Linking form and function of the fibrous joints in the skull: a new quantification scheme for cranial sutures using the extant fish Polypterus endlicherii

The aim of this study is to connect specific sutural morphologies with the specific types of deformation they experience. To meet this goal, we quantified the morphologies of the interfrontal (IF), interparietal (IP), and frontoparietal (FP) sutures in the extant fish Polypterus endlicherii, and used our published measurements of in vivo deformation of these sutures during feeding to infer how suture morphology and function are connected. Specifically, we found that three relatively simple measures of cross-sectional suture complexity (i.e., the ratio of total sutural length to its shortest end-to-end length; amount of sutural overlap; and size of the largest interdigitation) can be used to distinguish between the IF, FP, and IP sutures, which exhibit very different cross-sectional shapes and responses to loading. Interestingly, these differences in cross-sectional morphology are not reflected by the linear traces of these sutures on the surface of the skull, implying that cross-sectional shape of a suture must be known to infer the loading conditions it experiences. Plotting the three cross-sectional metrics against one another to yield a sutural morphospace shows that the IF, IP, and FP sutures define regions that are largely distinct from one another. Our previous measurements of strain across these sutures suggested that the FP region would lie between the IF and IP regions; instead, the FP region is largely set apart from the other two fields. Based on this discovery, and on the locations of cranial muscles, we propose a new model of deformation in the skull of P. endlicherii during feeding, in which rotation parallel to the skull roof is combined with bending, subjecting the FP suture to complex shearing. Finally, although the sutures of P. endlicherii appear to be significantly less complex than those of mammals, these fish sutures show a similar range of morphologies and perform similar functions as do mammalian sutures.     

Fractal analysis of the Orce skull sutures

Methods of fractal geometry (Mandelbrot, 1983) are used here to analyse the relative complexity of the sagittal and lambdoid sutures visible in the skull fragment formed by parts of an occipital squame and parietals found in a sealed deposit at the early Lower Pleistocene site of Venta Micena (Orce, Granada, Spain), generally regarded as human bone but occasionally suggested as belonging to an equid. For comparison with the fossil, corresponding sutures of various primates (hominids, pongids and cercopithecids) and two other groups of mammals (equids and ruminants) were analysed using the computer program FRACTAL-D (Slice, 1989) in order to determine their fractal dimensions as a measure of differential sutural design complexity. The results show that the fractal dimension of the Venta Micena skull sutures lies within the range of variation for infant specimens of both modern and Plio-Pleistocene hominids. Sutural complexity in young pongids and cercopithecids overlaps the range of fractal dimensions found in hominids, whereas values obtained from equids and ruminants are significantly greater than those for all the primates analysed here. Therefore, in terms of fractal dimension measures of relative complexity, the sutures preserved in the Venta Micena fossil could not have belonged to an equid (pace Agusti & Moyà-Sola, 1987); rather, its fractal dimension is consistent with the attribution of the fossil to an infant of Homo sp.     

Linkage mechanisms in the vertebrate skull: Structure and function of three‐dimensional,parallel transmission systems

Many musculoskeletal systems, including the skulls of birds, fishes, and some lizards consist of interconnected chains of mobile skeletal elements, analogous to linkage mechanisms used in engineering. Biomechanical studies have applied linkage models to a diversity of musculoskeletal systems, with previous applications primarily focusing on two‐dimensional linkage geometries, bilaterally symmetrical pairs of planar linkages, or single four‐bar linkages. Here, we present new, three‐dimensional (3D), parallel linkage models of the skulls of birds and fishes and use these models (available as free kinematic simulation software), to investigate structure–function relationships in these systems. This new computational framework provides an accessible and integrated workflow for exploring the evolution of structure and function in complex musculoskeletal systems. Linkage simulations show that kinematic transmission, although a suitable functional metric for linkages with single rotating input and output links, can give misleading results when applied to linkages with substantial translational components or multiple output links. To take into account both linear and rotational displacement we define force mechanical advantage for a linkage (analogous to lever mechanical advantage) and apply this metric to measure transmission efficiency in the bird cranial mechanism. For linkages with multiple, expanding output points we propose a new functional metric, expansion advantage, to measure expansion amplification and apply this metric to the buccal expansion mechanism in fishes. Using the bird cranial linkage model, we quantify the inaccuracies that result from simplifying a 3D geometry into two dimensions. We also show that by combining single‐chain linkages into parallel linkages, more links can be simulated while decreasing or maintaining the same number of input parameters. This generalized framework for linkage simulation and analysis can accommodate linkages of differing geometries and configurations, enabling novel interpretations of the mechanics of force transmission across a diversity of vertebrate feeding mechanisms and enhancing our understanding of musculoskeletal function and evolution. J. Morphol. 277:1570–1583, 2016. © 2016 Wiley Periodicals, Inc.     

Rigid-body analysis of a lizard skull: modelling the skull of Uromastyx hardwickii

Lizard skulls vary greatly in their detailed morphology. Theoretical models and practical studies have posited a definite relationship between skull morphology and bite performance, but this can be difficult to demonstrate in vivo. Computer modelling provides an alternative approach, as long as hard and soft tissue components can be integrated and the model can be validated. An anatomically accurate three-dimensional computer model of an Uromastyx hardwickii skull was developed for rigid-body dynamic analysis. The Uromastyx jaw was first opened under motion control, and then muscle forces were applied to produce biting simulations where bite forces and joint forces were calculated. Bite forces comparable to those reported in the literature were predicted, and detailed muscular force information was produced along with additional information on the stabilizing role of temporal ligaments in late jaw closing.     

The skull of the Upper Cretaceous snake Dinilysia patagonica Smith‐Woodward, 1901, and its phylogenetic position revisited

The cranial anatomy of Dinilysia patagonica, a terrestrial snake from the Upper Cretaceous of Argentina, is redescribed and illustrated, based on high‐resolution X‐ray computed tomography and better preparations made on previously known specimens, including the holotype. Previously unreported characters reinforce the intriguing mosaic nature of the skull of Dinilysia, with a suite of plesiomorphic and apomorphic characters with respect to extant snakes. Newly recognized plesiomorphies are the absence of the medial vertical flange of the nasal, lateral position of the prefrontal, lizard‐like contact between vomer and palatine, floor of the recessus scalae tympani formed by the basioccipital, posterolateral corners of the basisphenoid strongly ventrolaterally projected, and absence of a medial parietal pillar separating the telencephalon and mesencephalon, amongst others. We also reinterpreted the structures forming the otic region of Dinilysia, confirming the presence of a crista circumfenestralis, which represents an important derived ophidian synapomorphy. Both plesiomorphic and apomorphic traits of Dinilysia are treated in detail and illustrated accordingly. Results of a phylogenetic analysis support a basal position of Dinilysia, as the sister‐taxon to all extant snakes. The fossil taxa Yurlunggur, Haasiophis, Eupodophis, Pachyrhachis, and Wonambi appear as derived snakes nested within the extant clade Alethinophidia, as stem‐taxa to the crown‐clade Macrostomata. The hypothesis of a sister‐group relationship between Dinilysia and Najash rionegrina, as suggested by some authors, is rejected by the results of our analysis. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 194–238.     

A new psittacosaur from Inner Mongolia and the parrot-like structure and function of the psittacosaur skull

We describe a new species of psittacosaur, Psittacosaurus gobiensis, from the Lower Cretaceous of Inner Mongolia and outline a hypothesis of chewing function in psittacosaurs that in many respects parallels that in psittaciform birds. Cranial features that accommodate increased bite force in psittacosaurs include an akinetic skull (both cranium and lower jaws) and differentiation of adductor muscle attachments comparable to that in psittaciform birds. These and other features, along with the presence of numerous large gastroliths, suggest that psittacosaurs may have had a high-fibre, nucivorous (nut-eating) diet.Psittacosaurs, alone among ornithischians, generate oblique wear facets from tooth-to-tooth occlusion without kinesis in either the upper or lower jaws. This is accomplished with a novel isognathous jaw mechanism that combines aspects of arcilineal (vertical) and propalinal (horizontal) jaw movement. Here termed clinolineal (inclined) jaw movement, the mechanism uses posteriorly divergent tooth rows, rather than kinesis, to gain the added width for oblique occlusion. As the lower tooth rows are drawn posterodorsally into occlusion, the increasing width between the upper tooth rows accommodates oblique shear. With this jaw mechanism, psittacosaurs were able to maintain oblique shearing occlusion in an akinetic skull designed to resist high bite forces.     

Biomechanics of the rostrum and the role of facial sutures

The rostrum is a large diameter, thin-walled tubular structure that receives loads from the teeth. The rostrum can be conceptualized both as a rigid structure and as an assemblage of several bones that interface at sutures. Using miniature pigs, we measured in vivo strains in rostral bones and sutures to gain a better understanding of how the rostrum behaves biomechanically. Strains in the premaxillary and nasal bones were low but the adjacent maxillary-premaxillary, internasal, and intermaxillary suture strains were larger by an order of magnitude. While this finding emphasizes the composite nature of the rostrum, we also found evidence in the maxillary and nasal bones for rigid structural behavior. Namely, maxillary strain is consistent with a short beam model under shear deformation from molar loading. Strain in the nasal bones is only partially supported by a long beam model; rather, a complex pattern of dorsal bending of the rostrum from incisor contact and lateral compression is suggested. Torsion of the maxilla is ruled out due to the bilateral occlusion of pigs and the similar working and balancing side strains, although it may be important in mammals with a unilateral bite. Torsional loading does appear important in the premaxillae, which demonstrate working and balancing side changes in strain orientation. These differences are attributed to asymmetrical incisor contact occurring at the end of the power stroke.     

Limb development in the gekkonid lizard gonatodes albogularis: A reconsideration of homology in the lizard carpus and tarsus

Despite the attention squamate lizards have received in the study of digit and limb loss, little is known about limb morphogenesis in pentadactyl lizards. Recent developmental studies have provided a basis for understanding lizard autopodial element homology based on developmental and comparative anatomy. In addition, the composition and identity of some carpal and tarsal elements of lizard limbs, and reptiles in general, have been the theme of discussions about their homology compared to non‐squamate Lepidosauromorpha and basal Amniota. The study of additional embryonic material from different lizard families may improve our understanding of squamate limb evolution. Here, we analyze limb morphogenesis in the gekkonid lizard Gonatodes albogularis describing patterns of chondrogenesis and ossification from early stages of embryonic development to hatchlings. Our results are in general agreement with previous developmental studies, but we also show that limb development in squamates probably involves more chondrogenic elements for carpal and tarsal morphogenesis, as previously recognized on the grounds of comparative anatomy. We provide evidence for the transitory presence of distal carpale 1 and intermedium in the carpus and tibiale, intermedium, distal centralia, and distal tarsale 2 in the tarsus. Hence, we demonstrate that some elements that were believed to be lost in squamate evolution are conserved as transitory elements during limb development. However, these elements do not represent just phylogenetic burden but may be important for the morphogenesis of the lizard autopodium. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

BMP9 induces osteogenesis and adipogenesis in the immortalized human cranial suture progenitors from the patent sutures of craniosynostosis patients

The cranial suture complex is a heterogeneous tissue consisting of osteogenic progenitor cells and mesenchymal stem cells (MSCs) from bone marrow and suture mesenchyme. The fusion of cranial sutures is a highly coordinated and tightly regulated process during development. Craniosynostosis is a congenital malformation caused by premature fusion of cranial sutures. While the progenitor cells derived from the cranial suture complex should prove valuable for studying the molecular mechanisms underlying suture development and pathogenic premature suture fusion, primary human cranial suture progenitors (SuPs) have limited life span and gradually lose osteoblastic ability over passages. To overcome technical challenges in maintaining sufficient and long‐term culture of SuPs for suture biology studies, we establish and characterize the reversibly immortalized human cranial suture progenitors (iSuPs). Using a reversible immortalization system expressing SV40 T flanked with FRT sites, we demonstrate that primary human suture progenitor cells derived from the patent sutures of craniosynostosis patients can be efficiently immortalized. The iSuPs maintain long‐term proliferative activity, express most of the consensus MSC markers and can differentiate into osteogenic and adipogenic lineages upon BMP9 stimulation in vitro and in vivo. The removal of SV40 T antigen by FLP recombinase results in a decrease in cell proliferation and an increase in the endogenous osteogenic and adipogenic capability in the iSuPs. Therefore, the iSuPs should be a valuable resource to study suture development, intramembranous ossification and the pathogenesis of craniosynostosis, as well as to explore cranial bone tissue engineering.     

Brief communication: Ectocranial suture closure in Pongo: Pattern and phylogeny

Ectocranial suture fusion patterns have been shown to contain biological and phylogenetic information. Previously the patterns of Homo, Pan, and Gorilla have been described. These data reflect the phylogenetic relationships among these species. In this study, we applied similar methodology to Pongo to determine the suture synostosis progression of this genus, and to allow comparison to previously reported data on other large‐bodied hominoids. We hypothesized these data would strengthen the argument that suture synostosis patterns reflect the phylogeny of primate taxa. Results indicate that the synostosis of vault sutures in Pongo is similar to that reported for Gorilla (excluding Pan and Homo). However, the lateral‐anterior pattern of fusion, in which there is a strong superior to inferior pattern, for Pongo is unique among these species, reflecting its phylogenetic distinctness among great ape taxa. Am J Phys Anthropol, 2010. © 2010 Wiley‐Liss, Inc.     

Anatomical study of the skull of amphisbaenian Diplometopon zarudnyi (Squamata,Amphisbaenia)

This study investigates the amphisbaenian species skull which includes cranium, lower jaw and hyoid apparatus. The medial dorsal bones comprise the premaxilla, nasal, frontal and parietal. The premaxilla carries a large medial tooth and two lateral ones. The nasals are paired bones and separated by longitudinal suture. Bones of circumorbital series are frontal, orbitosphenoid and maxilla. The occipital ring consists of basioccipital, supraoccipital and exooccipital. Supraoccipital and basioccipital are single bones while the exo-occipitals are paired. The bones of the palate comprise premaxilla, maxilla, septomaxilla, palatine, pterygoid, ectopterygoid, basisphenoid, parasphenoid, orbitosphenoid and laterosphenoid. Prevomer and pterygoid teeth are absent. Palatine represent by two separate bones. The temporal bones are clearly visible. The lower jaw consists of the dentary, articular, coronoid, supra-angular, angular and splenial. The hyoid apparatus is represented by a Y-shaped structure. The mandible is long and is suspended from the braincase via relatively short quadrate. There is an extensive contact between the long angular and the large triangular coronoid. Thus inter-mandibular joint is bridged completely by the angular and consequently, the lower jaws are relatively rigid and kinetic. The maxillae are suspended from the braincase largely by ligaments and muscles rather than through bony articulation. In conclusion, the skull shape affects feeding strategy in Diplometopon zarudnyi. The prey is ingested and transported via a rapid maxillary raking mechanism.     

Computational structural modelling of coronary stent deployment: a review

The finite element (FE) method is a powerful investigative tool in the field of biomedical engineering, particularly in the analysis of medical devices such as coronary stents whose performance is extremely difficult to evaluate in vivo. In recent years, a number of FE studies have been carried out to simulate the deployment of coronary stents, and the results of these studies have been utilised to assess and optimise the performance of these devices. The aim of this paper is to provide a thorough review of the state-of-the-art research in this area, discussing the aims, methods and conclusions drawn from a number of significant studies. It is intended that this paper will provide a valuable reference for future research in this area.     

Diversity and seasonal dynamic of a lizard assemblage in a Neotropical semiarid habitat

We assessed the lizard assemblage from a priority conservation Caatinga area from northeastern Brazil, through a pluriannual ecological approach, to expand the understanding on biodiversity patterns of Neotropical semiarid habitats. The studied area presented one of the richest lizard faunas among Caatinga sites, being composed primarily by species typical from open landscapes. The local species composition was more similar to assemblages from adjacent Caatinga ecoregions than to those from other areas within the same ecoregion. The inventoried lizard assemblage consisted of a few common species and a majority of low abundance ones, and its overall richness and abundance did not differ between rainy and dry months. Our findings demonstrated that the composition of lizard assemblages did not match with the current proposed Caatinga ecoregions, and revealed that the studied assemblage followed a lognormal species-abundance distribution, showing no significant seasonal fluctuation in richness and abundance.     

Comparative morphology and functional interpretation of the sutures in the dermal skull roof of temnospondyl amphibians

Sutures in the dermal skull roof of several Palaeozoic temnospondyl amphibians were studied, including Archegosaurus decheni, Sclerocephalus haeuseri, Cheliderpeton latirostre (Archegosauridae), Acanthostomatops vorax (Zatrachydidae), Onchiodon labyrinthicus (Eryopidae), Micromelerpeton credneri and Branchierpeton amblystomum (Dissorophoidea). Lamellae, flat bevels, butt joints, steep walls, and grooves are the sutural types occurring. Morphological sutural differences cannot be used as taxonomically relevant characters, as they mainly differ in their functional manifestations. Similarities of sutural patterns in taxa not closely related are most probably based on convergence, therefore sutural morphology allows functional conclusions rather than revealing phylogenetic relationships. The basic sutural pattern is described in Micromelerpeton credneri. Different textures and obliquity patterns of the sutural surfaces are characteristic of each region of the skull roof. In all temnospondyls studied, the median plane and the margin of the skull roof show a nearly uniform sutural morphology; the circumorbital region is most complex. Although the skulls studied are akinetic, mesokinetic movements of dermal elements along their sutures must have been permitted to guarantee the dynamic stability of the skull construction.