Mechanical compensation in the evolution of the early hominin feeding apparatus.

Australopiths, a group of hominins from the Plio-Pleistocene of Africa, are characterized by derived traits in their crania hypothesized to strengthen the facial skeleton against feeding loads and increase the efficiency of bite force production. The crania of robust australopiths are further thought to be stronger and more efficient than those of gracile australopiths. Results of prior mechanical analyses have been broadly consistent with this hypothesis, but here we show that the predictions of the hypothesis with respect to mechanical strength are not met: some gracile australopith crania are as strong as that of a robust australopith, and the strength of gracile australopith crania overlaps substantially with that of chimpanzee crania. We hypothesize that the evolution of cranial traits that increased the efficiency of bite force production in australopiths may have simultaneously weakened the face, leading to the compensatory evolution of additional traits that reinforced the facial skeleton. The evolution of facial form in early hominins can therefore be thought of as an interplay between the need to increase the efficiency of bite force production and the need to maintain the structural integrity of the face.

Mapping the genicular arteries to provide a caution zone during knee surgery.

INTRODUCTION: Reports from the current literature show a lack of detail with depictions of the genicular arteries (GA). The intricate anatomy and infrequency of operating in the posterior knee may lead to surgeons being unfamiliar with the anatomy. The goal of this cadaveric study was to quantitatively map the arteries and create a caution zone that can be utilized when preparing and performing surgical procedures involving the knee. MATERIALS AND METHODS: The left knees of 46 cadavers were used. The distance of the GAs were from the joint line (JL) (+, superior to JL; -, inferior to JL) was measured in two locations: popliteal artery (PA) branch point and medial/lateral knee. The angle the artery traveled between these two points in the posterior knee was measured. A caution map was created. RESULTS: The superolateral GA branched from PA at +47.3 mm and traveled superiorly at 57.7° to +52.2 mm at the lateral knee. The superomedial GA branched from PA at +55.2 mm and traveled superiorly at 66.8° to +57.3 mm at the medial knee. The inferolateral GA branched from PA at -0.6 mm. It traveled superiorly at 74.1° or inferiorly at 62.1° to -1.0 mm at the lateral knee. The inferomedial GA branched from the PA at +9.9 mm. It traveled inferiorly at 21.2° to -33 mm at the medial knee. CONCLUSION: The GAs have a predictable pattern of location in the knee. There is a mismatch between medical textbooks and reality regarding arterial depictions. Knowledge regarding where the arteries are located may help reduce vascular complications in patients in the future.

Taking a big bite: Working together to better understand the evolution of feeding in primates.

The study of adaptation requires the integration of an array of different types of data. A single individual can find such integration daunting, if not impossible. In an effort to clarify the role of diet in the evolution of the primate craniofacial and dental apparatus, we assembled a team of researchers that have various types and degrees of expertise. This interaction has provided a range of insights for all contributors, and this has helped to refine questions, clarify the possibilities and limitations that laboratory and field settings offer, and further explore the ways in which laboratory and field data can be suitably integrated. A complete and accurate picture of dietary adaptation cannot be gained in isolation. Collaboration provides the bridge to a more holistic view of primate biology and evolution.

Primate dietary ecology in the context of food mechanical properties.

Substantial variation exists in the mechanical properties of foods consumed by primate species. This variation is known to influence food selection and ingestion among non-human primates, yet no large-scale comparative study has examined the relationships between food mechanical properties and feeding strategies. Here, we present comparative data on the Young's modulus and fracture toughness of natural foods in the diets of 31 primate species. We use these data to examine the relationships between food mechanical properties and dietary quality, body mass, and feeding time. We also examine the relationship between food mechanical properties and categorical concepts of diet that are often used to infer food mechanical properties. We found that traditional dietary categories, such as folivory and frugivory, did not faithfully track food mechanical properties. Additionally, our estimate of dietary quality was not significantly correlated with either toughness or Young's modulus. We found a complex relationship among food mechanical properties, body mass, and feeding time, with a potential interaction between median toughness and body mass. The relationship between mean toughness and feeding time is straightforward: feeding time increases as toughness increases. However, when considering median toughness, the relationship with feeding time may depend upon body mass, such that smaller primates increase their feeding time in response to an increase in median dietary toughness, whereas larger primates may feed for shorter periods of time as toughness increases. Our results emphasize the need for additional studies quantifying the mechanical and chemical properties of primate diets so that they may be meaningfully compared to research on feeding behavior and jaw morphology.

Age-related variation in the mechanical properties of foods processed by Sapajus libidinosus.

OBJECTIVES: The diet of tufted capuchins (Sapajus) is characterized by annual or seasonal incorporation of mechanically protected foods. Reliance on these foods raises questions about the dietary strategies of young individuals that lack strength and experience to access these resources. Previous research has demonstrated differences between the feeding competencies of adult and juvenile tufted capuchins. Here we test the hypothesis that, compared to adults, juveniles will process foods with lower toughness and elastic moduli. MATERIALS AND METHODS: We present data on variation in the toughness and elastic modulus of food tissues processed by Sapajus libidinosus during the dry season at Fazenda Boa Vista, Brazil. Food mechanical property data were collected using a portable universal mechanical tester. RESULTS: Results show that food tissues processed by the capuchins showed significant differences in toughness and stiffness. However, we found no relationship between an individual's age and mean or maximum food toughness or elastic modulus, indicating both juvenile and adult S. libidinosus are able to process foods of comparable properties. DISCUSSION: Although it has been suggested that juveniles avoid mechanically protected foods, age-related differences in feeding competence are not solely due to variation in food toughness or stiffness. Other factors related to food type (e.g., learning complex behavioral sequences, achieving manual dexterity, obtaining physical strength to lift stone tools, or recognizing subtle cues about food state) combined with food mechanical properties better explain variation in juvenile feeding competency.

The effects of ecology and evolutionary history on robust capuchin morphological diversity.

Recent molecular work has confirmed the long-standing morphological hypothesis that capuchins are comprised of two distinct clades, the gracile (untufted) capuchins (genus Cebus, Erxleben, 1777) and the robust (tufted) capuchins (genus Sapajus Kerr, 1792). In the past, the robust group was treated as a single, undifferentiated and cosmopolitan species, with data from all populations lumped together in morphological and ecological studies, obscuring morphological differences that might exist across this radiation. Genetic evidence suggests that the modern radiation of robust capuchins began diversifying ∼2.5 Ma, with significant subsequent geographic expansion into new habitat types. In this study we use a morphological sample of gracile and robust capuchin craniofacial and postcranial characters to examine how ecology and evolutionary history have contributed to morphological diversity within the robust capuchins. We predicted that if ecology is driving robust capuchin variation, three distinct robust morphotypes would be identified: (1) the Atlantic Forest species (Sapajus xanthosternos, S. robustus, and S. nigritus), (2) the Amazonian rainforest species (S. apella, S. cay and S. macrocephalus), and (3) the Cerrado-Caatinga species (S. libidinosus). Alternatively, if diversification time between species pairs predicts degree of morphological difference, we predicted that the recently diverged S. apella, S. macrocephalus, S. libidinosus, and S. cay would be morphologically comparable, with greater variation among the more ancient lineages of S. nigritus, S. xanthosternos, and S. robustus. Our analyses suggest that S. libidinosus has the most derived craniofacial and postcranial features, indicative of inhabiting a more terrestrial niche that includes a dependence on tool use for the extraction of imbedded foods. We also suggest that the cranial robusticity of S. macrocephalus and S. apella are indicative of recent competition with sympatric gracile capuchin species, resulting in character displacement.

Viewpoints: diet and dietary adaptations in early hominins: the hard food perspective.

Recent biomechanical analyses examining the feeding adaptations of early hominins have yielded results consistent with the hypothesis that hard foods exerted a selection pressure that influenced the evolution of australopith morphology. However, this hypothesis appears inconsistent with recent reconstructions of early hominin diet based on dental microwear and stable isotopes. Thus, it is likely that either the diets of some australopiths included a high proportion of foods these taxa were poorly adapted to consume (i.e., foods that they would not have processed efficiently), or that aspects of what we thought we knew about the functional morphology of teeth must be wrong. Evaluation of these possibilities requires a recognition that analyses based on microwear, isotopes, finite element modeling, and enamel chips and cracks each test different types of hypotheses and allow different types of inferences. Microwear and isotopic analyses are best suited to reconstructing broad dietary patterns, but are limited in their ability to falsify specific hypotheses about morphological adaptation. Conversely, finite element analysis is a tool for evaluating the mechanical basis of form-function relationships, but says little about the frequency with which specific behaviors were performed or the particular types of food that were consumed. Enamel chip and crack analyses are means of both reconstructing diet and examining biomechanics. We suggest that current evidence is consistent with the hypothesis that certain derived australopith traits are adaptations for consuming hard foods, but that australopiths had generalized diets that could include high proportions of foods that were both compliant and tough.

Venous Tributaries of the Lip: Implications for Lip Filler Injection.

BACKGROUND: Demand for lip filler injection continues to increase. Despite the current literature's acknowledgement of the role both venous and arterial vasculature play in minor and major side effects, research addressing the venous vasculature of the lower one-third of the face is scarce. METHODS: A photographic analysis of the venous vasculature of 26 participants was performed using a vein transilluminator to display the venous flow around the perioral region. The data were analyzed for commonalities among participants and then compared with common lip filler injection techniques and locations. RESULTS: Venous tributaries were identified in all patients, with slight variation in pattern, superior to the upper vermilion border between the nasolabial fold and philtral column on each side of the mouth. Venous tributaries were noted approximately 1 to 1.5 cm lateral to the oral commissures extending inferiorly to the chin and along the labiomental crease. Four areas of venous pooling were deemed significant: a small area approximately 2 mm superior to the Cupid's bow, along the middle tubercle of the upper lip, along the wet-dry line of the lower lip; and centrally along the vermilion border between the lower lip tubercles. CONCLUSIONS: Perioral venous mapping provides a guide for injectors performing lip enhancement procedures in identifying areas at risk for injury because of venous pooling. Avoiding these anatomically vulnerable regions can minimize the potential for inflammation and ecchymosis associated with intravenous injection and prevent dissatisfactory aesthetic results because of lumps, excessive bruising, swelling, or asymmetry.

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.

Relationships between the diet and dentition of Asian leaf monkeys.

Colobines have been generally described as primates that use the anterior teeth minimally, but the posterior teeth extensively, to process leaves and related food items. However, variation among leaf monkeys in both anterior and posterior dental morphology has been recognized for decades. In this study, we turn to Hylander's (Science 189 (1975) 1095-1098) analysis of anterior incisor row length and Kay's (Adaptations for foraging in nonhuman primates, 1984) examination of relative molar crest length to test hypotheses proposed by them for Asian colobines. We present findings based on data from the largest Asian colobine sample measured to date. Our findings for incisor row length and molar cresting are not amenable to broad generalizations. In those instances when our morphological findings concur with those of Hylander (Science 189 (1975) 1095-1098) and Kay and Hylander (The ecology of arboreal folivores, 1978), the ecological evidence seldom supports the morphological predictions. The disassociation between diet and dental patterns may be a consequence of differential selection by fallback foods, anthropogenic disturbance or climatic shifts limiting preferred diets, or the use of food types as opposed to food mechanical properties for dietary categorization. We also found that in the case of both incisor row length and molar crest length, the patterns for males and females differed markedly. The reasons for these differences may in part be ascribed to the metabolic challenges faced by females and subsequent niche partitioning. We propose integrated analyses of the ingestive and digestive systems of our study taxa to clarify relationships among behavior, dental morphology, and diet in extant and extinct colobines.

The feeding biomechanics and dietary ecology of Australopithecus africanus.

The African Plio-Pleistocene hominins known as australopiths evolved a distinctive craniofacial morphology that traditionally has been viewed as a dietary adaptation for feeding on either small, hard objects or on large volumes of food. A historically influential interpretation of this morphology hypothesizes that loads applied to the premolars during feeding had a profound influence on the evolution of australopith craniofacial form. Here, we test this hypothesis using finite element analysis in conjunction with comparative, imaging, and experimental methods. We find that the facial skeleton of the Australopithecus type species, A. africanus, is well suited to withstand premolar loads. However, we suggest that the mastication of either small objects or large volumes of food is unlikely to fully explain the evolution of facial form in this species. Rather, key aspects of australopith craniofacial morphology are more likely to be related to the ingestion and initial preparation of large, mechanically protected food objects like large nuts and seeds. These foods may have broadened the diet of these hominins, possibly by being critical resources that australopiths relied on during periods when their preferred dietary items were in short supply. Our analysis reconciles apparent discrepancies between dietary reconstructions based on biomechanics, tooth morphology, and dental microwear.

Spatial Visualization of Human Anatomy through Art Using Technical Drawing Exercises.

Spatial visualization, the ability to mentally rotate three-dimensional (3D) images, plays a significant role in anatomy education. This study examines the impact of technical drawing exercises on the improvement of spatial visualization and anatomy education in a Neuroscience course. First-year medical students (n = 84) were randomly allocated into a control group (n = 41) or art-training group (n = 43). Variables including self-reported artistic drawing ability, previous technical drawing experience, or previous anatomy laboratory exposure were gathered. Participants who self-identified as artistic individuals were equally distributed between the two groups. Students in the art-training group attended four 1-hour sessions to solve technical drawing worksheets. All participants completed two Mental Rotations Tests (MRT), which were used to assess spatial visualization. Data were also collected from two neuroscience written examinations and an anatomical "tag test" practical examination. Participants in the art-training and control groups improved on the MRT. The mean of written examination two was significantly higher (P = 0.007) in the art-training group (12.95) than the control group (11.48), and higher (P = 0.027) in those without technical drawing experience (12.44) than those with (11.00). The mean of the anatomical practical was significantly higher (P = 0.010) in those without artistic ability (46.24) than those with (42.00). These results suggest that completing technical drawing worksheets may aid in solving anatomy-based written examination questions on complex brain regions, but further research is needed to determine its implication on anatomy practical scores. These results propose a simple method of improving spatial visualization in anatomy education.

A finite element analysis of masticatory stress hypotheses.

Understanding how the skull transmits and dissipates forces during feeding provides insights into the selective pressures that may have driven the evolution of primate skull morphology. Traditionally, researchers have interpreted masticatory biomechanics in terms of simple global loading regimes applied to simple shapes (i.e., bending in sagittal and frontal planes, dorsoventral shear, and torsion of beams and cylinders). This study uses finite element analysis to examine the extent to which these geometric models provide accurate strain predictions in the face and evaluate whether simple global loading regimes predict strains that approximate the craniofacial deformation pattern observed during mastication. Loading regimes, including those simulating peak loads during molar chewing and those approximating the global loading regimes, were applied to a previously validated finite element model (FEM) of a macaque (Macaca fascicularis) skull, and the resulting strain patterns were compared. When simple global loading regimes are applied to the FEM, the resulting strains do not match those predicted by simple geometric models, suggesting that these models fail to generate accurate predictions of facial strain. Of the four loading regimes tested, bending in the frontal plane most closely approximates strain patterns in the circumorbital region and lateral face, apparently due to masseter muscle forces acting on the zygomatic arches. However, these results indicate that no single simple global loading regime satisfactorily accounts for the strain pattern found in the validated FEM. Instead, we propose that FE models replace simple cranial models when interpreting bone strain data and formulating hypotheses about craniofacial biomechanics.

Analysis of Nipple-Areola Complex Localization Using Male Cadavers: Considerations for Gender-Affirming Surgery.

BACKGROUND: Masculinizing chest reconstruction is the most common gender-affirming surgery in transgender males. Despite the current literature's acknowledgment of the vital role that proper placement of the nipple-areola complex (NAC) plays in a masculine chest contour, there is still much debate regarding the best anatomical landmarks to achieve the desired result. OBJECTIVES: The primary aim of this study is to determine which landmarks for NAC placement can be applied across diverse body types and aid surgeons in creating a masculine chest. METHODS: Twenty-five formaldehyde-embalmed male cadavers were analyzed by conducting various measurements of the NAC, nipple, and surrounding bony and muscular landmarks to identify the most consistent landmarks for proper NAC placement. Linear regression analyses were run to determine how the distance between nipple to respective landmarks varied based on antemortem body mass index (BMI), height, weight, and age. RESULTS: The measurements for the inferior and lateral borders of the pectoralis major muscle (PMM) displayed the least amount of variance of all the anatomical landmarks studied. Additionally, there was no significant change in these pectoral measurements with varying BMI, height, weight, or age, indicating that these measurements are reliable landmarks for NAC placement across various body types. The average NAC placement in relation to the inferior and lateral borders of PMM was around 2.5 and 2.0 cm, respectively. CONCLUSIONS: Our cadaveric analysis indicates that aesthetically pleasing masculine chest results can be produced consistently across varying body types when adhering to a simple pectoral approach in NAC placement.

The Critical Shoulder Angle as a Diagnostic Measure for Osteoarthritis and Rotator Cuff Pathology.

The purpose of this study was to correlate critical shoulder angle (CSA), a measurement that takes into account both glenoid tilt and the acromial index (AI), with shoulder pathologies as presented in an earlier study by Moor et al. (2013). Based on Moor et al.'s predicted normal CSA range of 30-35°, we hypothesized that a greater-than-normal CSA would be correlated to or associated with rotator cuff pathology, while a smaller-than-normal CSA would be associated with osteoarthritis (OA). Following Moore et al., we utilized Grashey radiographic imaging because it provides the clearest view of the entire glenoid fossa and acromion. We analyzed 323 anterior-posterior (AP) radiographs to identify and measure the CSA, classifying each patient into one of five groups [none reported (n=94), mild OA (n=156), moderate OA (n=36), severe OA (n=37), and rotator cuff pathology (n=40)]. Our results were statistically significant, supporting the association of smaller CSAs with OA and larger CSAs with rotator cuff pathology. CSA measurements could provide a new means for identifying shoulder pathology and thereby reduce the need for costly and timely imaging techniques. CSA values could also provide useful information to utilize preventatively with interventions such as physical therapy to alter the CSA and reduce the prevalence of OA and shoulder arthroplasties. This study builds on the findings of Moore et al. in creating a correlation between CSA and shoulder pathology.

Ingestive behaviors in bearded capuchins (Sapajus libidinosus).

The biomechanical and adaptive significance of variation in craniodental and mandibular morphology in fossil hominins is not always clear, at least in part because of a poor understanding of how different feeding behaviors impact feeding system design (form-function relationships). While laboratory studies suggest that ingestive behaviors produce variable loading, stress, and strain regimes in the cranium and mandible, understanding the relative importance of these behaviors for feeding system design requires data on their use in wild populations. Here we assess the frequencies and durations of manual, ingestive, and masticatory behaviors from more than 1400 observations of feeding behaviors video-recorded in a wild population of bearded capuchins (Sapajus libidinosus) at Fazenda Boa Vista in Piauí, Brazil. Our results suggest that ingestive behaviors in wild Sapajus libidinosus were used for a range of food material properties and typically performed using the anterior dentition. Coupled with previous laboratory work indicating that ingestive behaviors are associated with higher mandibular strain magnitudes than mastication, these results suggest that ingestive behaviors may play an important role in craniodental and mandibular design in capuchins and may be reflected in robust adaptations in fossil hominins.

Hard plant tissues do not contribute meaningfully to dental microwear: evolutionary implications.

Reconstructing diet is critical to understanding hominin adaptations. Isotopic and functional morphological analyses of early hominins are compatible with consumption of hard foods, such as mechanically-protected seeds, but dental microwear analyses are not. The protective shells surrounding seeds are thought to induce complex enamel surface textures characterized by heavy pitting, but these are absent on the teeth of most early hominins. Here we report nanowear experiments showing that the hardest woody shells - the hardest tissues made by dicotyledonous plants - cause very minor damage to enamel but are themselves heavily abraded (worn) in the process. Thus, hard plant tissues do not regularly create pits on enamel surfaces despite high forces clearly being associated with their oral processing. We conclude that hard plant tissues barely influence microwear textures and the exploitation of seeds from graminoid plants such as grasses and sedges could have formed a critical element in the dietary ecology of hominins.

The importance of fallback foods in primate ecology and evolution.

The role of fallback foods in shaping primate ranging, socioecology, and morphology has recently become a topic of particular interest to biological anthropologists. Although the use of fallback resources has been noted in the ecological and primatological literature for a number of decades, few attempts have been made to define fallback foods or to explore the utility of this concept for primate evolutionary biologists and ecologists. As a preface to this special issue of the American Journal of Physical Anthropology devoted to the topic of fallback foods in primate ecology and evolution, we discuss the development and use of the fallback concept and highlight its importance in primatology and paleoanthropology.

Fallback foraging as a way of life: using dietary toughness to compare the fallback signal among capuchins and implications for interpreting morphological variation.

The genus Cebus is one of the best extant models for examining the role of fallback foods in primate evolution. Cebus includes the tufted capuchins, which exhibit skeletal features for the exploitation of hard and tough foods. Paradoxically, these seemingly "specialized" taxa belong to the most ubiquitous group of closely related primates in South America, thriving in a range of different habitats. This appears to be a consequence of their ability to exploit obdurate fallback foods. Here we compare the toughness of foods exploited by two tufted capuchin species at two ecologically distinct sites; C. apella in a tropical rainforest, and C. libidinosus in a cerrado forest. We include dietary data for one untufted species (C. olivaceus) to assess the degree of difference between the tufted species. These data, along with information on skeletal morphology, are used to address whether or not a fallback foraging species exhibits a given suite of morphological and behavioral attributes, regardless of habitat. Both tufted species ingest and masticate a number of exceedingly tough plant tissues that appear to be used as fallback resources, however, C. libidinosus has the toughest diet both in terms of median and maximal values. Morphologically, C. libidinosus is intermediate in absolute symphyseal and mandibular measurements, and in measures of postcranial robusticity, but exhibits a higher intermembral index than C. apella. We propose that this incongruence between dietary toughness and skeletal morphology is the consequence of C. libidinosus' use of tools while on the ground for the exploitation of fallback foods.

Indentation as a technique to assess the mechanical properties of fallback foods.

A number of living primates feed part-year on seemingly hard food objects as a fallback. We ask here how hardness can be quantified and how this can help understand primate feeding ecology. We report a simple indentation methodology for quantifying hardness, elastic modulus, and toughness in the sense that materials scientists would define them. Suggested categories of fallback foods-nuts, seeds, and root vegetables-were tested, with accuracy checked on standard materials with known properties by the same means. Results were generally consistent, but the moduli of root vegetables were overestimated here. All these properties are important components of what fieldworkers mean by hardness and help understand how food properties influence primate behavior. Hardness sensu stricto determines whether foods leave permanent marks on tooth tissues when they are bitten on. The force at which a food plastically deforms can be estimated from hardness and modulus. When fallback foods are bilayered, consisting of a nutritious core protected by a hard outer coat, it is possible to predict their failure force from the toughness and modulus of the outer coat, and the modulus of the enclosed core. These forces can be high and bite forces may be maximized in fallback food consumption. Expanding the context, the same equation for the failure force for a bilayered solid can be applied to teeth. This analysis predicts that blunt cusps and thick enamel will indeed help to sustain the integrity of teeth against contacts with these foods up to high loads.