Laura B. Porro

In vivo bone strain and finite-element modeling of the craniofacial haft in catarrhine primates.

Hypotheses regarding patterns of stress, strain and deformation in the craniofacial skeleton are central to adaptive explanations for the evolution of primate craniofacial form. The complexity of craniofacial skeletal morphology makes it difficult to evaluate these hypotheses with in vivo bone strain data. In this paper, new in vivo bone strain data from the intraorbital surfaces of the supraorbital torus, postorbital bar and postorbital septum, the anterior surface of the postorbital bar, and the anterior root of the zygoma are combined with published data from the supraorbital region and zygomatic arch to evaluate the validity of a finite‐element model (FEM) of a macaque cranium during mastication. The behavior of this model is then used to test hypotheses regarding the overall deformation regime in the craniofacial haft of macaques. This FEM constitutes a hypothesis regarding deformation of the facial skeleton during mastication. A simplified verbal description of the deformation regime in the macaque FEM is as follows. Inferior bending and twisting of the zygomatic arches about a rostrocaudal axis exerts inferolaterally directed tensile forces on the lateral orbital wall, bending the wall and the supraorbital torus in frontal planes and bending and shearing the infraorbital region and anterior zygoma root in frontal planes. Similar deformation regimes also characterize the crania of Homo and Gorilla under in vitro loading conditions and may be shared among extant catarrhines. Relatively high strain magnitudes in the anterior root of the zygoma suggest that the morphology of this region may be important for resisting forces generated during feeding.

Lower limits of ornithischian dinosaur body size inferred from a new Upper Jurassic heterodontosaurid from North America

The extremes of dinosaur body size have long fascinated scientists. The smallest (<1 m length) known dinosaurs are carnivorous saurischian theropods, and similarly diminutive herbivorous or omnivorous ornithischians (the other major group of dinosaurs) are unknown. We report a new ornithischian dinosaur, Fruitadens haagarorum, from the Late Jurassic of western North America that rivals the smallest theropods in size. The largest specimens of Fruitadens represent young adults in their fifth year of development and are estimated at just 65–75 cm in total body length and 0.5–0.75 kg body mass. They are thus the smallest known ornithischians. Fruitadens is a late-surviving member of the basal dinosaur clade Heterodontosauridae, and is the first member of this clade to be described from North America. The craniodental anatomy and diminutive body size of Fruitadens suggest that this taxon was an ecological generalist with an omnivorous diet, thus providing new insights into morphological and palaeoecological diversity within Dinosauria. Late-surviving (Late Jurassic and Early Cretaceous) heterodontosaurids are smaller and less ecologically specialized than Early (Late Triassic and Early Jurassic) heterodontosaurids, and this ecological generalization may account in part for the remarkable 100-million-year-long longevity of the clade.

Free body analysis, beam mechanics, and finite element modeling of the mandible of Alligator mississippiensis

The mechanical behavior of mammalian mandibles is well‐studied, but a comprehensive biomechanical analysis (incorporating detailed muscle architecture, accurate material properties, and three‐dimensional mechanical behavior) of an extant archosaur mandible has never been carried out. This makes it unclear how closely models of extant and extinct archosaur mandibles reflect reality and prevents comparisons of structure–function relationships in mammalian and archosaur mandibles. We tested hypotheses regarding the mechanical behavior of the mandible of Alligator mississippiensis by analyzing reaction forces and bending, shear, and torsional stress regimes in six models of varying complexity. Models included free body analysis using basic lever arm mechanics, 2D and 3D beam models, and three high‐resolution finite element models of the Alligator mandible, incorporating, respectively, isotropic bone without sutures, anisotropic bone with sutures, and anisotropic bone with sutures and contact between the mandible and the pterygoid flange. Compared with the beam models, the Alligator finite element models exhibited less spatial variability in dorsoventral bending and sagittal shear stress, as well as lower peak values for these stresses, suggesting that Alligator mandibular morphology is in part designed to reduce these stresses during biting. However, the Alligator models exhibited greater variability in the distribution of mediolateral and torsional stresses than the beam models. Incorporating anisotropic bone material properties and sutures into the model reduced dorsoventral and torsional stresses within the mandible, but led to elevated mediolateral stresses. These mediolateral stresses were mitigated by the addition of a pterygoid‐mandibular contact, suggesting important contributions from, and trade‐offs between, material properties and external constraints in Alligator mandible design. Our results suggest that beam modeling does not accurately represent the mechanical behavior of the Alligator mandible, including important performance metrics such as magnitude and orientation of reaction forces, and mediolateral bending and torsional stress distributions. J.Morphol. 2011.

The impact of bone and suture material properties on mandibular function in Alligator mississippiensis: testing theoretical phenotypes with finite element analysis.

The functional effects of bone and suture stiffness were considered here using finite element models representing three different theoretical phenotypes of an Alligator mississippiensis mandible. The models were loaded using force estimates derived from muscle architecture in dissected specimens, constrained at the 18th and 19th teeth in the upper jaw and 19th tooth of the lower jaw, as well as at the quadrate‐articular joint. Stiffness was varied systematically in each theoretical phenotype. The three theoretical phenotypes included: (i) linear elastic isotropic bone of varying stiffness and no sutures; (ii) linear elastic orthotropic bone of varying stiffness with no sutures; and (iii) linear elastic isotropic bone of a constant stiffness with varying suture stiffness. Variation in the isotropic material properties of bone primarily resulted in changes in the magnitude of principal strain. By comparison, variation in the orthotropic material properties of bone and isotropic material properties of sutures resulted in: a greater number of bricks becoming either more compressive or more tensile, changing between being either dominantly compressive or tensile, and having larger changes in the orientation of maximum principal strain. These data indicate that variation in these model properties resulted in changes to the strain regime of the model, highlighting the importance of using biologically verified material properties when modeling vertebrate bones. When bones were compared within each set, the response of each to changing material properties varied. In two of the 12 bones in the mandible, varied material properties within sutures resulted in a decrease in the magnitude of principal strain in bricks adjacent to the bone/suture interface and decreases in stored elastic energy. The varied response of the mandibular bones to changes in suture stiffness highlights the importance of defining the appropriate functional unit when addressing relationships of performance and morphology.

In vivo bone strain and finite element modeling of the mandible of Alligator mississippiensis.

Forces experienced during feeding are thought to strongly influence the morphology of the vertebrate mandible; in vivo strain data are the most direct evidence for deformation of the mandible induced by these loading regimes. Although many studies have documented bone strains in the mammalian mandible, no information is available on strain magnitudes, orientations or patterns in the sauropsid lower jaw during feeding. Furthermore, strain gage experiments record the mechanical response of bone at a few locations, not across the entire mandible. In this paper, we present bone strain data recorded at various sites on the lower jaw of Alligator mississippiensis during in vivo feeding experiments. These data are used to understand how changes in loading regime associated with changes in bite location are related to changes in strain regime on the working and balancing sides of the mandible. Our results suggest that the working side mandible is bent dorsoventrally and twisted about its long‐axis during biting, and the balancing side experiences primarily dorsoventral bending. Strain orientations are more variable on the working side than on the balancing side with changes in bite point and between experiments; the balancing side exhibits higher strain magnitudes. In the second part of this paper, we use principal strain orientations and magnitudes recorded in vivo to evaluate a finite element model of the alligator mandible. Our comparison demonstrates that strain orientations and mandibular deformation predicted by the model closely match in vivo results; however, absolute strain magnitudes are lower in the finite element model.

Anatomy and cranial functional morphology of the small-bodied dinosaur Fruitadens haagarorum from the Upper Jurassic of the USA.

Background Heterodontosaurids are an important but enigmatic and poorly understood early radiation of ornithischian dinosaurs. The late-surviving heterodontosaurid Fruitadens haagarorum from the Late Jurassic (early Tithonian) Morrison Formation of the western USA is represented by remains of several small (<1 metre total body length, <1 kg body mass) individuals that include well-preserved but incomplete cranial and postcranial material. Fruitadens is hypothesized to represent one of the smallest known ornithischian dinosaurs. Methodology/Principal Findings We describe the cranial and postcranial anatomy of Fruitadens in detail, providing comparisons to all other known heterodontosaurid taxa. High resolution micro-CT data provides new insights into tooth replacement and the internal anatomy of the tooth-bearing bones. Moreover, we provide a preliminary functional analysis of the skull of late-surviving heterodontosaurids, discuss the implications of Fruitadens for current understanding of heterodontosaurid monophyly, and briefly review the evolution and biogeography of heterodontosaurids. Conclusions/Significance The validity of Fruitadens is supported by multiple unique characters of the dentition and hindlimb as well as a distinct character combination. Fruitadens shares highly distinctive appendicular characters with other heterodontosaurids, strengthening monophyly of the clade on the basis of the postcranium. Mandibular morphology and muscle moment arms suggest that the jaws of late-surviving heterodontosaurids, including Fruitadens, were adapted for rapid biting at large gape angles, contrasting with the jaws of the stratigraphically older Heterodontosaurus, which were better suited for strong jaw adduction at small gapes. The lack of wear facets and plesiomorphic dentition suggest that Fruitadens used orthal jaw movements and employed simple puncture-crushing to process food. In combination with its small body size, these results suggest that Fruitadens was an ecological generalist, consuming select plant material and possibly insects or other invertebrates.

Descriptive Anatomy and Three-Dimensional Reconstruction of the Skull of the Early Tetrapod Acanthostega gunnari Jarvik, 1952

The early tetrapod Acanthostega gunnari is an iconic fossil taxon exhibiting skeletal morphology reflecting the transition of vertebrates from water onto land. Computed tomography data of two Acanthostega skulls was segmented using visualization software to digitally separate bone from matrix and individual bones of the skull from each other. A revised description of cranial and lower jaw anatomy in this taxon based on CT data includes new details of sutural morphology, the previously undescribed quadrate and articular bones, and the mandibular symphysis. Sutural morphology is used to infer loading regime in the skull during feeding, and suggests Acanthostega used its anterior jaws to initially seize prey while smaller posterior teeth were used to restrain struggling prey during ingestion. Novel methods were used to repair and retrodeform the skull, resulting in a three-dimensional digital reconstruction that features a longer postorbital region and more strongly hooked anterior lower jaw than previous attempts while supporting the presence of a midline gap between the nasals and median rostrals.

Inverse dynamic modelling of jumping in the red-legged running frog Kassina maculata

ABSTRACT Although the red-legged running frog, Kassina maculata, is secondarily a walker/runner, it retains the capacity for multiple locomotor modes, including jumping at a wide range of angles (nearly 70 deg). Using simultaneous hind limb kinematics and single-foot ground reaction forces, we performed inverse dynamics analyses to calculate moment arms and torques about the hind limb joints during jumping at different angles in K. maculata. We show that forward thrust is generated primarily at the hip and ankle, while body elevation is primarily driven by the ankle. Steeper jumps are achieved by increased thrust at the hip and ankle and greater downward rotation of the distal limb segments. Because of its proximity to the GRF vector, knee posture appears to be important in controlling torque directions about this joint and, potentially, torque magnitudes at more distal joints. Other factors correlated with higher jump angles include increased body angle in the preparatory phase, faster joint openings and increased joint excursion, higher ventrally directed force, and greater acceleration and velocity. Finally, we demonstrate that jumping performance in K. maculata does not appear to be compromised by presumed adaptation to walking/running. Our results provide new insights into how frogs engage in a wide range of locomotor behaviours and the multi-functionality of anuran limbs. Summary: Experimental data and inverse dynamic modelling demonstrate how forward thrust and elevation are produced in the frog hind limb, allowing frogs to jump at a wide range of angles.

Digital dissection of the model organism Xenopus laevis using contrast‐enhanced computed tomography

The African clawed frog, Xenopus laevis, is one of the most widely used model organisms in biological research. However, the most recent anatomical description of X. laevis was produced nearly a century ago. Compared with other anurans, pipid frogs – including X. laevis – exhibit numerous unusual morphological features; thus, anatomical descriptions of more ‘typical’ frogs do not detail many aspects of X. laevis skeletal and soft‐tissue morphology. The relatively new method of using iodine‐based agents to stain soft tissues prior to high‐resolution X‐ray imaging has several advantages over gross dissection, such as enabling dissection of very small and fragile specimens, and preserving the three‐dimensional topology of anatomical structures. Here, we use contrast‐enhanced computed tomography to produce a high‐resolution three‐dimensional digital dissection of a post‐metamorphic X. laevis to successfully visualize: skeletal and muscular anatomy; the nervous, respiratory, digestive, excretory and reproductive systems; and the major sense organs. Our digital dissection updates and supplements previous anatomical descriptions of this key model organism, and we present the three‐dimensional data as interactive portable document format (PDF) files that are easily accessible and freely available for research and educational purposes. The data presented here hold enormous potential for applications beyond descriptive purposes, particularly for biological researchers using this taxon as a model organism, comparative anatomy and biomechanical modelling.

Digital preparation and osteology of the skull of Lesothosaurus diagnosticus (Ornithischia: Dinosauria)

Several skulls of the ornithischian dinosaur Lesothosaurus diagnosticus (Lower Jurassic, southern Africa) are known, but all are either incomplete, deformed, or incompletely prepared. This has hampered attempts to provide a comprehensive description of skull osteology in this crucial early dinosaurian taxon. Using visualization software, computed tomographic scans of the Lesothosaurus syntypes were digitally segmented to remove matrix, and identify and separate individual cranial and mandibular bones, revealing new anatomical details such as sutural morphology and the presence of several previously undescribed elements. Together with visual inspection of exposed skull bones, these CT data enable a complete description of skull anatomy in this taxon. Comparisons with our new data suggest that two specimens previously identified as Lesothosaurus sp. (MNHN LES 17 and MNHN LES 18) probably represent additional individuals of Lesothosaurus diagnosticus.