Ontogenetic changes in femoral cross-sectional geometry during childhood locomotor development.

OBJECTIVES: The femur is a major weight-bearing bone that is variably loaded throughout growth as children transition through locomotory states prior to the attainment of a mature bipedal gait. Here, we document ontogenetic trends in femoral cross-sectional geometry (CSG) and explore how changes in loading regime may impact the structural arrangement of cortical bone along the length of the developing diaphysis. MATERIALS AND METHODS: Micro-CT scans of 110 immature femora were generated from a documented archaeological sample ranging in age from birth to 8.5 years old. CSG properties indicative of relative bone strength and bending rigidity were analyzed from cross-sections extracted at 35%, 50% and 65% of total intermetaphyseal length. RESULTS: Infants experience a marked redistribution of cortical bone between birth and 7 months facilitating a more advantageous mechanical structure for early load bearing behaviors as bone is displaced further from the section centroid. Early walkers are characterized by a mediolaterally reinforced cross-section that becomes more circular as gait continues to develop. DISCUSSION: During ontogeny the femur undergoes distinct morphological phases, which correspond with changes in loading regime. This study illustrates the importance of loading conditions in shaping immature bone morphology. Nonmechanical factors such as changes in hormonal environmental can also impact on this dynamic.

Sex differences in the patterning of age-related bone loss in the human hallucal metatarsal in rural and urban populations.

OBJECTIVES: Age-degenerative features of the metatarsals are poorly known despite the importance of metatarsal bone properties for investigating mobility patterns. We assessed the role of habitual activity in shaping the patterning and magnitude of sexual dimorphism in age-related bone loss in the hallucal metatarsal. MATERIALS AND METHODS: Cross-sections were extracted at midshaft from micro-computed tomography scan models of individuals from medieval rural (Abingdon Vineyard) and early industrial urban (Spitalfields) settings (n = 71). A suite of cross-sectional geometry dimensions and biomechanical properties were compared between populations. RESULTS: The rural group display generally stronger and larger metatarsals that show a greater capacity to resist torsion and that have comparatively greater bending strength along the medio-lateral plane. Men in both groups show greater values of cortical area than women, but only in the urban group do men show lower magnitudes of age-related decline compared to females. Women in rural and urban populations show different patterns of age-related decline in bone mass, particularly old women in the urban group show a marked decline in cortical area that is absent for women in the rural group. DISCUSSION: Lifetime exposure to hard, physical activity in an agricultural setting has contributed to the attainment of greater bone mass and stronger bones in young adults. Furthermore, over the life-course, less of this greater amount of bone is lost, such that sustained activity levels may have acted to buffer against age-related decline, and this is most pronounced for women, who are expected to experience greater bone loss later in life than men.

Age estimation of immature human skeletal remains from the dimensions of the girdle bones in the postnatal period.

OBJECTIVES: This study provides classical calibration regression formulae for age estimation from the dimensions of unfused shoulder and pelvic girdle bones. MATERIALS AND METHODS: Age estimation models were derived from a sample of 160 known age and sex individuals (63 females and 97 males) aged birth to 12 years, selected from Portuguese and English skeletal collections. The sample was divided into two age groups at the age of 2 years, and formulae were obtained for the sexes separately and combined. RESULTS: Measurements of the pelvis provide more precise age estimates than the shoulder. In the younger age group, the height and width of the ilium, and the height of the glenoid yield the most precise age estimates. In the older age group, the length of the clavicle provides the most precise estimates, followed by measurements of the pubis and ischium. DISCUSSION: In the younger individuals (<2 years), age estimates based on measurements of the pelvic girdle seem to be as or more precise than those based on the length of long bones. In older individuals (≥2 years), estimates based on the measurements of the girdles are less precise than those based on the length of long bones. These age estimation formulae may be useful for fragmentary and incomplete material in archaeological and forensic contexts. The formulae are suitable for a variety of archeological populations living under adverse conditions. These conditions are similar to some "developing" countries, and hence the formulae may also be applicable to modern forensic remains from such environments.

Quantification of 3D curvature in the iliac crest: Ontogeny and implications for sex determination in juveniles.

OBJECTIVES: Sex-specific differences in the shape of the iliac crest have been reported based on quantification of simplified curvature using two-dimensional variables or qualitative identification of faint/marked S-shaped curvature. An objective and quantitative approach for the evaluation of iliac crest curvature in juveniles was developed. Using three-dimensional data, ontogenetic variation in iliac curvature was investigated in relation to size and chronological age and its accuracy for sex estimation among juveniles was evaluated. MATERIALS AND METHODS: Three-dimensional geometric morphometric landmarks were collected from virtual models of 53 surface-scanned ilia and used to: (1) quantify differences between male and female shape, and (2) to calculate a measure of sinuosity by extracting linear measurements relative to a plane constructed in virtual space. RESULTS: Males were found to exhibit on average more pronounced curvature of the iliac crest than females in equivalent age groups, and the difference between sexes was more marked on the posterior section of the bone than on the anterior section. Classification accuracy was higher for measurements of sinuosity relative to a three-dimensional plane (62-87%) than for landmarks describing crest shape (64-74%). CONCLUSIONS: An increased likelihood of correct male identification (reduced rate of false positive identification) was observed in older children (>5 years). This reflects divergence of male and female iliac crest shape with size (or age), and the development of a unique, more curved iliac crest shape in males. The results suggest more conspicuous male trait expression than female trait expression.

Age estimation of immature human skeletal remains from the metaphyseal and epiphyseal widths of the long bones in the post-natal period.

OBJECTIVES: This study provides regression and classical calibration models for the estimation of age from the dimensions of the metaphyses and epiphyses of the six long bones. METHODS: A sample of 148 known sex and age individuals (56 females and 92 males), aged between birth and 12 years of age, selected from Portuguese and English skeletal collections. Age estimation models were obtained separately for each sex and for the sexes combined, for the entire sample and for the sample divided into two subsamples at the age of 2 years. RESULTS: The best performing epiphysis is the proximal epiphysis of the tibia. For the metaphyses, the distal metaphysis of the tibia and the proximal metaphysis of the femur are best for individuals below and above 2 years of age, respectively. Growth of the metaphyses is similar to that of the diaphysis. Results suggest that age can be as accurately estimated from the width of the metaphyses and epiphyses as from the length of the diaphyses, or as from the length of the deciduous and permanent teeth. CONCLUSIONS: These models may be useful for fragmentary material in both archaeological and forensic contexts. However, due to the background of the samples, the models would be most applicable to individuals exposed to adverse environmental conditions during growth and development. Metaphyseal and epiphyseal widths may be less affected by ecological conditions than diaphyseal length.

Earliest evidence for caries and exploitation of starchy plant foods in Pleistocene hunter-gatherers from Morocco.

Dental caries is an infectious disease that causes tooth decay. The high prevalence of dental caries in recent humans is attributed to more frequent consumption of plant foods rich in fermentable carbohydrates in food-producing societies. The transition from hunting and gathering to food production is associated with a change in the composition of the oral microbiota and broadly coincides with the estimated timing of a demographic expansion in Streptococcus mutans, a causative agent of human dental caries. Here we present evidence linking a high prevalence of caries to reliance on highly cariogenic wild plant foods in Pleistocene hunter-gatherers from North Africa, predating other high caries populations and the first signs of food production by several thousand years. Archaeological deposits at Grotte des Pigeons in Morocco document extensive evidence for human occupation during the Middle Stone Age and Later Stone Age (Iberomaurusian), and incorporate numerous human burials representing the earliest known cemetery in the Maghreb. Macrobotanical remains from occupational deposits dated between 15,000 and 13,700 cal B.P. provide evidence for systematic harvesting and processing of edible wild plants, including acorns and pine nuts. Analysis of oral pathology reveals an exceptionally high prevalence of caries (51.2% of teeth in adult dentitions), comparable to modern industrialized populations with a diet high in refined sugars and processed cereals. We infer that increased reliance on wild plants rich in fermentable carbohydrates and changes in food processing caused an early shift toward a disease-associated oral microbiota in this population.

A virtual geometric morphometric approach to the quantification of long bone bilateral asymmetry and cross-sectional shape.

OBJECTIVE: The widespread use of three-dimensional digitization means that models of whole bone geometry are routinely captured for many applications in virtual anthropology. In this article, we test whether a geometric morphometric (GMM) approach can be used to accurately quantify directional bilateral asymmetry in cross-sectional shape using virtual bone models. METHODS: We introduce a method to (1) orient virtual long bone models and extract cross sections and (2) collect GMM landmarks to analyze cross-sectional shape. Landmark data, captured using polar radii, were analyzed for an example study set of paired humeri from Andaman Islanders sampled at 35%, 50%, and 65% locations. RESULTS: The GMM method can (1) detect significant differences in left and right cross-sectional shape and (2) allow the directionality of shape change (distribution of bone from centroid) to be evaluated in the context of whole outline shape and in relation to the axis of maximum bending rigidity. The protocol may be used with models created from laser (surface) scan or computed tomography (CT) scan data, and applied to cross-sectional images that were collected using periosteal molding techniques. CONCLUSIONS: The results demonstrate directional bilateral asymmetry in shape, but do not recover the same signal for size measurements. Our method offers a pathway to quantify both the pattern of variation in shape and the relationship between size and shape variation, opening new questions about how those patterns manifest over ontogeny, change temporally or differ in relation to the nature and intensity of the activity, and bone loading conditions.

Shape, size, and maturity trajectories of the human ilium.

Morphological traits of the ilium have consistently been more successful for juvenile sex determination than have techniques applied to other skeletal elements, however relatively little is known about the ontogeny and maturation of size and shape dimorphism in the ilium. We use a geometric morphometric approach to quantitatively separate the ontogeny of size and shape of the ilium, and analyze interpopulation differences in the onset, rate and patterning of sexual dimorphism. We captured the shape of three traits for a total of 191 ilia from Lisbon (Portugal) and London (UK) samples of known age and sex (0-17 years). Our results indicate that a) there is a clear dissociation between the ontogeny of size and shape in males and females, b) the ontogeny of size and shape are each defined by non-linear trajectories that differ between the sexes, c) there are interpopulation differences in ontogenetic shape trajectories, which point to population-specific patterning in the attainment of sexual dimorphism, and d) the rate of shape maturation and size maturation is typically higher for females than males. Male and female shape differences in the ilium are brought about by trajectory divergence. Differences in size and shape maturation between the sexes suggest that maturity may confound our ability to discriminate between the sexes by introducing variation not accounted for in age-based groupings. The accuracy of sex determination methods using the ilium may be improved by the use of different traits for particular age groups, to capture the ontogenetic development of shape in both sexes.

Age estimation of immature human skeletal remains from the diaphyseal length of the long bones in the postnatal period.

Age at death in immature human skeletal remains has been estimated from the diaphyseal length of the long bones, but few studies have actually been designed specifically for the purpose of age estimation and those which have, show important caveats. This study uses regression and classical calibration to model the relationship between age and diaphyseal length of the six long bones, in a sample of 184 known sex and age individuals (72 females and 112 males), younger than 13 years of age, selected from Portuguese and English skeletal collections. Age estimation models based on classical calibration were obtained for each of the six long bones, and separately for each sex and for the sexes combined, and also for the entire sample and when it is subdivided into two subsamples at the age of 2 years. Comparisons between inverse and classical calibration show there is a systematic bias in age estimations obtained from inverse calibration. In the classical calibration models, the length of the femur provides the most accurate estimates of age. Age estimates are more accurate for the male subsample and for individuals under the age of 2 years. These results and a test of previously published methods caution against inverse calibration as a technique for developing age estimation methods even from the immature skeleton. Age estimation methods developed using cemetery collections of identified human skeletons should not be uncritically applied to present-day populations from the same region since many populations have experienced dramatic secular trends in growth and adult height over the last century.

Isotopic and trace element evidence of dietary transitions in early life.

CONTEXT: Nursing behaviour has significant implications for individual health and population dynamics. Reconstruction of infant and early childhood diet has become a key focus in studies of past populations. OBJECTIVE: This paper reviews knowledge about how trace element and stable isotope composition of bone and dental tissues changes during development and how this may influence interpretation of biology and behaviour in past populations. METHODS: Studies of recent populations and osteological samples were reviewed. RESULTS: Four distinct stages of nutritional intake are recognized: gestation, exclusive breastfeeding, a transitional or weaning stage and fully weaned. The trace element and stable isotope composition of developing foetal and infant tissues differs during each of these stages due to differences in the source of nutrients and changes in metabolic parameters. CONCLUSION: Interpretation of early lifetime diet from trace element or stable isotope values requires a comprehensive and validated physiological model for changes during and at the transition between dietary stages. Micro-sampling of dental tissues offers the potential to reconstruct an individual record of early lifetime diet within a chronological framework and has several advantages over traditional cross-sectional techniques. Further research is necessary to understand the factors influencing intra-tooth variability in recovered signals and their chronological interpretation.

Weaning behaviour in human evolution.

Human life history incorporates early weaning, a prolonged period of post-weaning dependency and slow somatic growth, late onset of female reproduction, reduced birth spacing and a significant post-reproductive female lifespan, combined with rapid early brain growth. Weaned human offspring lack the cognitive skills and physical capacity required to locate, procure and prepare foods that are appropriate for their immature state and sufficient for their high energy requirements. During the weaning process and throughout childhood human offspring are supported by the provision of energy dense and easily digestible foods. Changes in weaning behaviour during human evolution imply a shift in the balance between maternal costs of lactation and the risk of poor offspring outcome, and may have been driven by an increase in infant nutritional and metabolic requirements, a reduction reproductive lifespan resulting in selection for reduced birth spacing or a change in other factors affecting offspring survival and fitness.

Body mass and stature estimation based on the first metatarsal in humans.

Archaeological assemblages often lack the complete long bones needed to estimate stature and body mass. The most accurate estimates of body mass and stature are produced using femoral head diameter and femur length. Foot bones including the first metatarsal preserve relatively well in a range of archaeological contexts. In this article we present regression equations using the first metatarsal to estimate femoral head diameter, femoral length, and body mass in a diverse human sample. The skeletal sample comprised 87 individuals (Andamanese, Australasians, Africans, Native Americans, and British). Results show that all first metatarsal measurements correlate moderately to highly (r = 0.62-0.91) with femoral head diameter and length. The proximal articular dorsoplantar diameter is the best single measurement to predict both femoral dimensions. Percent standard errors of the estimate are below 5%. Equations using two metatarsal measurements show a small increase in accuracy. Direct estimations of body mass (calculated from measured femoral head diameter using previously published equations) have an error of just over 7%. No direct stature estimation equations were derived due to the varied linear body proportions represented in the sample. The equations were tested on a sample of 35 individuals from Christ Church Spitalfields. Percentage differences in estimated and measured femoral head diameter and length were less than 1%. This study demonstrates that it is feasible to use the first metatarsal in the estimation of body mass and stature. The equations presented here are particularly useful for assemblages where the long bones are either missing or fragmented, and enable estimation of these fundamental population parameters in poorly preserved assemblages.

Using stable isotope analysis to examine the effect of economic change on breastfeeding practices in Spitalfields, London, UK.

Breastfeeding patterns were subject to a number of fads in 18th and 19th century Britain. Feeding infants by hand, rather than maternal breastfeeding or wet-nursing, became more prevalent among both the wealthy and poor. Substitute foods may have been a convenient alternative for mothers employed away from the household. This study used stable isotope ratio analysis to examine the weaning schedule in the 18th and 19th century skeletal assemblage from Spitalfields, London, UK. Analysis of 72 juvenile ribs revealed δ(15) N elevations of 2-3‰ above the adult mean for individuals up to the age of two, while elevations of 1-2‰ were observed in δ(13) C for the first year of life. This suggests that the introduction of solid foods took place before the end of the first year, and that breastfeeding had entirely ceased by 2 years of age. The age at death of many of these infants is known from historical records, and can be used to pinpoint the amount of time required for the breast milk signal to be observed in the stable isotope ratios of rib collagen. Results show that a δ(15) N elevation can be detected in the ribs of individuals who died as young as 5-6 weeks. Not all individuals at Spitalfields were breastfed, and there may not have been a single uniformly practiced weaning scheme. There is, however, more evidence for prolonged breastfeeding during the 19th century than the 18th century.

Unlocking evidence of early diet from tooth enamel.

Recent developments in microspatial analysis of enamel chemistry provide the resolution needed to reconstruct detailed chronological records of an individual's early life history. Evidence of nutritional history, residential mobility, and exposure to heavy metals can potentially be retrieved from archaeological and even fossil teeth. Understanding the pattern and timing of incorporation of each trace element or stable isotope into enamel is crucial to the interpretation of the primary data. Here, we use laser ablation inductively coupled plasma mass spectrometry and ArcGIS software to map variation in calcium-normalized strontium intensities across thin sections of enamel from exfoliated deciduous teeth. Differences in calcium-normalized strontium intensities across each tooth reflect variation in tooth mineralization, implying that sampling location must be taken into account in interpreting results. Chronologically consistent shifts in calcium-normalized strontium intensities in teeth from children with known nursing histories reflect the onset and duration of breastfeeding and the introduction of nonmaternal sources of food. This tool is likely to be valuable for studying weaning and nursing behavior in the past. The distribution of normalized strontium intensities presented here is consistent with a model for the differential incorporation of strontium and calcium into enamel during the secretory and maturational phases of formation.

Age estimation of immature human skeletal remains from mandibular and cranial bone dimensions in the postnatal period.

Age estimation is one of the crucial first steps in the identification of human skeletal remains in both forensic and archeological contexts. In the postnatal period, age is traditionally estimated from dental development or skeletal growth, typically long bone diaphyseal length. However, in many occasions other methods are required. This study provides alternative means of estimating age of juvenile remains from the size of several cranial bones and the mandible. A sample of 185 identified juvenile skeletons between birth and 13 years of age from two European collections were used (Lisbon and Spitalfields). Measurements of the frontal, occipital-lateralis, occipital-basilaris, occipital-squamous, zygomatic, maxilla, and mandible were used to calculate classical calibration regression formulae for the sexes combined. The sample was divided into three age groups birth-2 years, 2-6 years, and 2-12.9 years, depending on bone and its growth trajectory. For all the bones, measurements of the youngest age groups yielded the most precise age estimates. The vault bones on average yielded the best performing models, with the frontal bone having the most precise of all. The mandible performed on par with the best performing cranial bones, particularly in individuals under the age of 2 years. This study provides one of the most comprehensive approaches to juvenile age estimation based on bones of the skull, providing a resource that potentially can help estimate age of juvenile skeletons from a variety of circumstances.

The relationship of accentuated lines in enamel to weaning stress in juvenile baboons (Papio hamadryas anubis).

Weaning conflict may represent an evolutionary conflict of interest between parent and offspring, an honest signal of need on the part of a weanling, or both. Accentuated lines visible in histological sections of teeth are indicators of stress during enamel formation and have been hypothesised to form in baboon teeth during weaning. We analysed growth increments in 5 tooth sections from 2 Ugandan baboons (Papio hamadryas anubis), using polarised light microscopy, to determine when stresses occurred during the weaning process. Dietary transitions were reconstructed using normalised strontium intensities (Sr/Ca) in enamel. Accentuated lines were cross-matched between teeth from the same animal and plotted by month. The highest frequency of stress was experienced at around 6 months in 1 baboon, coinciding with an inferred reduction in suckling frequency, and at 11 months in another, coinciding with the inferred cessation of suckling. Because accentuated lines appear to indicate weaning stress at dietary transitions, weaning conflict may represent an honest signal of need on the part of the weanling.

Morphometric criteria for sexing juvenile human skeletons using the ilium.

Previous attempts to sex juvenile skeletons have focused on the application of qualitative or semi-quantitative techniques. This study applies a variety of geometric morphometric methods, including eigenshape analysis, to this problem. Six metric criteria for the ilia were tested with the aim of investigating previous ideas concerning sexually diagnostic characters. This study uses 25 ilia from juveniles of known age and sex from Christ Church, Spitalfields, London. Ninety-six percent of juvenile ilia were correctly identified as male or female using the shape of the greater sciatic notch. Identification accuracy is shown to improve with age for several criteria. Males were identified to a higher accuracy than females. Application of geometric techniques improves the understanding of the relationship between age, sex, and shape and the clarity with which these relationships can be quantified. Archaeological and forensic relevance of the results are discussed with recommendations for future application.

Tracking dietary transitions in weanling baboons (Papio hamadryas anubis) using strontium/calcium ratios in enamel.

Strontium and calcium are incorporated into developing teeth in a manner that reflects changing physiological concentrations in the body. A new model predicts changes in strontium/calcium (Sr/Ca) ratios in response to dietary transitions experienced at birth and during the weaning period. Microsampling of longitudinal thin sections of tooth enamel using laser ablation inductively coupled plasma mass spectrometry provides a basis for the systematic evaluation of variation in Sr/Ca ratios within the tooth crown. Incremental growth markers in enamel are used to determine the age of onset of enamel mineralization at each sampling point. Thin sections of 5 teeth from 2 wild-caught baboons (Papio hamadryas anubis) were systematically analysed using this technique. Intra- and intertooth analyses of Sr/Ca ratios reveal a pattern of dietary development during the period of enamel formation that is consistent with observational data on the timing of weaning behaviour in anubis baboons.

90,000 year-old specialised bone technology in the Aterian Middle Stone Age of North Africa.

The question of cognitive complexity in early Homo sapiens in North Africa is intimately tied to the emergence of the Aterian culture (~145 ka). One of the diagnostic indicators of cognitive complexity is the presence of specialised bone tools, however significant uncertainty remains over the manufacture and use of these artefacts within the Aterian techno-complex. In this paper we report on a bone artefact from Aterian Middle Stone Age (MSA) deposits in Dar es-Soltan 1 cave on the Atlantic coast of Morocco. It comes from a layer that can be securely dated to ~90 ka. The typological characteristics of this tool, which suggest its manufacture and use as a bone knife, are comparatively similar to other bone artefacts from dated Aterian levels at the nearby site of El Mnasra and significantly different from any other African MSA bone technology. The new find from Dar es-Soltan 1 cave combined with those from El Mnasra suggest the development of a bone technology unique to the Aterian.

On the reliability of a geometric morphometric approach to sex determination: a blind test of six criteria of the juvenile ilium.

Despite the attention of many studies, researchers still struggle to identify criteria with which to sex juvenile remains at levels of accuracy and reproducibility comparable with those documented for adults. This study uses a sample of 82 juvenile ilia from an identified Portuguese population (Lisbon collection) to test the cross-applicability of a new approach by Wilson et al. [23] that uses geometric morphometric methods to sex the subadult ilium. Further, we evaluate the wider applicability of these methods for forensic casework, extending the age range of the original study by examining an additional 19 juvenile ilia from the St. Brides and Spitalfields collections, housed in London. Levels of accuracy for the Portuguese sample (62.2-89.0%) indicate that the methods can be used to document dimorphism in another sample. Discriminant functions are sample-specific, indicated by not better than average classification using cross-validation. We propose a methodological update, whereby we recommend disuse of the auricular surface morphology criterion, based upon reduced success rates and inadequate accuracy of female identification. We show, in addition to population differences, differences in the ontogeny of dimorphism may lead to differing degrees of success for female identification using some criteria. The success rates are highest between the ages of 11.00 and 14.99 years (93.3% males, 80.0% females).