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David S Strait

Publications and source records attributed to David S Strait.

7 recordsLinked to original sources

A comparison of cortical elastic properties in the craniofacial skeletons of three primate species and its relevance to the study of human evolution.

When a force is applied to an object, the resulting pattern of strain is a function of both the object's geometry and its elastic properties. Thus, knowledge of elastic properties in craniofacial cortical bone is indispensable for exploring the biomechanics and adaptation of primate skulls. However, elastic properties, such as density and stiffness, cannot be measured in all species, particularly extinct species known only from fossils. In order for advanced engineering techniques such as finite element analysis (FEA) to be applied to questions of primate and hominid craniofacial functional morphology, it is important to understand interspecific patterns of variation in elastic properties. We hypothesized that closely related species would have similar patterns of bone elastic properties, and that similarities with extant species should allow reasonable predictions of elastic properties in the skeletons of extinct primate species. In this study, we tested this hypothesis by measuring elastic properties in five areas of the external cortex of the baboon craniofacial skeleton using an ultrasonic technique, and by comparing the results to existing data from macaque and human crania. Results showed that cortical density, thickness, elastic and shear moduli, and anisotropy varied among areas in the baboon cranium. Similar variation had previously been found in rhesus and human crania, suggesting area-specific elastic patterns in the skulls of each species. Comparison among species showed differences, suggesting species-specific patterns. These patterns were more similar between macaques and baboons for density, maximum elastic and shear stiffness, and anisotropy than between either of these and humans. This finding demonstrates that patterns of cortical elastic properties are generally similar in closely related primate species with similar craniofacial morphology. Thus, reasonable estimates of cortical bone elastic properties should be possible for extinct species through the study of phylogenetically related and functionally similar modern forms. For example, reasonable elastic property estimates of cortical bone from fossil hominid skulls should be possible once adequate information about such properties in extant great apes is added to our current data from humans, macaques, and baboons. Such data should eventually allow FEA of craniofacial function in fossil hominids.

Animals↗

Examining affinities of the Taung child by developmental simulation.

As a well-preserved juvenile and the type specimen of Australopithecus africanus, the Taung child figures prominently in taxonomic, ontogenetic, and phylogenetic analyses of fossil hominins. Despite general agreement about allocation of Sterkfontein and Makapansgat fossils to this species, limited morphological comparisons have been possible between these adult specimens and the juvenile Taung. Here, we used developmental simulation to estimate the adult form of the Taung child, and directly compare its morphology to that of other fossil hominins. Specimens were represented by 50 three-dimensional landmarks superimposed by generalized Procrustes analysis. The simulation process applied developmental trajectories from extant hominine species to the Taung fossil in order to generate its adult form. Despite differences found in the developmental patterns of these modern species, simulations tested on extant juveniles-transforming them into "adults" using trajectories from other species-revealed that these differences have negligible impact on adult morphology. This indicates that morphology already present by occlusion of the first permanent molar is the primary determinant of adult form, thereby supporting use of extant trajectories to estimate the morphology of an extinct species. The simulated Taung adult was then compared to other adult fossils. As these comparisons required assumptions about the pattern and magnitude of developmental change, additional analyses were performed to evaluate these two parameters separately. Results of all analyses overwhelmingly rejected the possibility that the Taung child was a juvenile robust australopith, but were consistent with the hypothesis that the Taung and Sterkfontein fossils are conspecific. Between Sts 5 and Sts 71, the latter is more likely to resemble the adult form of the Taung child.

Animals↗

Fusion patterns of craniofacial sutures in rhesus monkey skulls of known age and sex from Cayo Santiago.

Bones of the face and cranial vault meet at sutural boundaries. These sutures are of great importance for craniofacial growth. Although the effects that the sutures have on modulating craniofacial strains have been investigated, how sutural fusion influences primate craniofacial biomechanics and adaptation are less considered. Confounding this problem is the lack of any systematic data on patterns of craniofacial sutural fusion from animals of known age and sex. This study examined the status of 28 sutures in Macaca mulatta skulls from a collection of animals of known age and sex from Cayo Santiago, Puerto Rico. Survival analysis showed that most animals died before all sutures fused. There was high variation in the age at which individual sutures or sutural sections were fused in M. mulatta, and significant differences in the amount of sutural fusion among regions and between males and females. Intensive fusion of sutures took place between ages 5 and 15. Sutures in the facial area tended to be less fused than in the cranial vault. Between adolescence and adulthood, males tended to have more sutural fusion than females, especially in the facial area. These differences might be biomechanical adaptations during ontogeny to craniofacial sexual dimorphism. These findings enrich our understanding of variation in sutural morphology in rhesus monkeys. Comparative information across primate species is essential for understanding the biomechanics of craniofacial form throughout primate evolution.

Adaptation, Physiological↗

Finite element analysis in functional morphology.

This article reviews the fundamental principles of the finite element method and the three basic steps (model creation, solution, and validation and interpretation) involved in using it to examine structural mechanics. Validation is a critical step in the analysis, without which researchers cannot evaluate the extent to which the model represents or is relevant to the real biological condition. We discuss the method's considerable potential as a tool to test biomechanical hypotheses, and major hurdles involved in doing so reliably, from the perspective of researchers interested in functional morphology and paleontology. We conclude with a case study to illustrate how researchers deal with many of the factors and assumptions involved in finite element analysis.

Anatomy↗

Modeling masticatory muscle force in finite element analysis: sensitivity analysis using principal coordinates analysis.

Our work on a finite element model of the skull of Macaca aims to investigate the functional significance of specific features of primate skulls and to determine to which of the input variables (elastic properties, muscle forces) the model behavior is most sensitive. Estimates of muscle forces acting on the model are derived from estimates of physiological cross-sectional areas (PCSAs) of the jaw muscles scaled by relative electromyographic (EMG) amplitudes recorded in vivo. In this study, the behavior of the model was measured under different assumptions regarding the PCSAs of the jaw muscles and the latency between EMG activity in those muscles and the resulting force production. Thirty-six different loading regimes were applied to the model using four different PCSA sets and nine different PCSA scaling parameters. The four PCSA sets were derived from three different macaque species and one genus average, and the scaling parameters were either EMGs from 10, 20, 30, 40, 50 and 60 msec prior to peak bite force, or simply 100%, 50%, or 25% of peak muscle force. Principal coordinates analysis was used to compare the deformations of the model produced by the 36 loading regimes. Strain data from selected sites on the model were also compared with in vivo bone strain data. The results revealed that when varying the external muscle forces within these boundaries, the majority of the variation in model behavior is attributable to variation in the overall magnitude rather than the relative amount of muscle force generated by each muscle. Once this magnitude-related variation in model deformation was accounted for, significant variation was attributable to differences in relative muscle recruitment between working and balancing sides. Strain orientations at selected sites showed little variation across loading experiments compared with variation documented in vivo. These data suggest that in order to create an accurate and valid finite element model of the behavior of the primate skull at a particular instant during feeding, it is important to include estimates of the relative recruitment levels of the masticatory muscles. However, a lot can be learned about patterns of skull deformation, in fossil species for example, by applying external forces proportional to the estimated relative PCSAs of the jaw adductors.

Animals↗

Modeling elastic properties in finite-element analysis: how much precision is needed to produce an accurate model?

The influence of elastic properties on finite-element analysis was investigated using a finite-element model of a Macaca fascicularis skull. Four finite-element analyses were performed in which the model was assigned different sets of elastic properties. In analysis 1, elastic properties were modeled isotropically using published data obtained from human limb bones. Analyses 2-4 used data obtained from skulls of a closely allied species, M. mulatta, but varied as to how those data were incorporated into the model. In analysis 2, the model was assigned a single set of isotropic elastic properties. In analysis 3, each region within the model was assigned its own set of isotropic elastic properties. Finally, in analysis 4, each region received its own set of orthotropic elastic properties. Although a qualitative assessment indicates that the locations of strain concentrations across the model are broadly similar in all analyses, a quantitative assessment of strain indicates some differences between the analyses. When strain data from the finite-element analyses were compared to strain data derived from in vivo experiments, it was found that the model deformed most realistically using the orthotropic elastic properties employed in analysis 4. Results suggest that finite-element analyses can be adversely affected when elastic properties are modeled imprecisely, and that modelers should attempt to obtain elastic properties data about the species and skeletal elements that are the subjects of their analyses.

Anatomy↗

Inferring hominoid and early hominid phylogeny using craniodental characters: the role of fossil taxa.

Recent discoveries of new fossil hominid species have been accompanied by several phylogenetic hypotheses. All of these hypotheses are based on a consideration of hominid craniodental morphology. However, Collard and Wood (2000) suggested that cladograms derived from craniodental data are inconsistent with the prevailing hypothesis of ape phylogeny based on molecular data. The implication of their study is that craniodental characters are unreliable indicators of phylogeny in hominoids and fossil hominids but, notably, their analysis did not include extinct species. We report here on a cladistic analysis designed to test whether the inclusion of fossil taxa affects the ability of morphological characters to recover the molecular ape phylogeny. In the process of doing so, the study tests both Collard and Wood's (2000) hypothesis of character reliability, and the several recently proposed hypotheses of early hominid phylogeny. One hundred and ninety-eight craniodental characters were examined, including 109 traits that traditionally have been of interest in prior studies of hominoid and early hominid phylogeny, and 89 craniometric traits that represent size-corrected linear dimensions measured between standard cranial landmarks. The characters were partitioned into two data sets. One set contained all of the characters, and the other omitted the craniometric characters. Six parsimony analyses were performed; each data set was analyzed three times, once using an ingroup that consisted only of extant hominoids, a second time using an ingroup of extant hominoids and extinct early hominids, and a third time excluding Kenyanthropus platyops. Results suggest that the inclusion of fossil taxa can play a significant role in phylogenetic analysis. Analyses that examined only extant taxa produced most parsimonious cladograms that were inconsistent with the ape molecular tree. In contrast, analyses that included fossil hominids were consistent with that tree. This consistency refutes the basis for the hypothesis that craniodental characters are unreliable for reconstructing phylogenetic relationships. Regarding early hominids, the relationships of Sahelanthropus tchadensis and Ardipithecus ramidus were relatively unstable. However, there is tentative support for the hypotheses that S. tchadensis is the sister taxon of all other hominids. There is support for the hypothesis that A. anamensis is the sister taxon of all hominids except S. tchadensis and Ar. ramidus. There is no compelling support for the hypothesis that Kenyanthropus platyops shares especially close affinities with Homo rudolfensis. Rather, K. platyops is nested within the Homo + Paranthropus + Australopithecus africanus clade. If K. platyops is a valid species, these relationships suggest that Homo and Paranthropus are likely to have diverged from other hominids much earlier than previously supposed. There is no support for the hypothesis that A. garhi is either the sister taxon or direct ancestor of the genus Homo. Phylogenetic relationships indicate that Australopithecus is paraphyletic. Thus, A. anamensis and A. garhi should be allocated to new genera.

Animals↗