Microwear, mechanics and the feeding adaptations of Australopithecus africanus

Recent studies of dental microwear and craniofacial mechanics have yielded contradictory interpretations regarding the feeding ecology and adaptations of Australopithecus africanus. As part of this debate, the methods used in the mechanical studies have been criticized. In particular, it has been claimed that finite element analysis has been poorly applied to this research question. This paper responds to some of these mechanical criticisms, highlights limitations of dental microwear analysis, and identifies avenues of future research.     

Dietary adaptations of South African australopiths: inference from enamel prism attitude

The angle at which enamel prisms approach the wear surface holds information with regard to the stiffness of the tissue, as well as its wear resistance. Hence, analyses of prism orientation may shed light on questions of whether the thick enamel in hominins has evolved to confer stiffness or wear resistance to the teeth and may thus inform about the diet and behavioural ecology of these species. This was explored for Paranthropus robustus and Australopithecus africanus, whereby a distinction was made between prisms at the Phase I and Phase II facets. The results were compared with those obtained for Theropithecus, Macaca, and Potamochoerus for whom behavioural and/or experimental data are available, and were interpreted against simple mechanical principles. The South African hominins differ significantly in their relationships between wear facets and prism angulations. Teeth of P. robustus are better adapted to more vertical loads during mastication (Phase I), whereas those of A. africanus are better adapted to cope with more laterally-directed loads (Phase II) commonly associated with roll-crush and mastication. Overall, teeth of P. robustus appear stiffer, while those of A. africanus seem more wear resistant.     

Observations on the Feeding and Symptomatology of Xiphinema and Longidorus on Selected Host Roots

In vitro feeding of Xiphinema brevicolle, X. index and Longidorus africanus on roots of host seedlings is described. Both Xiphinema spp. fed mainly along roots rather than at tips and up to several days at a single site. Feeding of L. africanus was confined to root tips and lasted up to 15 min. No visible short term reaction of roots parasitized by the Xiphinema spp. could be discerned, but both swelling and cessation of growth of root tips were observed within 20 hr after feeding by L. africanus. Long-term (12-month) symptoms on roots of several host plants caused by cultured populations of X. brevicolle, X. index, X. italiae, L. africanus and L. brevicaudatus are described. All the Xiphinema spp. caused a thinning and distinct darkening of root systems and, at some sites, a breakdown of the cortex. Both species of Longidorus caused stubby and swollen root tips. Root symptom severity was in proportion to nematode population levels.     

Downsizing a giant: re-evaluating Dreadnoughtus body mass

Estimates of body mass often represent the founding assumption on which biomechanical and macroevolutionary hypotheses are based. Recently, a scaling equation was applied to a newly discovered titanosaurian sauropod dinosaur (Dreadnoughtus), yielding a 59 300 kg body mass estimate for this animal. Herein, we use a modelling approach to examine the plausibility of this mass estimate for Dreadnoughtus. We find that 59 300 kg for Dreadnoughtus is highly implausible and demonstrate that masses above 40 000 kg require high body densities and expansions of soft tissue volume outside the skeleton several times greater than found in living quadrupedal mammals. Similar results from a small sample of other archosaurs suggests that lower-end mass estimates derived from scaling equations are most plausible for Dreadnoughtus, based on existing volumetric and density data from extant animals. Although volumetric models appear to more tightly constrain dinosaur body mass, there remains a clear need to further support these models with more exhaustive data from living animals. The relative and absolute discrepancies in mass predictions between volumetric models and scaling equations also indicate a need to systematically compare predictions across a wide size and taxonomic range to better inform studies of dinosaur body size.     

Fatigue creep damage at the cement–bone interface: An experimental and a micro-mechanical finite element study

The goal of this study was to quantify the micromechanics of the cement–bone interface under tensile fatigue loading using finite element analysis (FEA) and to understand the underlying mechanisms that play a role in the fatigue behavior of this interface. Laboratory cement–bone specimens were subjected to a tensile fatigue load, while local displacements and crack growth on the specimen's surface were monitored. FEA models were created from these specimens based upon micro-computed tomography data. To accurately model interfacial gaps at the interface between the bone and cement, a custom-written erosion algorithm was applied to the bone model. A fatigue load was simulated in the FEA models while monitoring the local displacements and crack propagation. The results showed the FEA models were able to capture the general experimental creep damage behavior and creep stages of the interface. Consistent with the experiments, the majority of the deformation took place at the contact interface. Additionally, the FEA models predicted fatigue crack patterns similar to experimental findings. Experimental surface cracks correlated moderately with FEA surface cracks (r²=0.43), but did not correlate with the simulated crack volume fraction (r²=0.06). Although there was no relationship between experimental surface cracks and experimental creep damage displacement (r²=0.07), there was a strong relationship between the FEA crack volume fraction and the FEA creep damage displacement (r²=0.76). This study shows the additional value of FEA of the cement–bone interface relative to experimental studies and can therefore be used to optimize its mechanical properties.     

Predicting trophic relations in ecological networks: A test of the Allometric Diet Breadth Model

Few food web theory hypotheses/predictions can be readily tested using likelihoods of reproducing the data. Simple probabilistic models for food web structure, however, are an exception as their likelihoods were recently derived. Here I test the performance of a more complex model for food web structure that is grounded in the allometric scaling of interactions with body size and the theory of optimal foraging (Allometric Diet Breadth Model—ADBM). This deterministic model has been evaluated by measuring the fraction of trophic relations it correctly predicts. I contrasted this value with that produced by simpler models based on body sizes and found that the quantitative information on allometric scaling and optimal foraging does not significantly increase model fit. Also, I present a method to compute the p-value for the fraction of trophic interactions correctly predicted by the ADBM, or any other model, with respect to three probabilistic models. I find that the ADBM predicts significantly more links than random graphs, but other models can outperform it. Although optimal foraging and allometric scaling may improve our understanding of food webs, the ADBM needs to be modified or replaced to find support in the data.     

On the importance of 3D,geometrically accurate,and subject-specific finite element analysis for evaluation of in-vivo soft tissue loads

Pressure ulcers are a type of local soft tissue injury due to sustained mechanical loading and remain a common issue in patient care. People with spinal cord injury (SCI) are especially at risk of pressure ulcers due to impaired mobility and sensory perception. The development of load improving support structures relies on realistic tissue load evaluation e.g. using finite element analysis (FEA). FEA requires realistic subject-specific mechanical properties and geometries. This study focuses on the effect of geometry. MRI is used for the creation of geometrically accurate models of the human buttock for three able-bodied volunteers and three volunteers with SCI. The effect of geometry on observed internal tissue deformations for each subject is studied by comparing FEA findings for equivalent loading conditions. The large variations found between subjects confirms the importance of subject-specific FEA.     

GrowMatch: An Automated Method for Reconciling In Silico/In Vivo Growth Predictions

Genome-scale metabolic reconstructions are typically validated by comparing in silico growth predictions across different mutants utilizing different carbon sources with in vivo growth data. This comparison results in two types of model-prediction inconsistencies; either the model predicts growth when no growth is observed in the experiment (GNG inconsistencies) or the model predicts no growth when the experiment reveals growth (NGG inconsistencies). Here we propose an optimization-based framework, GrowMatch, to automatically reconcile GNG predictions (by suppressing functionalities in the model) and NGG predictions (by adding functionalities to the model). We use GrowMatch to resolve inconsistencies between the predictions of the latest in silico Escherichia coli (iAF1260) model and the in vivo data available in the Keio collection and improved the consistency of in silico with in vivo predictions from 90.6% to 96.7%. Specifically, we were able to suggest consistency-restoring hypotheses for 56/72 GNG mutants and 13/38 NGG mutants. GrowMatch resolved 18 GNG inconsistencies by suggesting suppressions in the mutant metabolic networks. Fifteen inconsistencies were resolved by suppressing isozymes in the metabolic network, and the remaining 23 GNG mutants corresponding to blocked genes were resolved by suitably modifying the biomass equation of iAF1260. GrowMatch suggested consistency-restoring hypotheses for five NGG mutants by adding functionalities to the model whereas the remaining eight inconsistencies were resolved by pinpointing possible alternate genes that carry out the function of the deleted gene. For many cases, GrowMatch identified fairly nonintuitive model modification hypotheses that would have been difficult to pinpoint through inspection alone. In addition, GrowMatch can be used during the construction phase of new, as opposed to existing, genome-scale metabolic models, leading to more expedient and accurate reconstructions.     

Verification of predicted specimen-specific natural and implanted patellofemoral kinematics during simulated deep knee bend

Verified computational models represent an efficient method for studying the relationship between articular geometry, soft-tissue constraint, and patellofemoral (PF) mechanics. The current study was performed to evaluate an explicit finite element (FE) modeling approach for predicting PF kinematics in the natural and implanted knee. Experimental three-dimensional kinematic data were collected on four healthy cadaver specimens in their natural state and after total knee replacement in the Kansas knee simulator during a simulated deep knee bend activity. Specimen-specific FE models were created from medical images and CAD implant geometry, and included soft-tissue structures representing medial–lateral PF ligaments and the quadriceps tendon. Measured quadriceps loads and prescribed tibiofemoral kinematics were used to predict dynamic kinematics of an isolated PF joint between 10° and 110° femoral flexion. Model sensitivity analyses were performed to determine the effect of rigid or deformable patellar representations and perturbed PF ligament mechanical properties (pre-tension and stiffness) on model predictions and computational efficiency.Predicted PF kinematics from the deformable analyses showed average root mean square (RMS) differences for the natural and implanted states of less than 3.1° and 1.7 mm for all rotations and translations. Kinematic predictions with rigid bodies increased average RMS values slightly to 3.7° and 1.9 mm with a five-fold decrease in computational time. Two-fold increases and decreases in PF ligament initial strain and linear stiffness were found to most adversely affect kinematic predictions for flexion, internal–external tilt and inferior–superior translation in both natural and implanted states. The verified models could be used to further investigate the effects of component alignment or soft-tissue variability on natural and implant PF mechanics.     

Development of a surrogate model based on patient weight,bone mass and geometry to predict femoral neck strains and fracture loads

Osteoporosis and related bone fractures are an increasing global burden in our ageing society. Areal bone mineral density assessed through dual energy X-ray absorptiometry (DEXA), the clinically accepted and most used method, is not sufficient to assess fracture risk individually. Finite element (FE) modelling has shown improvements in prediction of fracture risk, better than aBMD from DEXA, but is not practical for widespread clinical use. The aim of this study was to develop an adaptive neural network (ANN)-based surrogate model to predict femoral neck strains and fracture loads obtained from a previously developed population-based FE model. The surrogate model performance was assessed in simulating two loading conditions: the stance phase of gait and a fall.The surrogate model successfully predicted strains estimated by FE (r² = 0.90–0.98 for level gait load case, r² = 0.92–0.96 for the fall load case). Moreover, an ANN model based on three measurements obtainable in clinics (femoral neck length (level gait) or maximum femoral neck diameter (fall), femoral neck bone mass, body weight) was able to give reasonable predictions (r² = 0.84–0.94) for all of the strain metrics and the estimated femoral neck fracture load. Overall, the surrogate model has potential for clinical applications as they are based on simple measures of geometry and bone mass which can be derived from DEXA images, accurately predicting FE model outcomes, with advantages over FE models as they are quicker and easier to perform.     

Premolar microwear and tooth use in Australopithecus afarensis

The mandibular third premolar (P₃) of Australopithecus afarensis is notable for extensive morphological variability (e.g., metaconid presence/absence, closure of the anterior fovea, root number) and temporal trends in crown length and shape change over its 700 Ka time range. Hominins preceding A. afarensis have unicuspid, mesiodistally elongated P₃s with smaller talonids, and subsequent australopiths have bicuspid, more symmetrically-shaped P₃ crowns with expanded talonids. For these features, A. afarensis is intermediate and, thus, evinces the incipient stages of P₃ molarization. Here, we examine A. afarensis P₃ Phase II microwear and compare it with that of Australopithecus africanus and Cercocebus atys, an extant hard-object specialist, to assess whether the role of the P₃ in food processing changed over time in A. afarensis. Premolar Phase II microwear textures are also compared with those of the molars to look for evidence of functional differentiation along the tooth row (i.e., that foods with different mechanical properties were processed by separate regions of the postcanine battery). Microwear textures were also examined along the mesial protoconid crest, the site of occlusion with the maxillary canine, of the A. afarensis P₃ and compared with the same region in Pan troglodytes to determine whether microwear can be useful for identifying changes in the occlusal relationship between the P₃ and maxillary canine in early Australopithecus. Finally, temporal trends in P₃ Phase II and mesial microwear are considered. Results indicate that 1) both the P₃ and molar Phase II facets of A. afarensis have less complex microwear textures than in A. africanus or C. atys; 2) A. afarensis P₃ and molar Phase II textures differ, though not to the extent seen in taxa that eat hard and tough items; 3) microwear along the A. afarensis mesial protoconid crest is clearly distinct from that of the P. troglodytes, indicating that there is no honing equivalent in A. afarensis; and 4) there is little evidence of change over time in A. afarensis P₃ microwear on either the mesial or Phase II facet. In sum, these results provide no evidence that A. afarensis routinely loaded either its premolars or molars to process hard objects or that A. afarensis P₃ function changed over time.     

Finite element analyses of human vertebral bodies embedded in polymethylmethalcrylate or loaded via the hyperelastic intervertebral disc models provide equivalent predictions of experimental strength

Quantitative computer tomography (QCT)-based finite element (FE) models of vertebral body provide better prediction of vertebral strength than dual energy X-ray absorptiometry. However, most models were validated against compression of vertebral bodies with endplates embedded in polymethylmethalcrylate (PMMA). Yet, loading being as important as bone density, the absence of intervertebral disc (IVD) affects the strength. Accordingly, the aim was to assess the strength predictions of the classic FE models (vertebral body embedded) against the in vitro and in silico strengths of vertebral bodies loaded via IVDs. High resolution peripheral QCT (HR-pQCT) were performed on 13 segments (T11/T12/L1). T11 and L1 were augmented with PMMA and the samples were tested under a 4° wedge compression until failure of T12. Specimen-specific model was generated for each T12 from the HR-pQCT data. Two FE sets were created: FE-PMMA refers to the classical vertebral body embedded model under axial compression; FE-IVD to their loading via hyperelastic IVD model under the wedge compression as conducted experimentally. Results showed that FE-PMMA models overestimated the experimental strength and their strength prediction was satisfactory considering the different experimental set-up. On the other hand, the FE-IVD models did not prove significantly better (Exp/FE-PMMA: R²=0.68; Exp/FE-IVD: R²=0.71, p=0.84). In conclusion, FE-PMMA correlates well with in vitro strength of human vertebral bodies loaded via real IVDs and FE-IVD with hyperelastic IVDs do not significantly improve this correlation. Therefore, it seems not worth adding the IVDs to vertebral body models until fully validated patient-specific IVD models become available.     

Transitional fossils and the origin of turtles

The origin of turtles is one of the most contentious issues in systematics with three currently viable hypotheses: turtles as the extant sister to (i) the crocodile–bird clade, (ii) the lizard–tuatara clade, or (iii) Diapsida (a clade composed of (i) and (ii)). We reanalysed a recent dataset that allied turtles with the lizard–tuatara clade and found that the inclusion of the stem turtle Proganochelys quenstedti and the ‘parareptile’ Eunotosaurus africanus results in a single overriding morphological signal, with turtles outside Diapsida. This result reflects the importance of transitional fossils when long branches separate crown clades, and highlights unexplored issues such as the role of topological congruence when using fossils to calibrate molecular clocks.     

A novel methodology for generating 3D finite element models of the hip from 2D radiographs

Finite element (FE) modelling has been proposed as a tool for estimating fracture risk and patient-specific FE models are commonly based on computed tomography (CT). Here, we present a novel method to automatically create personalised 3D models from standard 2D hip radiographs. A set of geometrical parameters of the femur were determined from seven a–p hip radiographs and compared to the 3D femoral shape obtained from CT as training material; the error in reconstructing the 3D model from the 2D radiographs was assessed. Using the geometry parameters as the input, the 3D shape of another 21 femora was built and meshed, separating a cortical and trabecular compartment. The material properties were derived from the homogeneity index assessed by texture analysis of the radiographs, with focus on the principal tensile and compressive trabecular systems. The ability of these FE models to predict failure load as determined by experimental biomechanical testing was evaluated and compared to the predictive ability of DXA. The average reconstruction error of the 3D models was 1.77 mm (±1.17 mm), with the error being smallest in the femoral head and neck, and greatest in the trochanter. The correlation of the FE predicted failure load with the experimental failure load was r²=64% for the reconstruction FE model, which was significantly better (p<0.05) than that for DXA (r²=24%). This novel method for automatically constructing a patient-specific 3D finite element model from standard 2D radiographs shows encouraging results in estimating patient-specific failure loads.     

Using 3D finite element models verified the importance of callus material and microstructure in biomechanics restoration during bone defect repair

Background: There is lack of further observations on the microstructure and material property of callus during bone defect healing and the relationships between callus properties and the mechanical strength. Methods: Femur bone defect model was created in rabbits and harvested CT data to reconstruct finite element models at 1 and 2 months. Three types of assumed finite element models were compared to study the callus properties, which assumed the material elastic property as heterogeneous (R-model), homogenous (H-model) or did not change from 1 to 2 months (U-model). Results: The apparent elastic moduli increased at 2 months (from 355.58 ± 132.67 to 1139.30 ± 967.43 MPa) in R-models. But there was no significant difference in apparent elastic moduli between R-models (355.58 ± 132.67 and 1139.30 ± 967.43 MPa) and H-models (344.79 ± 138.73 and 1001.52 ± 692.12 MPa) in 1 and 2 months. A significant difference of apparent elastic moduli was found between the R-model (1139.30 ± 967.43 MPa) and U-model group (207.15 ± 64.60 MPa) in 2 months. Conclusions: This study showed that the callus structure stability remodeled overtime to achieve a more effective structure, while the material quality of callus tissue is a very important factor for callus strength. At the meantime, this study showed an evidence that the material heterogeneity maybe not as important as it is in bone fracture model.     

Examining tissue composition,whole-bone morphology and mechanical behavior of GorabPrx1 mice tibiae: A mouse model of premature aging

Gerodermia osteodysplastica (GO) is a segmental progeroid disorder caused by loss-of-function mutations in the GORAB gene, associated with early onset osteoporosis and bone fragility. A conditional mouse model of GO (Gorab^Prx1) was generated in which the Gorab gene was deleted in long bones. We examined the biomechanical/functional relevance of the Gorab^Prx1 mutants as a premature aging model by characterizing bone composition, tissue-level strains, and whole-bone morphology and mechanical properties of the tibia. MicroCT imaging showed that Gorab^Prx1 tibiae had an increased anterior convex curvature and decreased cortical cross-sectional area, cortical thickness and moments of inertia, compared to littermate control (LC) tibiae. Fourier transform infrared (FTIR) imaging indicated a 34% decrease in mineral/matrix ratio and a 27% increase in acid phosphate content in the posterior metaphyseal cortex of the Gorab^Prx1 tibiae (p < .05), suggesting delayed mineralization. In vivo strain gauge measurement and finite element analysis showed ∼two times higher tissue-level strains within the Gorab^Prx1 tibiae relative to LC tibiae when subjected to axial compressive loads of the same magnitude. Three-point bending tests suggested that Gorab^Prx1 tibiae were weaker and more brittle, as indicated by decreasing whole-bone strength (46%), stiffness (55%), work-to-fracture (61%) and post-yield displacement (47%). Many of these morphological and biomechanical characteristics of the Gorab^Prx1 tibia recapitulated changes in other animal models of skeletal aging. Future studies are necessary to confirm how our observations might guide the way to a better understanding and treatment of GO.     

Finite element analysis to characterize how varying patellar loading influences pressure applied to cartilage: model evaluation

A finite element analysis (FEA) modeling technique has been developed to characterize how varying the orientation of the patellar tendon influences the patellofemoral pressure distribution. To evaluate the accuracy of the technique, models were created from MRI images to represent five knees that were previously tested in vitro to determine the influence of hamstrings loading on patellofemoral contact pressures. Hamstrings loading increased the lateral and posterior orientation of the patellar tendon. Each model was loaded at 40°, 60°, and 80° of flexion with quadriceps force vectors representing the experimental loading conditions. The orientation of the patellar tendon was represented for the loaded and unloaded hamstrings conditions based on experimental measures of tibiofemoral alignment. Similar to the experimental data, simulated loading of the hamstrings within the FEA models shifted the center of pressure laterally and increased the maximum lateral pressure. Significant (p < 0.05) differences were identified for the center of pressure and maximum lateral pressure from paired t-tests carried out at the individual flexion angles. The ability to replicate experimental trends indicates that the FEA models can be used for future studies focused on determining how variations in the orientation of the patellar tendon related to anatomical or loading variations or surgical procedures influence the patellofemoral pressure distribution.     

A computational study of invariant I5 in a nearly incompressible transversely isotropic model for white matter

The aligned axonal fiber bundles in white matter make it suitable to be modeled as a transversely isotropic material. Recent experimental studies have shown that a minimal form, nearly incompressible transversely isotropic (MITI) material model, is capable of describing mechanical anisotropy of white matter. Here, we used a finite element (FE) computational approach to demonstrate the significance of the fifth invariant (I₅) when modeling the anisotropic behavior of white matter in the large-strain regime. We first implemented and validated the MITI model in an FE simulation framework for large deformations. Next, we applied the model to a plate-hole structural problem to highlight the significance of the invariant I₅ by comparing with the standard fiber reinforcement (SFR) model. We also compared the two models by fitting the experiment data of asymmetric indentation, shear test, and uniaxial stretch of white matter. Our results demonstrated the significance of I₅ in describing shear deformation/anisotropy, and illustrated the potential of the MITI model to characterize transversely isotropic white matter tissues in the large-strain regime.     

Is the sheep a suitable model to study the mechanical alterations of disc degeneration in humans? A probabilistic finite element model study

Intervertebral disc degeneration is one major source of low back pain, which because of its complex multifactorial nature renders the treatment challenging and thus necessitates extensive research. Experimental animal models have proven valuable in improving our understanding of degenerative processes and potentially promising therapies. Currently, the sheep is the most frequently used large animal in vivo model in intervertebral disc research. However, despite its undoubted value for investigations of the complex biological and cellular aspects, to date, it is unclear whether the sheep is also suited to study the mechanical aspects of disc degeneration in humans.A parametric finite element (FE) model of the L4–5 spinal motion segment was developed. Using this model, the geometry and the material properties of both the human and the ovine spinal segment as well as different appearances of disc degeneration can be depicted. Under pure and combined loads, it was investigated whether degenerative changes to both the human and the ovine model equivalent caused the same mechanical response.Different patterns of degeneration resulted in large variations in the ranges of motion, intradiscal pressure, ligament and facet loads. In the human, but not in the ovine model, all these results differed significantly between different degrees of degeneration.This FE model study highlighted possible differences in the mechanical response to disc degeneration between human and ovine intervertebral discs and indicates the necessity of further, more detailed, investigations.