Molar occlusion and jaw mechanics of the Eocene primate Adapis

Wear facets on molars of the Eocene primate Adapis magnus are described. Striations on these wear facets indicate three separate directions of mandibular movement during mastication. One direction corresponds to a first stage of mastication involving orthal retraction of the mandible. The remaining two directions correspond to buccal and lingual phases of a second stage of mastication involving a transverse movement of the mandible. The mechanics of jaw adduction are analysed for both the orthal retraction and transverse stages of mastication. During the orthal retraction stage the greatest component of bite force is provided by the temporalis muscles acting directly against the food with the mandible functioning as a link rather than as a lever. A geometrical argument suggests that during the transverse stage of mastication bite force is provided by the temporalis muscles of both sides, the ipsilateral medial and lateral pterygoid muscles, and the contralateral masseter muscle.     

The mandibular corpus of female primates: taxonomic, dietary, and allometric correlates of interspecific variations in size and shape

Measurements were taken on skulls of 253 adult female anthropoid primates from 32 species, in order to determine patterns and possible causes for variation among species in the cross-sectional size and shape of the mandibular corpus under M1. When all 32 species are considered as a group, there is a tendency for corpus shape to become more robust with increasing body size. However, this does not hold for colobines or cercopithecines evaluated separately. When diets are classified into the general categories of folivory or frugivory, neither size-adjusted measurements of mandibular corpus breadth and height, nor estimates of the second moments of inertia or the polar moment of inertia of the mandibular cross section, show any relationship to dietary variation among species. Species reported to include hard nuts in their diets have larger mandibular cross sections than other species, and the size of the corpus is significantly correlated with size of the dentition and molar enamel thickness. A biomechanical model taking into account frictional effects of tooth-to-tooth contact indicates that mandibular corpus robusticity may not be related to a large horizontal component of force during mastication.     

Masticatory biomechanics and its relevance to early hominid phylogeny: an examination of palatal thickness using finite-element analysis

It has been proposed that morphological characters functionally related to mastication may be unreliable indicators of early hominid phylogeny. One hypothesis states that masticatory characters are highly prone to homoplasy. A second hypothesis states that such characters are likely to be morphologically integrated and thus violate the assumption of character independence implicit in all phylogenetic analyses. Evaluation of these hypotheses requires that masticatory features be accurately identified, but, to date, there have been relatively few attempts to test precisely which early hominid features are functionally related to chewing. This paper uses finite-element analysis to evaluate the functional relationships of a character--palatal thickness--that is one of several Paranthropus synapomorphies putatively related to mastication. A finite-element model of 145,680 elements was created from sixty-one 2-mm-thick CT scans of a Macaca fascicularis skull. The model was assigned the elastic properties of facial bone and loaded with muscle forces corresponding to the moment of centric occlusion during mastication. The model was constrained so as to produce a reaction force (corresponding to the bite force) at M(1). With a few exceptions, the strain patterns in the finite-element model compare well with those gathered from published and unpublished bone-strain experiments. The model was then modified to have a thick palate. The model was reloaded using an identical loading regime, and the strain patterns of the original and thick-palate models were compared. Although a thickened palate acts to reduce palatal strain, strains are elevated in other facial regions. This suggests that a thick palate would not have evolved in isolation as an adaptation to withstand masticatory stress. Rather, a thick palate may have evolved in concert with a suite of other facial features that share a stress-resistance function. This appears to be consistent with hypotheses positing that at least some facial features related to chewing evolved in an integrated fashion. More functional studies of other facial features are needed, as are formal studies of morphological integration.     

Loading patterns and jaw movements during mastication in Macaca fascicularis: a bone-strain, electromyographic, and cineradiographic analysis

Rosette strain gage, electromyography (EMG), and cineradiographic techniques were used to analyze loading patterns and jaw movements during mastication in Macaca fascicularis. The cineradiographic data indicate that macaques generally swallow frequently throughout a chewing sequence, and these swallows are intercalated into a chewing cycle towards the end of a power stroke. The bone strain and jaw movement data indicate that during vigorous mastication the transition between fast close and the power stroke is correlated with a sharp increase in masticatory force, and they also show that in most instances the jaws of macaques are maximally loaded prior to maximum intercuspation, i.e. during phase I (buccal phase) occlusal movements. Moreover, these data indicate that loads during phase II (lingual phase) occlusal movements are ordinarily relatively small. The bone strain data also suggest that the duration of unloading of the jaw during the power stroke of mastication is largely a function of the relaxation time of the jaw adductors. This interpretation is based on the finding that the duration from 100% peak strain to 50% peak strain during unloading closely approximates the half-relaxation time of whole adductor jaw muscles of macaques. The EMG data of the masseter and medial pterygoid muscles have important implications for understanding both the biomechanics of the power stroke and the external forces responsible for the "wishboning" effect that takes place along the mandibular symphysis and corpus during the power stroke of mastication. Although both medial pterygoid muscles reach maximum EMG activity during the power stroke, the activity of the working-side medial pterygoid peaks after the balancing-side medial pterygoid. Associated with the simultaneous increase of force of the working-side medial pterygoid and the decrease of force of the balancing-side medial pterygoid is the persistently high level of EMG activity of the balancing-side deep masseter (posterior portion). This pattern is of considerable significance because the direction of force of both the working-side medial pterygoid and the balancing-side deep masseter are well aligned to aid in driving the working-side lower molars across the upper molars in the medial direction during unilateral mastication.(ABSTRACT TRUNCATED AT 400 WORDS)     

Effects of limb mass distribution on the ontogeny of quadrupedalism in infant baboons (Papio cynocephalus) and implications for the evolution of primate quadrupedalism

Primate quadrupedal kinematics differ from those of other mammals. Several researchers have suggested that primate kinematics are adaptive for safe travel in an arboreal, small-branch niche. This study tests a compatible hypothesis that primate kinematics are related to their limb mass distribution patterns. Primates have more distally concentrated limb mass than most other mammals due to their grasping hands and feet. Experimental studies have shown that increasing distal limb mass by adding weights to the limbs of humans and dogs influences kinematics. Adding weights to distal limb elements increases the natural period of a limb's oscillation, leading to relatively long swing and stride durations. It is therefore possible that primates' distal limb mass is responsible for some of their unique kinematics. This hypothesis was tested using a longitudinal ontogenetic sample of infant baboons (Papio cynocephalus). Because limb mass distribution changes with age in infant primates, this project examined how these changes influence locomotor kinematics within individuals. The baboons in this sample showed a shift in their kinematics as their limb mass distributions changed during ontogeny. When their limb mass was most distally concentrated (at young ages), stride frequencies were relatively low, stride lengths were relatively long, and stance durations were relatively long compared to older ages when limb mass was more proximally concentrated. These results suggest that the evolution of primate quadrupedal kinematics was tied to the evolution of grasping hands and feet.     

Sexual dimorphism and mechanics of the human hip: A multivariate assessment

The present research was undertaken to determine the effects of sexual dimorphism in the human pelvis and femur on the mechanics of human locomotion. The analysis was based on six biomechanical variables determined from 25 male and 32 female skeletal remains from the Dickson Mound site. Discriminant function analysis indicates that the mechanical variables which primarily contribute to dimorphism are the moment arm of the gluteus medius and the torque produced by the abductors at the hip. These mechanical aspects of hip function produce greater pressure on the femoral head in females.     

Biplanar X-ray motion analysis of the lower jaw movement during incisor interaction and mastication in the beaver (Castor fiber L. 1758)

The first biplanar X-ray motion analysis of mastication and food processing for Castor fiber is presented. While particles are chipped off interaction of incisors involves variable movements of the lower mandible and thus incisors. After jaw opening the tip of the lower incisors can reach different positions anteriorly of the upper incisors. Then the mandible moves upwards and backwards and brings the tips of the incisors into contact. The lower incisors slide along the wear facet of the upper to the ledge when the cheek teeth occlude. The glenoid fossa and lower jaw condyle are in close contact during incisor contact and no transverse movements are observed. Mastication involves interaction of the cheek teeth with no contact of the incisors. When the cheek teeth are in occlusal contact the mandible is moved forward and transverse, or mediolateral. In consecutive power strokes the jaw is moved alternately to the right and left side. When the jaw opens it is brought into a more central but not totally centred position. During mastication the condyles are positioned posteriorly to the glenoid allowing lateral movement of the mandible. The lateral movement is particularly noticeable in the anterior part of the mandible. With the lateral movements of the incisors one glenoid has to move posteriorly, the other anteriorly.     

The effects of long-standing limb loss on anatomical reorganization of the somatosensory afferents in the brainstem and spinal cord

We examined the terminations of sensory afferents in the brainstem and spinal cord of squirrel monkeys and prosimian galagos 4-8 years after a therapeutic forelimb or hindlimb amputation within 2 months of birth. In each animal, the distributions of labeled sensory afferent terminations from remaining body parts proximal to the limb stump were much more extensive than in normal animals. These sprouted afferents extended into the portions of the dorsal horn of the spinal cord as well as the cuneate and external cuneate nuclei of the brainstem (forelimb amputees) or spinal Clarke's column (hindlimb amputee) related to the amputated limb. Such reorganization in sensory afferents along with reorganization of the motor efferents to muscles (Wu and Kaas, J Neurosci 19 : 7679-7697, 1999, Neuron 28 : 967-978, 2000) may provide a basis for mislocated phantom sensations of missing forelimb movements accompanying actual shoulder movements during cortical stimulation or movement imagery in patients with amputations.     

Arthropod touch reception: structure and mechanics of the basal part of a spider tactile hair

Our previous studies on spider tactile hairs concentrated on the mechanical behavior of the hair shaft and the electrophysiological properties of the sensory cells. Here we focus on the structure and mechanical properties of the coupling of the hair shaft and the sensory terminals. 1. The functional design of the coupling provides for a combination of high sensitivity and protection against mechanical damage and overstimulation. The dendritic sheath is not directly coupled to the hair shaft. Rather, there is terminal connecting material between the dendrites and the hair shaft. 2. The hair shaft forms a first-order lever. Its acentric axis of rotation is located ca.3.5 m from its inner end. Displacement of the hair tip is scaled down by a factor of ca.750:1, not even considering the outer hair shafts bending. 3. At threshold the dendrite sheath displacement is ca. 0.05 m by forces in the order of 0.4–4×10^–6 N. 4. The hair shaft bends within the socket even before contacting it. The elasticities representing its suspension and bending in the socket can be described quantitatively by measuring the hairs restoring moments (range: 10^–9 Nm) and bending at different degrees of deflection.     

Modeling a fibula transplant in mandibular reconstructions: evaluation of the effects of a minimal number of osteotomies on the contour of the jaw.

The fibula osteocutaneous free flap has become the preferred method for most cases of mandibular reconstruction after oncologic surgical ablation. To recreate the parabolic form of the mandible, the fibula has to be divided up into segments using a closed wedge osteotomy technique. The number of osteotomies is preferably kept to a minimum so that segmental periosteal circulation is not compromised and also to keep operating time to a minimum. The limited number of osteotomies creates an angular contour. The aim of this study was to establish the degree to which overcorrection or undercorrection would occur when a subtotal reconstruction from ramus to ramus was simulated using five bony segments and four osteotomies. The study was carried out using 30 preserved jaws; the contour lines of the jaws were transferred onto tracing paper using a cardboard template. The contour of the mandible was divided into five sections (ramus, body, symphysis, body, and ramus). Because of the cutting off of the curvature in the original jaw outline, the lateral side of the body will become narrower and the chin broader. This also results in an underprojection (displacement) of the chin. To follow the original contour of the jaw as accurately as possible, all these anomalies must be minimized. The amount of under- and overprojection is calculated for a displacement of 1.0, 1.5, 2.5, 5.0, 7.5, and 10 mm of the chin. The most accurate reconstruction of the mandibular contour is achieved with a displacement of 1.5 or 2.5 mm. To preserve sufficient periosteal circulation, the minimum width of bone segments must be 15 mm or more. This concerns especially the symphysis section. On the basis of a fibula thickness of 14 mm, the internal bone width of the symphysis section is calculated. With a displacement of 1.5 mm, the average internal width of the bone segment is 14.8 mm, with a range of 9.9 to 23.0 mm (95 percent confidence interval, 12.8 to 16.7 mm). Therefore, a displacement of 2.5 mm with an internal bone width of 16.4 mm is preferred (range, 11.9 to 24.8 mm; 95 percent confidence interval, 15.5 to 18.2 mm). The loss of lateral projection is minimal (5.8 mm) and the resulting chin width is acceptable (average, 35.0 mm). In conclusion, we propose that in a subtotal procedure, an acceptable jaw reconstruction can be achieved with a limited number of osteotomies. The bone length of the symphysis section remains within safe limits. If the defect is of limited dimensions, then the resulting jaw contour is even more accurate.     

Uricoteley:its nature and origin during the evolution of tetrapod vertebrates

The hepatic mechanism for detoxication of ammonia formed during amino acid gluconeogenesis in uricotelic vertebrates requires the intramitochondrial synthesis of glutamine by glutamine synthetase. This glutamine then serves as a precursor of uric acid in the cytosol. The evolutionary development of uricoteley thus required the localization of glutamine synthetase in liver mitochondria. The mechanism for the mitochondrial import of glutamine synthetase in uricotelic vertebrate liver is not yet known. Tortoises, extant relatives of the stem reptiles, possess both the ureotelic and uricotelic hepatic systems. It therefore seems likely that the genetic events allowing the mitochondrial localization of glutamine synthetase in liver occurred in the amniote amphibian ancestors of the stem reptiles. The selection of ureoteley by the theropsids and of uricoteley by the sauropsids were major events in the divergence and subsequent evolution of these two lines. Once established in the sauropsid line, uricoteley has persisted through to the higher reptiles, crocodilians, and birds. Uricoteley was in part responsible for the radiation of the archosaurs during the Triassic as a water-conserving mechanism in the adult, thereby allowing them to invade the arid environments of that period. Contrary to dogma, uricoteley was probably of minor significance in the development of the cleidoic egg. Neither mammalian nor avian embryonic liver tissues catabolize amino acids to any great extent, so it is inappropriate to attribute to them a kind of "waste" nitrogen metabolism.     

Convergence of forelimb and hindlimb Natural Pendular Period in baboons (Papio cynocephalus) and its implication for the evolution of primate quadrupedalism

The patterns of muscle mass distribution along the lengths of limbs may have important effects on the mechanics and energetics of quadrupedalism. Specifically, Myers and Steudel (J. Morphol. 234 (1997) 183) have shown that fore- and hindlimb Natural Pendular Periods (NPPs) may affect quadrupedal kinematics and must converge to reduce locomotor energetic costs. This study quantifies patterns of limb mass distribution in a live sample of Papio cynocephalus using limb inertial properties (mass, center of mass, mass moment of inertia, and radius of gyration). These inertial properties are calculated using a geometric modeling technique similar to that of Crompton et al. (Am. J. phys. Anthrop. 99 (1996) 547). The inertial properties in Papio are compared to those of Canis from Myers and Steudel (J. Morphol. 234 (1997) 183). The Papio sample has convergent fore- and hindlimb NPPs. Additionally, these limb NPPs are relatively large compared to those of Canis due to the relatively distally distributed limb mass in the Papio sample (relatively large limb masses, relatively distal centers of mass and radii of gyration, and relatively large limb mass moments of inertia). This relatively distal limb mass appears related to the grasping abilities of their hands and feet. Causal links are explored between limb shape adaptations for grasping hands and feet and the kinematics of primate quadrupedalism. In particular, if primates in general follow Papio's limb mass distribution pattern, then relatively large limb NPPs may lead to the relatively low stride frequencies already documented for primates. The kinematics of primate quadrupedalism appears to have been strongly influenced by both selection for grasping hands and feet and selection for reduced locomotor energetic costs.     

Relationship between the critical phases of the physiological state of insect during supercooling and its dielectric permeability

The relationship between critical temperatures causing the cold torpor and freezing of liquid fractions of the body of honeybees and their dielectric permeability has been studied. It has been shown that the temperature at which the freezing of liquid fractions occurs, as distinct from the temperature inducing the cold torpor, depends on age and seasonal generation of insects. It has been found that changes in the temperature causing the torpor and freezing correlate with jumps of dielectric permeability.     

Relationship between the flexibility and the motility of actin filaments: effects of pH

Both the sliding velocity of fluorescently labeled actin filament and its persistence length as an index of the bending flexibility of the filament were examined in the motility assay as varying the pH values of the solution for preparing actin filaments. When the pH value was varied from 5.0 to 9.0 in the solution in which actin filaments were formed from the constituent monomers, the motile performance of Mg²⁺ bound actin filaments (Mg-F-actin) was apparently suppressed compared to the case of Ca²⁺ bound ones (Ca-F-actin). The persistence length for Ca-F-actin gradually increased with the increase of the pH value while the similar length for Mg-F-actin remained rather independent of the value. The largest sliding velocity of the filament, on the other hand, obtained at the persistence length of roughly 6 μm for both cases of Mg-F-actin and Ca-F-actin.     

Electromyography of the gluteal muscles in Hylobates,Pongo, and pan: Implications for the evolution of hominid bipedality

The gluteal musculature of primates has been a focus of great research interest among those who study human evolution. Most current theorists agree that gluteus superficialis (= maximus) need not have changed its action in the step from pongid to hominid, but dispute has arisen over a purported change in action and role of the gluteus medius. To clarify the functions of gluteus medius, gluteus superficialis, and tensor fasciae femoris during ape locomotion, we conducted a telemetered electromyographic study of these muscles in two gibbons, one orangutan, and four chimpanzees as they walked bipedally on the ground and on a horizontal tree trunk, walked quadrupedally on the same substrates, and climbed a vertical tree trunk. The results indicate that the gluteus medius of apes is not, as has been previously suggested, primarily an extensor of the thigh; its action is chiefly that of medial rotation. The role of the gluteus medius during bipedality is the same in apes and humans–to provide side-to-side balance of the trunk at the hip. The change in the hominid lateral balance mechanism can be viewed as primarily osteological, allowing preservation of the same muscle function with an extended thigh. As a result, the stride length is increased and there occurs a diminution of the demands placed on other muscles to maintain anteroposterior balance at the hip and knee. Our data also support the view that vertical climbing may be specifically preadaptive to bipedalism. One may picture the earliest hominid as part biped, when on the ground traveling between scattered food trees, and part climber, when moving from the ground to food.     

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.