This study uses synchrotron radiation-based micro-computed tomography (CT) scans to reconstruct three-dimensional networks of Haversian systems in human cortical bone in order to observe and analyse interconnectivity of Haversian systems and the development of total Haversian networks across different ages. A better knowledge of how Haversian systems interact with each other is essential to improve understanding of remodeling mechanisms and bone maintenance; however, previous methodological approaches (e.g. serial sections) did not reveal enough detail to follow the specific morphology of Haversian branching, for example. Accordingly, the aim of the present study was to identify the morphological diversity of branching patterns and transverse connections, and to understand how they change with age. Two types of branching morphologies were identified: lateral branching, resulting in small osteon branches bifurcating off of larger Haversian canals; and dichotomous branching, the formation of two new osteonal branches from one. The reconstructions in this study also suggest that Haversian systems frequently target previously existing systems as a path for their course, resulting in a cross-sectional morphology frequently referred to as 'type II osteons'. Transverse connections were diverse in their course from linear to oblique to curvy. Quantitative assessment of age-related trends indicates that while in younger human individuals transverse connections were most common, in older individuals more evidence of connections resulting from Haversian systems growing inside previously existing systems was found. Despite these changes in morphological characteristics, a relatively constant degree of overall interconnectivity is maintained throughout life. Altogether, the present study reveals important details about Haversian systems and their relation to each other that can be used towards a better understanding of cortical bone remodeling as well as a more accurate interpretation of morphological variants of osteons in cross-sectional microscopy. Permitting visibility of reversal lines, synchrotron radiation-based micro-CT is a valuable tool for the reconstruction of Haversian systems, and future analyses have the potential to further improve understanding of various important aspects of bone growth, maintenance and health.
Xcambó is a Classic period Maya site (250-700 AD) situated on the northern coast of Yucatan, Mexico. Archaeological evidence suggests that the site began as a salt production center but adopted a more administrative role as a commercial port in the Late Classic period. Economic growth, depending on its magnitude, could have affected the daily occupations of Xcambó's inhabitants. However, this is difficult to infer from the archaeological record. The aim of this study was to directly evaluate this possibility through skeletal analysis. Since diaphyseal robusticity and shape are predominantly influenced by mechanical loading history, long bone cross-sections can be used to access activity patterns. To this end, humeri and femora of 47 male and 35 female adult specimens from two Xcambó population samples were scrutinized. Our analysis satisfies general archaeological expectations and provides additional information on the population's physical response to economic growth. Decreasing robusticity and femoral anterior-posterior rigidity indicate an overall decrease in physical workload and mobility, concomitant with the site's increasing administrative function. We also observed a significant decrease in sexual dimorphism, possibly attributable to the differential response of male and female physical work spheres during socioeconomic change. In general, our findings suggest even nonsubsistence based socioeconomic change can significantly affect the bone structure of a population, rendering activity analysis an important aspect of the reconstruction of living conditions of past populations.
This study compares two novel methods quantifying bone shaft tissue distributions, and relates observations on human humeral growth patterns for applications in anthropological and anatomical research. Microstructural variation in compact bone occurs due to developmental and mechanically adaptive circumstances that are 'recorded' by forming bone and are important for interpretations of growth, health, physical activity, adaptation, and identity in the past and present. Those interpretations hinge on a detailed understanding of the modeling process by which bones achieve their diametric shape, diaphyseal curvature, and general position relative to other elements. Bone modeling is a complex aspect of growth, potentially causing the shaft to drift transversely through formation and resorption on opposing cortices. Unfortunately, the specifics of modeling drift are largely unknown for most skeletal elements. Moreover, bone modeling has seen little quantitative methodological development compared with secondary bone processes, such as intracortical remodeling. The techniques proposed here, starburst point-count and 45° cross-polarization hand-drawn histomorphometry, permit the statistical and populational analysis of human primary tissue distributions and provide similar results despite being suitable for different applications. This analysis of a pooled archaeological and modern skeletal sample confirms the importance of extreme asymmetry in bone modeling as a major determinant of microstructural variation in diaphyses. Specifically, humeral drift is posteromedial in the human humerus, accompanied by a significant rotational trend. In general, results encourage the usage of endocortical primary bone distributions as an indicator and summary of bone modeling drift, enabling quantitative analysis by direction and proportion in other elements and populations.
Bone is a dynamic tissue, responding locally to differential mechanical loading and systemically to hormonal stimuli. Although the tendency is to consider compact bone a homogenous tissue, regional variations in microarchitecture are known to have quite different properties relating to processes of growth and mechanical loading. Specifically, we investigated an example of microstructural variation of the human, adult femoral endocortex, referred to here as the endosteal lamellar pocket (ELP). The femoral ELP is characterised as a histomorphological meta-feature, made distinct from general circumferential lamellae by its hemicircumferential lamellar orientation, medial positioning, radially oriented Volkmann's canals and decreased presence of Haversian systems. Our preliminary observations of mid-diaphyseal transverse thin-ground sections demonstrate the ELP as a regular meta-feature of adult human femora unearthed from the Maya archaeological site of Xcambó , Mexico. ELPs were easily identified in 38 of 45 examined individuals. Their size, shape and degree of secondary remodelling were variable. In comparison, ELP lamellae and those originating from the periosteum are divergent, or non-concentric in orientation. This divergence, along with the ELP's medial position, suggest it forms during growth through endosteal modelling drift. Possible considerations for ELP formation and persistence with age are discussed as well as its implications for the study of bone dynamics during growth and activity. Further research is necessary to quantitatively measure the morphology of ELPs and to identify any co-variance between ELP characteristics and age, sex or indicators of mechanical loading.
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