William C. H. Parr

Toward integration of geometric morphometrics and computational biomechanics: New methods for 3D virtual reconstruction and quantitative analysis of Finite Element Models

The ability to warp three-dimensional (3D) meshes from known biological morphology to fit other known, predicted or hypothetical morphologies has a range of potential applications in functional morphology and biomechanics. One of the most challenging of these applications is Finite Element Analysis (FEA), a potentially powerful non-destructive tool in the prediction of mechanical behaviour. Geometric morphometrics is another typically computer-based approach commonly applied in morphological studies that allows for shape differences between specimens to be quantified and analysed. There has been some integration of these two fields in recent years. Although a number of shape warping approaches have been developed previously, none are easily accessible. Here we present an easily accessed method for warping meshes based on freely available software and test the effectiveness of the approach in FEA using the varanoid lizard mandible as a model. We further present new statistical approaches, strain frequency plots and landmark point strains, to analyse FEA results quantitatively and further integrate FEA with geometric morphometrics. Using strain frequency plots, strain field, bending displacements and landmark point strain data we demonstrate that the mechanical behaviour of warped specimens reproduces that of targets without significant error. The influence of including internal cavity morphology in FEA models was also examined and shown to increase bending displacements and strain magnitudes in FE models. The warping approaches presented here will be useful in a range of applications including the generation and analysis of virtual reconstructions, generic models that approximate species means, hypothetical morphologies and evolutionary intermediaries.

Human remains from the Pleistocene-Holocene transition of southwest China suggest a complex evolutionary history for East Asians.

Background Later Pleistocene human evolution in East Asia remains poorly understood owing to a scarcity of well described, reliably classified and accurately dated fossils. Southwest China has been identified from genetic research as a hotspot of human diversity, containing ancient mtDNA and Y-DNA lineages, and has yielded a number of human remains thought to derive from Pleistocene deposits. We have prepared, reconstructed, described and dated a new partial skull from a consolidated sediment block collected in 1979 from the site of Longlin Cave (Guangxi Province). We also undertook new excavations at Maludong (Yunnan Province) to clarify the stratigraphy and dating of a large sample of mostly undescribed human remains from the site. Methodology/Principal Findings We undertook a detailed comparison of cranial, including a virtual endocast for the Maludong calotte, mandibular and dental remains from these two localities. Both samples probably derive from the same population, exhibiting an unusual mixture of modern human traits, characters probably plesiomorphic for later Homo, and some unusual features. We dated charcoal with AMS radiocarbon dating and speleothem with the Uranium-series technique and the results show both samples to be from the Pleistocene-Holocene transition: ∼14.3-11.5 ka. Conclusions/Significance Our analysis suggests two plausible explanations for the morphology sampled at Longlin Cave and Maludong. First, it may represent a late-surviving archaic population, perhaps paralleling the situation seen in North Africa as indicated by remains from Dar-es-Soltane and Temara, and maybe also in southern China at Zhirendong. Alternatively, East Asia may have been colonised during multiple waves during the Pleistocene, with the Longlin-Maludong morphology possibly reflecting deep population substructure in Africa prior to modern humans dispersing into Eurasia.

Finite element analysis of three patterns of internal fixation of fractures of the mandibular condyle

The most stable pattern of internal fixation for fractures of the mandibular condyle is a matter for ongoing discussion. In this study we investigated the stability of three commonly used patterns of plate fixation, and constructed finite element models of a simulated mandibular condylar fracture. The completed models were heterogeneous in the distribution of bony material properties, contained about 1.2 million elements, and incorporated simulated jaw-adducting musculature. Models were run assuming linear elasticity and isotropic material properties for bone. This model was considerably larger and more complex than previous finite element models that have been used to analyse the biomechanical behaviour of differing plating techniques. The use of two parallel 2.0 titanium miniplates gave a more stable configuration with lower mean element stresses and displacements over the use of a single miniplate. In addition, a parallel orientation of two miniplates resulted in lower stresses and displacements than did the use of two miniplates in an offset pattern. The use of two parallel titanium plates resulted in a superior biomechanical result as defined by mean element stresses and relative movement between the fractured fragments in these finite element models.

Finite element micro-modelling of a human ankle bone reveals the importance of the trabecular network to mechanical performance: New methods for the generation and comparison of 3D models

Most modelling of whole bones does not incorporate trabecular geometry and treats bone as a solid non-porous structure. Some studies have modelled trabecular networks in isolation. One study has modelled the performance of whole human bones incorporating trabeculae, although this required considerable computer resources and purpose-written code. The difference between mechanical behaviour in models that incorporate trabecular geometry and non-porous models has not been explored. The ability to easily model trabecular networks may shed light on the mechanical consequences of bone loss in osteoporosis and remodelling after implant insertion. Here we present a Finite Element Analysis (FEA) of a human ankle bone that includes trabecular network geometry. We compare results from this model with results from non-porous models and introduce protocols achievable on desktop computers using widely available softwares. Our findings show that models including trabecular geometry are considerably stiffer than non-porous whole bone models wherein the non-cortical component has the same mass as the trabecular network, suggesting inclusion of trabecular geometry is desirable. We further present new methods for the construction and analysis of 3D models permitting: (1) construction of multi-property, non-porous models wherein cortical layer thickness can be manipulated; (2) maintenance of the same triangle network for the outer cortical bone surface in both 3D reconstruction and non-porous models allowing exact replication of load and restraint cases; and (3) creation of an internal landmark point grid allowing direct comparison between 3D FE Models (FEMs).

Micro-biomechanics of the Kebara 2 hyoid and its implications for speech in Neanderthals.

The description of a Neanderthal hyoid from Kebara Cave (Israel) in 1989 fuelled scientific debate on the evolution of speech and complex language. Gross anatomy of the Kebara 2 hyoid differs little from that of modern humans. However, whether Homo neanderthalensis could use speech or complex language remains controversial. Similarity in overall shape does not necessarily demonstrate that the Kebara 2 hyoid was used in the same way as that of Homo sapiens. The mechanical performance of whole bones is partly controlled by internal trabecular geometries, regulated by bone-remodelling in response to the forces applied. Here we show that the Neanderthal and modern human hyoids also present very similar internal architectures and micro-biomechanical behaviours. Our study incorporates detailed analysis of histology, meticulous reconstruction of musculature, and computational biomechanical analysis with models incorporating internal micro-geometry. Because internal architecture reflects the loadings to which a bone is routinely subjected, our findings are consistent with a capacity for speech in the Neanderthals.

Evolution of Neck Vertebral Shape and Neck Retraction at the Transition to Modern Turtles: an Integrated Geometric Morphometric Approach

The unique ability of modern turtles to retract their head and neck into the shell through a side-necked (pleurodiran) or hidden-necked (cryptodiran) motion is thought to have evolved independently in crown turtles. The anatomical changes that led to the vertebral shapes of modern turtles, however, are still poorly understood. Here we present comprehensive geometric morphometric analyses that trace turtle vertebral evolution and reconstruct disparity across phylogeny. Disparity of vertebral shape was high at the dawn of turtle evolution and decreased after the modern groups evolved, reflecting a stabilization of morphotypes that correspond to the two retraction modes. Stem turtles, which had a very simple mode of retraction, the lateral head tuck, show increasing flexibility of the neck through evolution towards a pleurodiran-like morphotype. The latter was the precondition for evolving pleurodiran and cryptodiran vertebrae. There is no correlation between the construction of formed articulations in the cervical centra and neck mobility. An increasing mobility between vertebrae, associated with changes in vertebral shape, resulted in a more advanced ability to retract the neck. In this regard, we hypothesize that the lateral tucking retraction of stem turtles was not only the precondition for pleurodiran but also of cryptodiran retraction. For the former, a kink in the middle third of the neck needed to be acquired, whereas for the latter modification was necessary between the eighth cervical vertebra and first thoracic vertebra. Our paper highlights the utility of 3D shape data, analyzed in a phylogenetic framework, to examine the magnitude and mode of evolutionary modifications to vertebral morphology. By reconstructing and visualizing ancestral anatomical shapes, we provide insight into the anatomical features underlying neck retraction mode, which is a salient component of extant turtle classification.

Inter‐ and intra‐specific scaling of articular surface areas in the hominoid talus

The morphology of postcranial articular surfaces is expected to reflect their weight‐bearing properties, as well as the stability and mobility of the articulations to which they contribute. Previous studies have mainly confirmed earlier predictions of isometric scaling between articular surface areas and body mass; the exception to this is ‘male‐type’, convex articular surface areas, which may scale allometrically due to differences in locomotor strategies within the analysed samples. In the present study, we used new surface scanning technology to quantify more accurately articular surface areas and to test those predictions within the talus of hominoid primates, including modern humans. Our results, contrary to predictions, suggest that there are no generalised rules of articular scaling within the talus of hominoids. Instead, we suggest that articular scaling patterns are highly context‐specific, depending on the role of each articulation during locomotion, as well as taxon‐ and sex‐specific differences in locomotion and ontogenetic growth trajectories within any given sample. While this may prove problematic for inferring body mass based on articular surface area, it also offers new opportunities of gaining substantial insights into the locomotor patterns of extinct species.

Comparative Biomechanical Modeling of Metatherian and Placental Saber-Tooths: A Different Kind of Bite for an Extreme Pouched Predator.

Questions surrounding the dramatic morphology of saber-tooths, and the presumably deadly purpose to which it was put, have long excited scholarly and popular attention. Among saber-toothed species, the iconic North American placental, Smilodon fatalis, and the bizarre South American sparassodont, Thylacosmilus atrox, represent extreme forms commonly forwarded as examples of convergent evolution. For S. fatalis, some consensus has been reached on the question of killing behaviour, with most researchers accepting the canine-shear bite hypothesis, wherein both head-depressing and jaw closing musculatures played a role in delivery of the fatal bite. However, whether, or to what degree, T. atrox may have applied a similar approach remains an open question. Here we apply a three-dimensional computational approach to examine convergence in mechanical performance between the two species. We find that, in many respects, the placental S. fatalis (a true felid) was more similar to the metatherian T. atrox than to a conical-toothed cat. In modeling of both saber-tooths we found that jaw-adductor-driven bite forces were low, but that simulations invoking neck musculature revealed less cranio-mandibular stress than in a conical-toothed cat. However, our study also revealed differences between the two saber-tooths likely reflected in the modus operandi of the kill. Jaw-adductor-driven bite forces were extremely weak in T. atrox, and its skull was even better-adapted to resist stress induced by head-depressors. Considered together with the fact that the center of the arc described by the canines was closer to the jaw-joint in Smilodon, our results are consistent with both jaw-closing and neck musculature playing a role in prey dispatch for the placental, as has been previously suggested. However, for T. atrox, we conclude that the jaw-adductors probably played no major part in the killing bite. We propose that the metatherian presents a more complete commitment to the already extreme saber-tooth ‘lifestyle’.

Finite element analysis of a micromechanical model of bone and a new 3D approach to validation

Finite Element Analysis (FEA) is now widely used to analyse the mechanical behaviour of bone structures. Ideally, simulations are validated against experimental data. To date, validation of Finite Element Models (FEMs) has been 2 Dimensional (2D) only, being based on comparison with surface-mounted strain gauge readings. In this study we present a novel 3-Dimensional (3D) approach to validation that allows comparison of modelled with experimental results between any two points in 3D space throughout the structure, providing magnitude and direction data for comparison, internally and externally. Specifically, we validate a FEM of a rat tibia, including trabecular network geometry, using a material testing stage housed within a microCT scanner. We further apply novel landmark based morphometric approaches to more effectively compare modelled and experimental results. 542 landmark points on the cortical and trabecular bone surfaces of the model were selected and validated in 3D against experimental data. This approach may hold considerable potential in fields wherein a better understanding of the mechanical behaviour of trabecular networks is important, e.g., the studies of osteoporosis and trabecular loss after orthopaedic implant insertion.

Virtual Reconstruction and Prey Size Preference in the Mid Cenozoic Thylacinid, Nimbacinus dicksoni (Thylacinidae, Marsupialia)

Thylacinidae is an extinct family of Australian and New Guinean marsupial carnivores, comprizing 12 known species, the oldest of which are late Oligocene (∼24 Ma) in age. Except for the recently extinct thylacine (Thylacinus cynocephalus), most are known from fragmentary craniodental material only, limiting the scope of biomechanical and ecological studies. However, a particularly well-preserved skull of the fossil species Nimbacinus dicksoni, has been recovered from middle Miocene (∼16-11.6 Ma) deposits in the Riversleigh World Heritage Area, northwestern Queensland. Here, we ask whether N. dicksoni was more similar to its recently extinct relative or to several large living marsupials in a key aspect of feeding ecology, i.e., was N. dicksoni a relatively small or large prey specialist. To address this question we have digitally reconstructed its skull and applied three-dimensional Finite Element Analysis to compare its mechanical performance with that of three extant marsupial carnivores and T. cynocephalus. Under loadings adjusted for differences in size that simulated forces generated by both jaw closing musculature and struggling prey, we found that stress distributions and magnitudes in the skull of N. dicksoni were more similar to those of the living spotted-tailed quoll (Dasyurus maculatus) than to its recently extinct relative. Considering the Finite Element Analysis results and dental morphology, we predict that N. dicksoni likely occupied a broadly similar ecological niche to that of D. maculatus, and was likely capable of hunting vertebrate prey that may have exceeded its own body mass.