. Aggregate upper limb muscle marker correlated with: age, r = 0.44; humeral size, r = 0.44; and sex, r = 0.43; Ps < 0.001. Older individuals had greater muscle markers, as did larger individuals, and males. Rankings seemed to be confounded by the effect size had on the muscle markers. However, based on partial correlations controlling for size and age, the differences that remained between males and females could be used to reconstruct male activities of throwing in hunting and interpersonal aggression acts.