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Abstract: The origin of jaws and teeth remains contentious in vertebrate evolution. ‘Placoderms’ (Silurian-Devonian armoured jawed fishes) are central to debates on the origins of these anatomical structures. ‘Acanthothoracids’ are generally considered the most primitive ‘placoderms’. However, they are so far known mainly from disarticulated skeletal elements that are typically incomplete. The structure of the jaws—particularly the jaw hinge—is poorly known, leaving open questions about their jaw function and comparison with other placoderms and modern gnathostomes. Here we describe a near-complete ‘acanthothoracid’ upper jaw, allowing us to reconstruct the likely orientation and angle of the bite and compare its morphology with that of other known ‘placoderm’ groups. We clarify that the bite position is located on the upper jaw cartilage rather than on the dermal cheek and thus show that there is a highly conserved bite morphology among most groups of ‘placoderms’, regardless of their overall cranial geometry. Incorporation of the dermal skeleton appears to provide a sound biomechanical basis for jaw origins. It appears that ‘acanthothoracid’ dentitions were fundamentally similar in location to that of arthrodire ‘placoderms’, rather than resembling bony fishes. Irrespective of current phylogenetic uncertainty, the new data here resolve the likely general condition for ‘placoderms’ as a whole, and as such, ancestral morphology of known jawed vertebrates.

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Abstract: Accurately estimating developmental age and life history traits in fossils is crucial for identifying and classifying extinct species and understanding how biological attributes evolved. The evolution of life history traits such as growth pattern is far from clear in birds, and development has been studied in only a handful of modern species. The exceptionally rapid growth of modern birds means ageing methods based on annual incremental growth lines, used in other vertebrates, are inapplicable to birds and robust alternative methods remain to be established. Analysis of avian intracortical bone microstructure, which varies both with age and tissue deposition rate, is a promising approach already used in palaeobiology. However, current thin section-based histological methods are destructive. Moreover, to date, most microstructural studies in avian bone are qualitative, 2D, and involve a limited range of extant species. The objective of this study was to investigate cortical bone microstructure and developmental age and life history traits in living birds, to identify phenotypes which can then be applied to examination of the fossil record, using minimally-destructive high-resolution 3D imaging. First, the necessity of 3D measurement was tested: a combination of idealised, simulated datasets and real synchrotron-based computed tomography (SR CT) datasets were used to compare published methods for measuring key microstructural traits based on 2D sections and 3D volumes. Next, SR CT imaging and quantitative measurements were used to characterise age-related changes in bone microstructure in a range of extant bird species: growth series ducks and pheasants, and a smaller sample size in starlings, rock doves, partridges, and ostrich. The methods tested in modern material were applied to fossils as a proof-of concept. It was found that 3D quantification methods are required for measuring vascular canal orientation and osteocyte lacunar shape and volume, though 2D sections could be used to measure traits such as bone volume fraction (BV/TV) and osteocyte lacunar volume. In all species studied, juvenile, subadult, and adult species could be distinguished by their values of BV/TV, and further information could be added using measured values of vascular canal diameter as well as qualitative assessment. Using a synchrotron-based CT system, high-resolution 3D datasets comparable to modern bone samples were obtained from fossils, and preliminary estimates of developmental age can be made. Further work may reveal more changes within juvenile age stages, and better characterise the variation within extant birds, allowing more accurate interpretation of the fossil record. Therefore developmental studies in a greater number of extant bird species are required using larger sample sizes, to support and add to the results presented in this thesis.

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Abstract: Small-diameter, thin-walled pipes have applications in a wide range of industries including high-energy physics, heat transfer, nuclear, medical and communications. There are no standards that exist for welds less than 0.5 mm in width, and as such it is difficult to determine the likely performance of a thin-walled pipe weld. Porosity is largely inevitable in fusion welded joints and is a determining factor in the performance of a connection. This study focused on characterisation of the evolution of strains in soldered welds less than 0.5 mm in width, by incrementally tensile loading samples and studying them in-situ with Synchrotron X-Ray Computed Tomography and X-Ray Diffraction. Two sample geometries were studied, and porosity defects were present in both, although the levels of porosity size, number and area varied dramatically between the two samples. Lattice strain interpretation showed that crack propagation for such samples is not driven by porosity but that crack evolution occurs at the same location and load levels irrespective of the presence of pores. Residual stresses of up to 0.3% from the fusion welding process were seen in both samples and appear to have a greater impact on locations of failure than porosity. Porosity does cause differences in strains across directions, however high strains alone did not appear to cause premature failure. Hence, efforts to improve weld strength should in future focus more on reducing residual stresses than reducing porosity.

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Abstract: We present a flyscan compatible acquisition scheme for three-modal X-Ray Computed Tomography (CT) with two-dimensional phase sensitivity. Our approach is demonstrated using a “beam tracking” setup, through which a sample’s attenuation, phase (refraction) and scattering properties can be measured from a single frame, providing three complementary contrast channels. Up to now, such setups required the sample to be stepped at each rotation angle to sample signals at an adequate rate, to prevent resolution losses, anisotropic resolution, and under-sampling artefacts. However, the need for stepping necessitated a step-and-shoot implementation, which is affected by motors’ overheads and increases the total scan time. By contrast, our proposed scheme, by which continuous horizontal and vertical translations of the sample are integrated with its rotation (leading to a “cycloidal-spiral” trajectory), is fully compatible with continuous scanning (flyscans). This leads to greatly reduced scan times while largely preserving image quality and isotropic resolution.

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Abstract: Skeletal muscle contractions are critical for normal skeletal growth and morphogenesis but it is unclear how the detrimental effects of absent muscle on the bones and joints change over time. Joint shape and cavitation as well as rudiment length and mineralisation were assessed in multiple rudiments at two developmental stages [Theiler stage (TS)24 and TS27] in the splotch-delayed “muscle-less limb” mouse model and littermate controls. Chondrocyte morphology was quantified in 3D in the distal humerus at the same stages. As development progressed, the effects of absent muscle on all parameters except for cavitation become less severe. All major joints in muscle-less limbs were abnormally shaped at TS24, while, by TS27, most muscle-less limb joint shapes were normal or nearly normal. In contrast, any joints that were fused at TS24 did not cavitate by TS27. At TS24, chondrocytes in the distal humerus were significantly smaller in the muscle-less limbs than in controls, while by TS27, chondrocyte volume was similar between the two groups, offering a cell-level mechanism for the partial recovery in shape of muscle-less limbs. Mineralisation showed the most pronounced changes over gestation. At TS24, all muscle-less rudiments studied had less mineralisation than the controls, while at TS27, muscle-less limb rudiments had mineralisation extents equivalent to controls. In conclusion, the effects of muscle absence on prenatal murine skeletogenesis reduced in severity over gestation. Understanding how mammalian bones and joints continue to develop in an environment with abnormal fetal movements provides insights into conditions including hip dysplasia and arthrogryposis.