Ontogenetic biomechanics of tufted (Sapajus) and untufted (Cebus) capuchin mandibles.

OBJECTIVES: Cortical bone geometry is commonly used to investigate biomechanical properties of primate mandibles. However, the ontogeny of these properties is less understood. Here we investigate changes in cortical bone cross-sectional properties throughout capuchin ontogeny and compare captive versus wild, semi-provisioned groups. Tufted capuchins (Sapajus spp.) are known to consume relatively hard/tough foods, while untufted capuchins (Cebus spp.) exploit less mechanically challenging foods. Previous research indicates dietary differences are present early in development and adult Sapajus mandibles can resist higher bending/shear/torsional loads. MATERIALS AND METHODS: This study utilized microCT scans of 22 Cebus and 45 Sapajus from early infancy to adulthood from three sample populations: one captive Cebus, one captive Sapajus, and one semi-provisioned, free-ranging Sapajus. Mandibular cross-sectional properties were calculated at the symphysis, P3, and M1. If the tooth had not erupted, its position within the crypt was used. A series of one-way ANOVAs were performed to assess differences between and within the sample populations. RESULTS: Mandible robusticity increases across ontogeny for all three sample populations. Sapajus were better able to withstand bending and torsional loading even early in ontogeny, but no difference in shear resistance was found. Semi-provisioned, free-ranging Sapajus tend to show increased abilities to resist bending and torsional loading but not shear loading compared to captive Sapajus. DISCUSSION: This study helps advance our understanding of the primate masticatory system development and opens the door for further studies into adaptive plasticity in shaping the masticatory apparatus of capuchins and differences in captive versus free-ranging sample populations.

Ontogenetic changes in jaw leverage and skull shape in tufted and untufted capuchins.

The ontogeny of feeding is characterized by shifting functional demands concurrent with changes in craniofacial anatomy; relationships between these factors will look different in primates with disparate feeding behaviors during development. This study examines the ontogeny of skull morphology and jaw leverage in tufted (Sapajus) and untufted (Cebus) capuchin monkeys. Unlike Cebus, Sapajus have a mechanically challenging diet and behavioral observations of juvenile Sapajus suggest these foods are exploited early in development. Landmarks were placed on three-dimensional surface models of an ontogenetic series of Sapajus and Cebus skulls (n = 53) and used to generate shape data and jaw-leverage estimates across the tooth row for three jaw-closing muscles (temporalis, masseter, medial pterygoid) as well as a weighted combined estimate. Using geometric morphometric methods, we found that skull shape diverges early and shape is significantly different between Sapajus and Cebus throughout ontogeny. Additionally, jaw leverage varies with age and position on the tooth row and is greater in Sapajus compared to Cebus when calculated at the permanent dentition. We used two-block partial least squares analyses to identify covariance between skull shape and each of our jaw muscle leverage estimates. Sapajus, but not Cebus, has significant covariance between all leverage estimates at the anterior dentition. Our findings show that Sapajus and Cebus exhibit distinct craniofacial morphologies early in ontogeny and strong covariance between leverage estimates and craniofacial shape in Sapajus. These results are consistent with prior behavioral and comparative work suggesting these differences are a function of selection for exploiting mechanically challenging foods in Sapajus, and further emphasize that these differences appear quite early in ontogeny. This research builds on prior work that has highlighted the importance of understanding ontogeny for interpreting adult morphology.

Ontogenetic changes in bite force and gape in tufted capuchins.

Bite force and gape are two important performance metrics of the feeding system, and these metrics are inversely related for a given muscle size because of fundamental constraints in sarcomere length-tension relationships. How these competing performance metrics change in developing primates is largely unknown. Here, we quantified in vivo bite forces and gapes across ontogeny and examined these data in relation to body mass and cranial measurements in captive tufted capuchins, Sapajus spp. Bite force and gape were also compared across geometric and mechanical properties of mechanically challenging foods to investigate relationships between bite force, gape and food accessibility (defined here as the ability to breach shelled nuts). Bite forces at a range of gapes and feeding behavioral data were collected from a cross-sectional ontogenetic series of 20 captive and semi-wild tufted capuchins at the Núcleo de Procriação de Macacos-Prego Research Center in Araçatuba, Brazil. These data were paired with body mass, photogrammetric measures of jaw length and facial width, and food geometric and material properties. Tufted capuchins with larger body masses had absolutely higher in vivo bite forces and gapes, and animals with wider faces had absolutely higher bite forces. Bite forces and gapes were significantly smaller in juveniles compared with subadults and adults. These are the first primate data to empirically demonstrate the gapes at which maximum active bite force is generated and to demonstrate relationships to food accessibility. These data advance our understanding of how primates meet the changing performance demands of the feeding system during development.

Fiber-type phenotype of the jaw-closing muscles in Gorilla gorilla, Pan troglodytes, and Pan paniscus: A test of the Frequent Recruitment Hypothesis.

Skeletal muscle fiber types are important determinants of the contractile properties of muscle fibers, such as fatigue resistance and shortening velocity. Yet little is known about how jaw-adductor fiber types correlate with feeding behavior in primates. Compared with chimpanzees and bonobos, gorillas spend a greater percentage of their daily time feeding and shift to herbaceous vegetation when fruits are scarce. We thus used the African apes to test the hypothesis that chewing with unusually high frequency is correlated with the expression in the jaw adductors of a high proportion of type 1 (slow, fatigue-resistant) fibers at the expense of other fiber types (the Frequent Recruitment Hypothesis). We used immunohistochemistry to determine the presence and distribution of the four major myosin heavy chain (MHC) isoforms in the anterior superficial masseter (ASM), superficial anterior temporalis, and deep anterior temporalis of four Gorilla gorilla, two Pan paniscus, and four Pan troglodytes. Serial sections were stained against slow (MHC-1/-α-cardiac) and fast (MHC-2/-M) fibers. Fibers were counted and scored for staining intensity, and fiber cross-sectional areas (CSAs) were measured and used to estimate percentage of CSA of each MHC isoform. Hybrid fibers accounted for nearly 100% of fiber types in the masseter and temporalis of all three species, resulting in three main hybrid phenotypes. As predicted, the gorilla ASM and deep anterior temporalis comprised a greater percentage of CSA of the slower, fatigue-resistant hybrid fiber type, significantly so for the ASM (p = 0.015). Finally, the results suggest that fiber phenotype of the chewing muscles contributes to behavioral flexibility in ways that would go undetected in paleontological studies relying solely on morphology of the bony masticatory apparatus.

The Source and the Course of the Articular Branches to the T4-T8 Zygapophysial Joints.

OBJECTIVE: To define the source and the course of the articular branches to the midthoracic zygapophysial ("z") joints. DESIGN: Cadaveric dissection. SETTING: The Gross Anatomy Laboratory of the Duke University School of Medicine. SUBJECTS: Ten human cadaveric thoraces. METHODS: Gross and stereoscopic dissection of dorsal rami T4-T8 was performed bilaterally on 10 adult embalmed cadavers. The medial and lateral branches were traced to their origins from the dorsal rami, and the course of the articular nerves was documented through digital photography. Radio-opaque wire (20 gauge) was applied to the nerves. Fluoroscopic images were obtained to delineate their radiographic course with respect to osseous landmarks. RESULTS: Forty-eight inferior articular branches were identified. Three (6.3%) originated from the medial branch and 44 (91.7%) from the dorsal ramus. One was indeterminate. Fifty-one superior articular branches were identified. Eight (15.7%) originated from the medial branch and 43 (84.3%) from the dorsal ramus. In 12% of cases (6/50), there was side-to-side asymmetry in the origins of the articular branches. Nerves were commonly suspended in the intertransverse space. The articular branches contacted an osseous structure in only 39% of cases. As previously reported, a "descending branch" was not identified in any specimen. CONCLUSIONS: Articular branches to the T4-T8 z-joints have substantial inter- and intraspecimen variability of origin. They typically arise from the dorsal ramus rather than the medial branch and frequently do not contact any osseous structure to allow percutaneous needle placement.

Interpreting skeletal growth in the past from a functional and physiological perspective.

The study of juvenile skeletal remains can yield important insights into the health, behavior, and biological relationships of past populations. However, most studies of past skeletal growth have been limited to relatively simple metrics. Considering additional skeletal parameters and taking a broader physiological perspective can provide a more complete assessment of growth patterns and environmental and genetic effects on those patterns. We review here some alternative approaches to ontogenetic studies of archaeological and paleontological skeletal material, including analyses of body size (stature and body mass) and cortical bone structure of long bone diaphyses and the mandibular corpus. Together such analyses can shed new light on both systemic and localized influences on bone growth, and the metabolic and mechanical factors underlying variation in growth.

Dietary effects on development of the human mandibular corpus.

The extent to which the mandibular corpus exhibits developmental plasticity has important implications for interpreting variation in adult and juvenile mandibular morphology in the archaeological and paleontological record. Here, we examine ontogenetic changes in mandibular corpus breadth, rigidity, and strength in two population samples with contrasting diets: late prehistoric Tigara from Point Hope, Alaska, characterized by a very demanding masticatory regime, and proto-historic Arikara from the Sully Site in South Dakota, with a less demanding regime. A total of 52 juvenile and 11 adult Tigara, and 32 juvenile and 10 adult Arikara were included in the study. Juveniles ranged in age from 1 to 17 years, with good representation of younger (1-6-year-old) juveniles (20 Arikara, 18 Tigara). Superoinferior and buccolingual external and cortical bone breadths of mandibles were measured at the Pm(4) -M(1) and M(1) -M(2) junctions using calipers and biplanar radiographs, respectively. An asymmetrical hollow beam model was employed to reconstruct cross sections and calculate bending rigidities and strengths in the sagittal and transverse planes. Among adults, Tigara have greater transverse corpus width, bending rigidity, and strength, and ratios of transverse to sagittal dimensions than Arikara. This shape difference develops gradually during growth, with only weak trends among young juveniles, increasing to near-adult contrasts among adolescents. These results support a role for functional mechanical loading of the mandible during growth in producing adult differences in mandibular corpus morphology.

Awareness during anesthesia.

The number of patients who have preoperative anxiety over possibly "waking up" in the middle of surgery has increased dramatically over the last decade. McCleane and Cooper found that more than 50% of 247 patients were concerned that they would not be asleep during their surgery. Even after having an adequate anesthetic, 25% were still worried about being asleep with future anesthetics. With increased media coverage, these anxieties are not likely to go away anytime soon. For the patient, awareness or recall while under general anesthesia is a frightening experience that can lead to debilitating emotional injury and even post-traumatic stress disorder. For anesthesiologists, awareness under anesthesia ranks second only to death as a "dreaded" complication. This chapter reviews the incidence, etiology, psychological sequelae, medicolegal consequences, and prevention of awareness during anesthesia.