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Biomedical subjects

Brian G Richmond

Publications and source records attributed to Brian G Richmond.

6 recordsLinked to original sources

Limb-size proportions in Australopithecus afarensis and Australopithecus africanus.

Previous analyses have suggested that Australopithecus africanus possessed more apelike limb proportions than Australopithecus afarensis. However, due to the errors involved in estimating limb length and body size, support for this conclusion has been limited. In this study, we use a new Monte Carlo method to (1) test the hypothesis that A. africanus had greater upper:lower limb-size proportions than A. afarensis and (2) assess the statistical significance of interspecific differences among these taxa, extant apes, and humans. Our Monte Carlo method imposes sampling constraints that reduce extant ape and human postcranial measurements to sample sizes comparable to the fossil samples. Next, composite ratios of fore- and hindlimb geometric means are calculated for resampled measurements from the fossils and comparative taxa. Mean composite ratios are statistically indistinguishable (alpha=0.05) from the actual ratios of extant individuals, indicating that this method conserves each sample's central tendency. When applied to the fossil samples, upper:lower limb-size proportions in A. afarensis are similar to those of humans (p=0.878) and are significantly different from all great ape proportions (p< or =0.034), while Australopithecus africanus is more similar to the apes (p> or =0.180) and significantly different from humans and A. afarensis (p< or =0.031). These results strongly support the hypothesis that A. africanus possessed more apelike limb-size proportions than A. afarensis, suggesting that A. africanus either evolved from a more postcranially primitive ancestor than A. afarensis or that the more apelike limb-size proportions of A. africanus were secondarily derived from an A. afarensis-like ancestor. Among the extant taxa, limb-size proportions correspond with observed levels of forelimb- and hindlimb-dominated positional behaviors. In conjunction with detailed anatomical features linked to arboreality, these results suggest that arboreal posture and locomotion may have been more important components of the A. africanus behavioral repertoire relative to that of A. afarensis.

Animals↗

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↗

Cross-sectional geometry of the human forefoot.

The human forefoot presents an interesting biomechanical problem of clinical importance because loads are distributed unequally across multiple bones. The fact that long bones typically have a several-fold safety factor relative to peak loads suggests that metatarsal strengths should be related to their peak loads. This study is the first to systematically examine the cross-sectional geometric properties of the human forefoot and their relationship to external loads during walking and running. We report midlength cross-sectional geometric properties (CA, Ix, Iy, Imax, Imin, J, Zx, and Zy) of metatarsals (1-5) and the hallucial proximal phalanx of a shod industrial population (n = 40) obtained using computed tomography. We then examine the relationship between these measures of shaft strength and published plantar pressure data sets recorded during the following functional activities: standing, at the push-off stage of the walking cycle, the full walking cycle, and running. Cross-sectional geometric properties of the first ray are greater than those of other rays, even when scaled to bone length. This pattern corresponds to the high pressures recorded for the first ray during most activities. The relationships between cross-sectional geometric properties of the lateral metatarsals and peak plantar pressure data are more complex. Metatarsals 2-4 are weakest in most cross-sectional geometric properties. However, metatarsal 2, and to a lesser extent metatarsal 3, experience relatively high peak pressures. On average, geometric measures of axial and bending strengths (adjusted relative to body size) are lower in females than males, and in European Americans than in African Americans, which corresponds to the respective rates of general metatarsal stress fracture in these groups. The discrepancy between strength and plantar pressure values in metatarsals 2 and 3 is consistent with the high incidence of stress fractures in these bones and underscores the importance of soft tissues, such as the plantar fascia and flexor musculature, in moderating metatarsal shaft strain.

Black or African American↗

Early hominin limb proportions.

Recent analyses and new fossil discoveries suggest that the evolution of hominin limb length proportions is complex, with evolutionary reversals and a decoupling of proportions within and between limbs. This study takes into account intraspecific variation to test whether or not the limb proportions of four early hominin associated skeletons (AL 288-1, OH 62, BOU-VP-12/1, and KNM-WT 15000) can be considered to be significantly different from one another. Exact randomization methods were used to compare the differences between pairs of fossil skeletons to the differences observed between all possible pairs of individuals within large samples of Gorilla gorilla, Pan troglodytes, Pongo pygmaeus, and Homo sapiens. Although the difference in humerofemoral proportions between OH 62 and AL 288-1 does not exceed variation in the extant samples, it is rare. When humerofemoral midshaft circumferences are compared, the difference between OH 62 and AL 288-1 is fairly common in extant species. This, in combination with error associated with the limb lengths estimates, suggests that it may be premature to consider H. (or Australopithecus) habilis as having more apelike limb proportions than those in A. afarensis. The humerofemoral index of BOU-VP-12/1 differs significantly from both OH 62 and AL 288-1, but not from KNM-WT 15000. Published length estimates, if correct, suggest that the relative forearm length of BOU-VP-12/1 is unique among hominins, exceeding those of the African apes and resembling the proportions in Pongo. Evidence that A. afarensis exhibited a less apelike upper:lower limb design than A. africanus (and possibly H. habilis) suggests that, if A. afarensis is broadly ancestral to A. africanus, the latter did not simply inherit primitive morphology associated with arboreality, but is derived in this regard. The fact that the limb proportions of OH 62 (and possibly KNM-ER 3735) are no more human like than those of AL 288-1 underscores the primitive body design of H. habilis.

Animals↗