A model of temporomandibular joint function in anthropoid primates based on condylar movements during mastication

The hypothesis that the shape of the bony temporomandibular joint (TMJ) is functionally related to sagittal sliding of the condyle during mastication is tested, and a model of the relation of sagittal sliding to mandibular size, TMJ shape, and diet is developed. Sagittal sliding is defined as fore-aft motion of the condyle during mandibular translation and/or angular rotation. Ascending ramus height is used as a structural correlate of the distance between the condyle and the mandibular axis of rotation (CR). Cineradiographic data on sagittal sliding and gape during mastication in Ateles spp., Macaca fascicularis, Papio anubis, and Pan troglodytes in conjunction with comparative data on mandibular size and TMJ shape are used to evaluate the hypothesis. The results show that 1) linear and angular gape are highly positively correlated with sagittal sliding, 2) pure mandibular translation is rare during mastication, 3) the CR is rarely if ever located at the condyle during mastication, 4) angular gape should be standardized in interindividual comparisons of sagittal sliding, and 5) the height of the ascending ramus (and by inference the CR-to-condyle distance) is highly positively correlated with absolute sagittal sliding. Sagittal sliding relative to the length of the articular eminence was the variable used to explore the relation between TMJ shape and sliding. This variable standardized absolute sagittal sliding relative to joint size. The relative depth and orientation of the articular eminence were not correlated with relative sagittal sliding. The anteroposterior curvature of the condyle was highly negatively correlated with relative sagittal sliding. Flat condyles are associated with large amounts of relative sagittal sliding. A flat condyle increases joint contact area, which reduces joint stress. A flat condyle also increases joint congruence, and this may facilitate the combined sliding and rolling motion of the condyle when the sliding motion is relatively large. The shape of the entoglenoid process was also positively correlated with relative sagittal sliding. A relatively large entoglenoid process may help to guide sagittal sliding and prevent excessive mediolateral sliding of the condyle. The functional model makes a number of predictions about the correlations between food consistency and food object size, mandibular size, TMJ shape, and sagittal sliding of the condyle during mastication and incision. Am J Phys Anthropol 109:67–88, 1999. © 1999 Wiley-Liss, Inc.     

Spinal constraint modulates head instantaneous center of rotation and dictates head angular motion

The head is kinematically constrained to the torso through the spine and thus, the spine dictates the amount of output head angular motion expected from an input impact. Here, we investigate the spinal kinematic constraint by analyzing the head instantaneous center of rotation (HICOR) with respect to the torso in head/neck sagittal extension and coronal lateral flexion during mild loads applied to 10 subjects. We found the mean HICOR location was near the C5-C6 intervertebral joint in sagittal extension, and T2-T3 intervertebral joint in coronal lateral flexion. Using the impulse-momentum relationship normalized by subject mass and neck length, we developed a non-dimensional analytical ratio between output angular velocity and input linear impulse as a function of HICOR location. The ratio was 0.65 and 0.50 in sagittal extension and coronal lateral flexion respectively, implying 30% greater angular velocities in sagittal extension given an equivalent impulse. Scaling to subject physiology also predicts larger required impulses given greater subject mass and neck length to achieve equivalent angular velocities, which was observed experimentally. Furthermore, the HICOR has greater motion in sagittal extension than coronal lateral flexion, suggesting the head and spine can be represented with a single inverted pendulum in coronal lateral flexion, but requires a more complex representation in sagittal extension. The upper cervical spine has substantial compliance in sagittal extension, and may be responsible for the complex motion and greater extension angular velocities. In analyzing the HICOR, we can gain intuition regarding the neck’s role in dictating head motion during external loading.     

The instantaneous center of rotation of the mandible in nonhuman primates

Kinematic analyses of mandibular movement in humans demonstrate that the mandibular instantaneous center of rotation (ICoR) is commonly located near the level of the occlusal plane and varies in its position during a chewing sequence. Few data are available regarding the location of the ICoR in nonhuman primates and it remains unclear how the position of the ICoR varies in association with mastication and/or gape behaviors. ICoR was quantified throughout the gape cycle in five species of nonhuman primates (Macaca mulatta, Cebus apella, Chlorocebus aethiops, Eulemur fulvus, and Varecia variegata). The ICoR is commonly located below the mandibular condyle close to the occlusal plane and varies considerably both superoinferiorly and anteroposteriorly through the gape cycle. The path of the ICoR, and by inference condylar movement, in Macaca and Chlorocebus differs from humans whereas movement in Cebus resembles that of humans. Similarities between humans and Cebus in articular eminence and occlusal morphology may explain these resemblances. Food material properties had little influence on ICoR movement parameters.     

Evolutionary patterns of shape and functional diversification in the skull and jaw musculature of triggerfishes (Teleostei: Balistidae)

The robust skull and highly subdivided adductor mandibulae muscles of triggerfishes provide an excellent system within which to analyze the evolutionary processes underlying phenotypic diversification. We surveyed the anatomical diversity of balistid jaws using Procrustes‐based geometric morphometric analyses and a phylomorphospace approach to quantifying morphological transformation through evolution. We hypothesized that metrics of interspecific cranial shape would reveal patterns of phylogenetic diversification that are congruent with functional and ecological transformation. Morphological landmarks outlining skull and adductor mandibulae muscle shape were collected from 27 triggerfish species. Procrustes‐transformed skull shape configurations revealed significant phylogenetic and size‐influenced structure. Phylomorphospace plots of cranial shape diversity reveal groupings of shape between different species of triggerfish that are mostly consistent with phylogenetic relatedness. Repeated instances of convergence upon similar cranial shape by genetically disparate taxa are likely due to the functional demands of shared specialized dietary habits. This study shows that the diversification of triggerfish skulls occurs via modifications of cranial silhouette and the positioning of subdivided jaw adductor muscles. Using the morphometric data collected here as input to a biomechanical model of triggerfish jaw function, we find that subdivided jaw adductors, in conjunction with a unique cranial skeleton, have direct biomechanical consequences that are not always congruent with phylomorphospace patterns in the triggerfish lineage. The integration of geometric morphometrics with biomechanical modeling in a phylogenetic context provides novel insight into the evolutionary patterns and ecological role of muscle subdivisions in triggerfishes. J. Morphol. 277:737–752, 2016. © 2016 Wiley Periodicals, Inc.     

Testing amniote models of prey transport kinematics: a quantitative analysis of mouth opening patterns in lizards

Two models have been proposed to describe the prey transport kinematics of terrestrial vertebrates (Bramble and Wake, 1985; Reilly and Lauder, 1990). The critical difference between the models is the presence or absence of a slow open-II phase (SO-II) in the gape profile during mouth opening. Each of these models has been applied to lizards, however to date, lizard feeding kinematics have not been adequately quantified to assess the utility of these models for this clade. Neither model has been sufficiently tested due to the lack of a methodology to assess the specific differences between the models. We describe a method that uses explicit mathematical criteria to define the kinematic phases in tetrapod feeding. This "slope analysis& is used to precisely quantify and compare the transport kinematics of seven lizard species. Lizard transport kinematics were highly variable both within and across taxa. However, several common gape cycle patterns were identified. The predominant patterns were slow-fast opening (37.3%), fast opening only (22.9%) and slow opening only (21.2%). The most common pattern explicitly fits the prediction of the Reilly and Lauder model while the other two are similar to patterns observed in salamanders. Thus, lizards possess both the slow opening-fast opening pattern predicted for amniotes and the more primitive, simple opening pattern characteristic of more basal tetrapods. Plateau phases were found in only 12.8% of the profiles and only a fourth of these (3.4% of the total) explicitly fit the Bramble and Wake model (slow opening, plateau, fast opening) and two species never exhibited plateaus in their gape cycles. Thus, it is clear that the Bramble and Wake model is not supported as a generalized model for lizards or generalized tetrapods.     

Patterns of strain in the macaque ulna during functional activity

In vivo bone strain experiments were performed on the ulnae of three female rhesus macaques to test how the bone deforms during locomotion. The null hypothesis was that, in an animal moving its limbs predominantly in sagittal planes, the ulna experiences anteroposterior bending. Three rosette strain gauges were attached around the circumference of the bone slightly distal to midshaft. They permit a complete characterization of the ulna's loading environment. Strains were recorded during walking and galloping activities. Principal strains and strain directions relative to the long axis of the bone were calculated for each gauge site. In all three animals, the lateral cortex experienced higher tensile than compressive principal strains during the stance phase of walking. Compressive strains predominated at the medial cortex of two animals (the gauge on this cortex of the third animal did not function). The posterior cortex was subject to lower strains; the nature of the strain was highly dependent on precise gauge position. The greater principal strains were aligned closely with the long axis of the bone in two animals, whereas they deviated up to 45° from the long axis in the third animal. A gait change from walk to gallop was recorded for one animal. It was not accompanied by an incremental change in strain magnitudes. Strains are at the low end of the range of strain magnitudes recorded for walking gaits of nonprimate mammals. The measured distribution of strains in the rhesus monkey ulna indicates that mediolateral bending, rather than anteroposterior bending, is the predominant loading regime, with the neutral axis of bending running from anterior and slightly medial to posterior and slightly lateral. A variable degree of torsion was superimposed over this bending regime. Ulnar mediolateral bending is apparently caused by a ground reaction force vector that passes medial to the forearm. The macaque ulna is not reinforced in the plane of bending. The lack of buttressing in the loaded plane and the somewhat counterintuitive bending direction recommend caution with regard to conventional interpretations of long bone cross-sectional geometry. Am J Phys Anthropol 106:87–100, 1998. © 1998 Wiley-Liss, Inc.     

Evolutionary trends and their functional significance in chasmopine trilobites

In response to environmental pressures, speciation within the Chasmopinae involved the attempt to minimise the anterior blind spot. This was attempted in four different ways: by positioning the eyes progressively closer to the glabella, in the Chasmops-Toxochasmops lineage; on the sides of the cephalon, in Bolbochasmops and Rollmops; by elevating and reducing the size of the eyes, in Scopelochasmops ; and by the development of very large eyes in species of Chasmops and Toxochasmops. Development of different eye morphologies resulted in complementary cephalic morphological changes, involving frontal lobe convexity, position and orientation of glabellar furrows, size of hypostome and overall cephalic shape. The strong selection pressure acting on eye form in order to minimise the anterior blind spot suggests that the Chasmopinae may have led a predatory mode of life.     

植物功能型及其生态学意义

植物功能型是近年来在全球变化研究中引入的一个新的生态学术语,综 植物功能型概念的由来、划分植物及其生态学意义,用系统生物学的理论与方法理解植物功能型,并指出了植物功能型在全球变化研究中的意义。     

The hippocampal learning-behavior translation and the functional significance of hippocampal dysfunction in schizophrenia

How is hippocampal learning, including place learning, translated into behavior? The hippocampus integrates, along its septotemporal axis, substrates of rapid place learning, including entorhinal-hippocampal connectivity, with functional connectivity to subcortical sites and prefrontal cortex, which play central roles in behavioral-control functions, including sensorimotor, emotional, motivational, attentional, and executive functions. I present recent evidence that such integration, for which the intermediate hippocampus is a key neuroanatomical substrate, enables translation of rapid place learning into adaptive behavior. What are the clinical implications of the hippocampal learning-behavior translation? Focusing on hippocampal overactivity, which has emerged as a central feature of schizophrenia pathophysiology, I highlight how, due to functional connectivity enabling the learning-behavior translation, hippocampal dysfunction may cause not only memory deficits, but also neural-network disruptions underlying psychosis and attentional and executive deficits.     

Computational model of the movement of the human muscles of mastication during opening and closing of the jaw

Muscle fibre bundles comprising the four major muscles of mastication in the human being were studied in cadavers. Markers were placed along each muscle fibre bundle by means of serial dissections. The 3D coordinates of each marker were tabulated and imported to Cinema 4D, a software animation program. Origins and insertions of each fibre bundle were also digitized and imported, as were the coordinates of the surface of the skull, the mandible and temporomandibular joint. It was then possible to visualize the movement of all relevant fibre bundles during the passive motions of the mandible. An animated film depicts the positions of all relevant muscle fibres during passive movement of the mandible. The properties of the masseter muscle were documented as a prototype for the eventual study of all the muscles of mastication. One can now proceed to study the inverse problem, namely the forces within each fibre bundle that actively generate mandibular motion. It is hoped that these studies will aid in the management of conditions affecting the temporomandibular joint.     

Tibiofemoral kinematics and condylar motion during the stance phase of gait

Accurate knowledge of the dynamic knee motion in-vivo is instrumental for understanding normal and pathological function of the knee joint. However, interpreting motion of the knee joint during gait in other than the sagittal plane remains controversial. In this study, we utilized the dual fluoroscopic imaging technique to investigate the six-degree-of-freedom kinematics and condylar motion of the knee during the stance phase of treadmill gait in eight healthy volunteers at a speed of 0.67 m/s. We hypothesized that the 6DOF knee kinematics measured during gait will be different from those reported for non-weightbearing activities, especially with regards to the phenomenon of femoral rollback. In addition, we hypothesized that motion of the medial femoral condyle in the transverse plane is greater than that of the lateral femoral condyle during the stance phase of treadmill gait. The rotational motion and the anterior–posterior translation of the femur with respect to the tibia showed a clear relationship with the flexion–extension path of the knee during the stance phase. Additionally, we observed that the phenomenon of femoral rollback was reversed, with the femur noted to move posteriorly with extension and anteriorly with flexion. Furthermore, we noted that motion of the medial femoral condyle in the transverse plane was greater than that of the lateral femoral condyle during the stance phase of gait (17.4±2.0 mm vs. 7.4±6.1 mm, respectively; p<0.01). The trend was opposite to what has been observed during non-weightbearing flexion or single-leg lunge in previous studies. These data provide baseline knowledge for the understanding of normal physiology and for the analysis of pathological function of the knee joint during walking. These findings further demonstrate that knee kinematics is activity-dependent and motion patterns of one activity (non-weightbearing flexion or lunge) cannot be generalized to interpret a different one (gait).     

Differences in loading of the temporomandibular joint during opening and closing of the jaw

Kinematics of the human masticatory system during opening and closing of the jaw have been reported widely. Evidence has been provided that the opening and closing movement of the jaw differ from one another. However, different approaches of movement registration yield divergent expectations with regard to a difference in loading of the temporomandibular joint between these movements. Because of these diverging expectations, it was hypothesized that joint loading is equal during opening and closing. This hypothesis was tested by predicting loading of the temporomandibular joint during an unloaded opening and closing movement of the jaw by means of a three-dimensional biomechanical model of the human masticatory system. Model predictions showed that the joint reaction forces were markedly higher during opening than during closing. The predicted opening trace of the centre of the mandibular condyle was located cranially of the closing trace, with a maximum difference between the traces of 0.45 mm. The hypothesis, postulating similarity of joint loading during unloaded opening and closing of the jaw, therefore, was rejected. Sensitivity analysis showed that the reported differences were not affected in a qualitative sense by muscular activation levels, the thickness of the cartilaginous layers within the temporomandibular joint or the gross morphology of the model. Our predictions indicate that the TMJ is loaded more heavily during unloaded jaw opening than during unloaded jaw closing.     

A note on functional implications of morphological variation in the human mandible

This study was carried out on 56 mandibles belonging to skeletal remains recovered from archaeological excavations in Israel dated to 6.000 BP. or less, 2 Neandertal mandibles dating between 50.000–60.000 BP. and 2 early H. sapiens sapiens mandibles both dating to circa 92.000 yr BP. Mandibular body length, the distance from the anterior border of the symphysis to a line bisecting the first molar (distance 1), and the distance from the line bisecting the first molar to the mandibular angle (distance 2) were measured. Distance 1, showed little variation between specimens. However, distance 2 showed a significant difference between sexes and between early and late specimens. For all specimens examined there was a low nonsignificant correlation, between the length of the mandible and distance 1, while there was a high correlation between the length of the mandibular body and distance 2. There was little or no correlation between distance 1 and 2. We propose that the human mandible, as a lever arm, can be divided into two functional parts; an anterior part which shows little change over the last 90000 years, and a posterior part which differs in accordance with the length of the mandibular corpus. These changes in distance 2 appear to correlate to changes in body size and diet, suggesting that as proposed by Hylander (1988) chewing rather than incision has played the main role in evolutionary trends of the hominid mandible. This is also in accordance with mandibular growth during development where the lengthening of the jaw takes place mostly in the posterior part by remodeling in the ramus area (Enlow, 1990) both during individual development (ontogenesis) and through evolutionary changes (phylogenesis).     

The functional morphology of lingual prey capture in a scincid lizard,Tiliqua scincoides (Reptilia: Squamata)

We investigated the functional morphology of lingual prey capture in the blue‐tongued skink, Tiliqua scincoides, a lingual‐feeding lizard nested deep within the family Scincidae, which is presumed to be dominated by jaw‐feeding. We used kinematic analysis of high‐speed video to characterize jaw and tongue movements during prey capture. Phylogenetically informed principal components analysis of tongue morphology showed that, compared to jaw‐feeding scincids and lacertids, T. scincoides and another tongue‐feeding scincid, Corucia zebrata, are distinct in ways suggesting an enhanced ability for hydrostatic shape change. Lingual feeding kinematics show substantial quantitative and qualitative variation among T. scincoides individuals. High‐speed video analysis showed that T. scincoides uses significant hydrostatic elongation and deformation during protrusion, tongue‐prey contact, and retraction. A key feature of lingual prey capture in T. scincoides is extensive hydrostatic deformation to increase the area of tongue‐prey contact, presumably to maximize wet adhesion of the prey item. Adhesion is mechanically reinforced during tongue retraction through formation of a distinctive “saddle” in the foretongue that supports the prey item, reducing the risk of prey loss during retraction.     

Exploring feeding behaviour in deep‐sea dragonfishes (Teleostei: Stomiidae): jaw biomechanics and functional significance of a loosejaw

Deep‐sea dragonfishes (family Stomiidae) possess spectacular morphologies adapted to capturing large prey items in a seascape largely devoid of biomass, including large fang‐like teeth set on extremely long jaws. Perhaps the most intriguing aspect of dragonfish morphology is a lack of a floor to the oral cavity (i.e. there is no skin between the mandibular rami) in species of three dragonfish genera. The present study aimed to investigate the kinematic properties and performance of lower‐jaw adduction in stomiid fishes and to infer what functional advantages or constraints the ‘loosejaw’ confers. A computation model based on dynamic equilibrium predicted very fast jaw adduction for all species at gapes ranging from 90–120° in 66.6–103 ms. Simulations demonstrated that forces resisting lower‐jaw adduction in dragonfishes, and long‐jawed fishes in general, are substantially greater than those in fishes with shorter jaws. These forces constrain inlever length, resulting in relatively high mechanical advantages to attain fast adduction velocities. By reducing the surface area of the lower‐jaw system, loosejaws drastically reduce resistive forces. This has permitted loosejaw dragonfishes to evolve lower mechanical advantages that produce high displacement velocities with an extremely long jaw, a distinct asset in capturing large and scarce resources in the deep‐sea. In addition, loosejaws require a substantially reduced adductor mass to close long jaws at high velocities. These results reveal that the loosejaw condition is an adaptation that expands the morphological boundaries imposed by the dynamic limitations of a long jaw. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 224–240.     

The water vascular system and functional morphology of Paleozoic asteroids

Asteroids of all geologic ages share a single basic body form, surficial skeletal arrangement, and aspects of water vascular construction. In almost all described Paleozoic species, however, either podial pores to the interior of the arm were lacking, or they are directed laterally, above the adambulacrals. They are internal and above the ambulacrals in known post-Paleozoic species and the Pennsylvanian Calliasterella. Certain features of the ambulacral skeletal arrangement also differ. Calliasterella is the closest known Paleozoic relative of post-Paleozoic asteroids. Classifications of asteroids that stress only overall form and surticial skeletal arrangement erroneously include Paleozoic and Holocene species in common ordinal or even lower level groupings. Taxonomic revision is premature: however, most known Paleozoic asteroids represent primitive lineages. Transitional forms allow reconstruction of events leading to the modern arrangement. Ampullar and skeletal arrangements of post-Paleozoic asteroids appear to offer some functional advantages over those of their precursors, but as early as the Ordovician, diverse feeding habits had evolved and ecological roles paralleled those of Holocene species.