Ontogeny and functional adaptation of trabecular bone in the human foot
Trabecular bone forms the internal scaffolding of most bones, and consists of a microscopic lattice-like structure of interconnected bony struts. Experimental work has demonstrated that trabecular bone adapts its structural rigidity and orientation in response to the strains placed upon the skeleton during life, a concept popularly known as “Wolff’s Law” or “bone functional adaptation”. Anthropological work has focused on correlating variation in primate trabecular bone to locomotor and masticatory function, to provide a context for the interpretation of fossil morphology. However, intraspecies variation and its underlying mechanisms are still poorly understood. In this thesis, variation in trabecular bone structure is examined in the human foot in four archaeological populations. The aim is to tease apart the factors underlying variation in human trabecular microstructure to determine whether it may be a suitable proxy for inferring terrestrial mobility in past populations. µCT scanning is used to image the three-dimensional trabecular structure of the talus, calcaneus, and first metatarsal in samples from four archaeological populations. Trabecular structure is quantified in seventeen volumes of interest placed throughout the foot. Trabecular bone is influenced by a variety of factors including body mass, age, diet, temperature, genetics, sex, and mechanical loading. Before trabecular structure can be used to infer habitual behaviour, the effects of these factors need to be understood and ideally statistically accounted for. Therefore, the effects of variation in bone size and shape, body mass, age, and sex on human trabecular structure are examined in four populations. Significant effects of body mass and age are reported, but little sexual dimorphism was found within populations. Taking these results into account, variation in trabecular structure is compared between archaeological populations that were divided into high and low mobility categories. Results demonstrate that the four populations show similar patterns of trabecular variation throughout the foot, with a signal of terrestrial mobility level superimposed upon it. Terrestrial mobility is associated with greater bone volume fraction and thicker, more widely spaced, and less interconnected trabeculae. Ontogeny of trabecular bone in the human calcaneus is investigated in two archaeological populations in the final chapter of the thesis. Results indicate that calcaneal trabecular bone adapts predictably to changes in loading associated with phases of gait maturation and increases in body mass. This opens the possibility of using trabecular structure to serve as a proxy of neuromuscular development in juvenile hominins. This work demonstrates that trabecular bone may serve as a useful proxy of habitual behaviour in hominin fossils and past populations when all contributing factors are carefully considered and ideally statistically controlled for.