Sauropod haemal arches: morphotypes,new classification and phylogenetic aspects

Sauropod haemal arches are caudal bony structures that have been traditionally incorporated into two different types observed in different anatomical views: Y-shaped (anterior view) and forked (lateral view). This research proposes a classification combining information observed in anterior and lateral views. Four types of ‘Y-shaped’ and six types of ‘V-shaped’ chevrons were recognised. Complete chevron series in some eusauropods allows the comparison of topological equivalent structures along the tail and also among taxa. A basal titanosaur from Argentina exhibits mid-caudal chevron morphology in which more than one cranial and caudal process is present, arising from the distal blades as well as from the proximal rami, a condition not seen before in a sauropod dinosaur. The morphological variability seen in sauropod chevrons along the tail is in close relationship with the development and distribution of muscle. caudofemoralis longus, as seen in extant crocodiles and as previously proposed for non-avian theropods and for caudal centra and transverse processes of sauropod dinosaurs. Two new characters related to middle chevrons are proposed here, in which the transitional morphology is described.     

Spin-label studies of erythrocyte deformability. IV. Relation of electron spin resonance spectral change with deformation and orientation of erythrocytes in shear flow.

Electron spin resonance (ESR) spectra of spin-labeled human erythrocytes in shear flow are simulated to derive semi-empirical relations of the ESR spectral change with deformation and orientation of the cells by using a modified theoretical model developed for deformation and orientation of liquid drops. The six observed spectra at different shear stress values were simultaneously simulated by adjusting only two parameters. One parameter can be related to the ratio of the internal to the external viscosity, and the other to the elastic property of the cell membrane. From these results we have derived a semi-empirical relationship between the average deformation index or the orientation angle with a spectral measure, which characterizes the spectral shape change induced by shear stress. Thus, it becomes possible to obtain improved quantitative information on the rheological behavior of red blood cells by using the spin-label ESR method.     

A model for shear stress-induced deformation of a flow sensor on the surface of vascular endothelial cells

Fluid mechanical shear stress elicits humoral, metabolic, and structural responses in vascular endothelial cells (ECs); however, the mechanisms involved in shear stress sensing and transduction remain incompletely understood. Beyond being responsive to shear stress, ECs distinguish among and respond differently to different types of shear stress. Recent observations suggest that endothelial shear stress sensing may occur through direct interaction of the flow with cell-surface structures that act as primary flow sensors. This paper presents a mathematical model for the shear stress-induced deformation of a flow sensor on the EC surface. The sensor is modeled as a cytoskeleton-coupled viscoelastic structure exhibiting standard linear solid behavior. Since ECs respond differently to different types of flow, the deformation and resulting velocity of the sensor in response to steady, non-reversing pulsatile, and oscillatory flow have been studied. Furthermore, the sensitivity of the results to changes in various model parameters including the magnitude of applied shear stress, the constants that characterize the viscoelastic behavior, and the pulsatile flow frequency (f) has been investigated. The results have demonstrated that in response to a suddenly applied shear stress, the sensor exhibits a level of instantaneous deformation followed by gradual creeping to the long-term response. The peak deformation increases linearly with the magnitude of the applied shear stress and decreases for viscoelastic constants that correspond to stiffer sensors. While the sensor deformation depends on f for low f values, the deformation becomes f -independent above a critical threshold frequency. Finally, the peak sensor deformation is considerably larger for steady and non-reversing pulsatile flow than for oscillatory flow. If the extent of sensor deformation correlates with the intensity of flow-mediated endothelial signaling, then our results suggest possible mechanisms by which ECs distinguish among steady, non-reversing pulsatile, and oscillatory shear stress.     

Mapping the dynamics of shear stress-induced structural changes in endothelial cells

Hemodynamic shear stress regulates endothelial cell biochemical processes that govern cytoskeletal contractility, focal adhesion dynamics, and extracellular matrix (ECM) assembly. Since shear stress causes rapid strain focusing at discrete locations in the cytoskeleton, we hypothesized that shear stress coordinately alters structural dynamics in the cytoskeleton, focal adhesion sites, and ECM on a time scale of minutes. Using multiwavelength four-dimensional fluorescence microscopy, we measured the displacement of rhodamine-fibronectin and green fluorescent protein-labeled actin, vimentin, paxillin, and/or vinculin in aortic endothelial cells before and after onset of steady unidirectional shear stress. In the cytoskeleton, the onset of shear stress increased actin polymerization into lamellipodia, altered the angle of lateral displacement of actin stress fibers and vimentin filaments, and decreased centripetal remodeling of actin stress fibers in subconfluent and confluent cell layers. Shear stress induced the formation of new focal complexes and reduced the centripetal remodeling of focal adhesions in regions of new actin polymerization. The structural dynamics of focal adhesions and the fibronectin matrix varied with cell density. In subconfluent cell layers, shear stress onset decreased the displacement of focal adhesions and fibronectin fibrils. In confluent monolayers, the direction of fibronectin and focal adhesion displacement shifted significantly toward the downstream direction within 1 min after onset of shear stress. These spatially coordinated rapid changes in the structural dynamics of cytoskeleton, focal adhesions, and ECM are consistent with focusing of mechanical stress and/or strain near major sites of shear stress-mediated mechanotransduction.     

Biorheological views of endothelial cell responses to mechanical stimuli

Vascular endothelial cells are located at the innermost layer of the blood vessel wall and are always exposed to three different mechanical forces: shear stress due to blood flow, hydrostatic pressure due to blood pressure and cyclic stretch due to vessel deformation. It is well known that endothelial cells respond to these mechanical forces and change their shapes, cytoskeletal structures and functions. In this review, we would like to mainly focus on the effects of shear stress and hydrostatic pressure on endothelial cell morphology. After applying fluid shear stress, cultured endothelial cells show marked elongation and orientation in the flow direction. In addition, thick stress fibers of actin filaments appear and align along the cell long axis. Thus, endothelial cell morphology is closely related to the cytoskeletal structure. Further, the dynamic course of the morphological changes is shown and the related events such as changes in mechanical stiffness and functions are also summarized. When endothelial cells were exposed to hydrostatic pressure, they exhibited a marked elongation and orientation in a random direction, together with development of centrally located, thick stress fibers. Pressured endothelial cells also exhibited a multilayered structure with less expression of VE-cadherin unlike under control conditions. Simultaneous loading of hydrostatic pressure and shear stress inhibited endothelial cell multilayering and induced elongation and orientation of endothelial cells with well-developed VE-cadherin in a monolayer, which suggests that for a better understanding of vascular endothelial cell responses one has to take into consideration the combination of the different mechanical forces such as exist under in vivo mechanical conditions.     

Microscopic investigation of erythrocyte deformation dynamics

The understanding of erythrocyte deformation under conditions of high shear stress and short exposure time is central to the study of hemorheology and hemolysis within prosthetic blood contacting devices. A combined computational and experimental microscopic study was conducted to investigate the erythrocyte deformation and its relation to transient stress fields. A microfluidic channel system with small channels fabricated using polydimethylsiloxane on the order of 100 mum was designed to generate transient stress fields through which the erythrocytes were forced to flow. The shear stress fields were analyzed by three-dimensional computational fluid dynamics. Microscopic images of deforming erythrocytes were experimentally recorded to obtain the changes in cell morphology over a wide range of fluid dynamic stresses. The erythrocyte elongation index (EI) increased from 0 to 0.54 with increasing shear stress up to 123 Pa. In this shear stress range, erythrocytes behaved like fluid droplets, and deformed and flowed following the surrounding fluid. Cells exposed to shear stress beyond 123 Pa (up to 5170 Pa) did not exhibit additional elongation beyond EI=0.54. Two-stage deformation of erythrocytes in response to shear stress was observed: an initial linear elongation with increasing shear stress and a plateau beyond a critical shear stress.     

Biomechanics of cell rolling: shear flow, cell-surface adhesion, and cell deformability

The mechanics of leukocyte (white blood cell; WBC) deformation and adhesion to endothelial cells (EC) has been investigated using a novel in vitro side-view flow assay. HL-60 cell rolling adhesion to surface-immobilized P-selectin was used to model the WBC-EC adhesion process. Changes in flow shear stress, cell deformability, or substrate ligand strength resulted in significant changes in the characteristic adhesion binding time, cell-surface contact and cell rolling velocity. A 2-D model indicated that cell-substrate contact area under a high wall shear stress (20 dyn/cm2) could be nearly twice of that under a low stress (0.5 dyn/cm2) due to shear flow-induced cell deformation. An increase in contact area resulted in more energy dissipation to both adhesion bonds and viscous cytoplasm, whereas the fluid energy that inputs to a cell decreased due to a flattened cell shape. The model also predicted a plateau of WBC rolling velocity as flow shear stresses further increased. Both experimental and computational studies have described how WBC deformation influences the WBC-EC adhesion process in shear flow.     

Larval morphology of the lesser housefly, Fannia canicularis

The morphology of all larval instars of Fannia canicularis (Linnaeus) (Diptera: Fanniidae) is documented using a combination of light and scanning electron microscopy. The following structures are documented for all instars: antennal complex; maxillary palpus; facial mask; cephaloskeleton; ventral organ; anterior spiracle; Keilin's organ; posterior spiracle; fleshy processes, and anal pad. Structures reported for the first time for all instars include: two pairs of lateral prominences on the prothoracic segment; additional ventrolateral prominences on the second thoracic segment, and a papilla at the base of the posterior spiracle. Other structures reported for the first time are anterior spiracles in the first instar and a serrated tip on the mouthhook in the second instar. A trichoid sensillum on the posterior spiracular plate, representing a sensory organ otherwise unknown in the Calyptratae, is described in the second and third instars. Results are discussed and compared with existing knowledge on dipteran larval morphology.     

Foregut morphology and ontogeny of the spider crab Maja brachydactyla (Brachyura,Majoidea, Majidae)

We describe the morphology of the foregut of the spider crab Maja brachydactyla Balss, 1922, from first larval stage to adult, with detailed stage‐specific documentation using light and scanning electron microscopy. A total of 40 ossicles have been identified in the foregut of adults of M. brachydactyla using Alizarin‐Red staining. The morphological pattern of the ossicles and gastric mill is very similar to other Majoidea species with only a few variations. The foregut of the zoeae stages appeared as a small and simple cavity, with a cardio‐pyloric valve that separates the stomach into cardiac and pyloric regions. The pyloric filter is present from the first zoea, in contrast to the brachyuran species which have an extended larval development. Calcified structures have been identified in the cardio‐pyloric valve and pyloric region of the zoeal stages. The most significant changes in foregut morphology take place after the metamorphosis from ZII to megalopa, including the occurrence of the gastric mill. In the megalopa stage, the foregut ossicles are recognizable by their organization and general morphology, but are different from the adult phase in shape and number. Moreover, the gastric teeth show important differences: the cusps of the lateral teeth are sharp (no molariform); the dorsal tooth have a small, dentate cusp (not a well‐developed quadrangular cusp); and the accessory teeth are composed of one sharp peak (instead of four sharp peaks). The gastric mill ontogeny from megalopa to adult reveals intermediate morphologies during the earlier juvenile stages. The relationship between gastric mill structures with food preferences and their contribution to the brachyuran phylogeny are briefly discussed. J. Morphol. 276:1109–1122, 2015. © 2015 Wiley Periodicals, Inc.     

Mandibular growth and function in Archaeolemur

Ontogenetic changes in the morphology of the mandibular symphysis are described in Archaeolemur so as to infer the functional significance of symphyseal fusion in this subfossil Malagasy lemur. The first regions of the symphysis to show a more complex morphology were the lower and anterior borders of the joint and, to a lesser extent, the lingual borders of the superior and inferior transverse tori. During growth, these regions became increasingly rugose and encroached upon a centrally located, smooth, “oval” region, which may have been a principal pathway for neurovascular structures communicating with the unfused joint. In subadults, the symphysis was completely fused except for the lingual surface of the inferior transverse torus, where a patent suture and potential space were present between dentaries. Thus, in Archaeolemur there was an age- and size-related pattern of increased symphyseal ossification or fusion that was complete by adulthood. The morphology of the interlocking bony processes and the sequence of ossification in the symphysis suggest that increased dorsoventral shear stress during mastication was the most likely determinant of symphyseal fusion in Archaeolemur: The allometric pattern of greater symphyseal fusion may be linked to the presence of relatively greater dorsoventral shear in adults due to an increased recruitment of balancing-side jaw-muscle force. There is little indication that the symphysis of juvenile Archaeolemur was buttressed to resist forces associated with “wishboning” during mastication or vertical bending during incision. Our observations, as well as those of others, suggest that symphyseal fusion in primates occurs initially as a response to increased dorsoventral shear during mastication. Therefore, wishboning stress might only become a major determinant of symphyseal form and function in those taxa that develop a fused symphysis to counter increased dorsoventral shear. © 1994 Wiley-Liss, Inc.     

Osteocyte calcium signaling response to bone matrix deformation

Osteocytes embedded in calcified bone matrix have been widely believed to play important roles in mechanosensing to achieve adaptive bone remodeling in a changing mechanical environment. In vitro studies have clarified several types of mechanical stimuli such as hydrostatic pressure, fluid shear stress, and direct deformation influence osteocyte functions. However, osteocyte response to mechanical stimuli in the bone matrix has not been clearly understood. In this study, we observed the osteocyte calcium signaling response to the quantitatively applied deformation in the bone matrix. A novel experimental system was developed to apply deformation to cultured bone tissue with osteocytes on a microscope stage. As a mechanical stimulus to the osteocytes in bone matrix, in-plane shear deformation was applied using a pair of glass microneedles to bone fragments, obtained from 13-day-old embryonic chick calvariae. Deformation of bone matrix and cells was quantitatively evaluated using an image correlation method by applying for differential interference contrast images of the matrix and fluorescent images of immunolabeled osteocytes, together with imaging of the cellular calcium transient using a ratiometric method. As a result, it was confirmed that the newly developed system enables us to apply deformation to bone matrix and osteocytes successfully under the microscope without significant focal plane shift or deviation from the observation view field. The system could be a basis for further development to investigate the mechanosensing mechanism of osteocytes in bone matrix through examination of various types of rapid biochemical signaling responses and intercellular communication induced by matrix deformation.     

The effects of variable mass and geometry, pretwist, shear deformation and rotatory inertia on the resonant frequencies of intact long bones: A finite element model analysis

The influence of pretwist, nonuniformities in mass and flexural stiffness, rotatory inertia and shear deformation on the natural frequencies of intact bones is evaluated by means of a linear elastic, finite-element model which has been programmed for solution on the digital computer. Theoretical results are compared to the results on the forced vibration of intact canine radii obtained experimentally by Thompson. Surprisingly, inclusion of fairly large pretwist angles (from −14° to 12° for one specimen) had little affect on the first three frequencies of transverse vibration in either the cranial or lateral directions. Inclusion of shear deformation reduced the third-mode frequency in the stiffest (lateral) direction by about six per cent, otherwise shear deformation played a minor role in determining natural frequencies. Similarly, rotatory inertia had negligible influence up to the third natural frequency.

The predominant influence on the first three natural frequencies of transverse vibration could be attributed to the variations in mass and flexural stiffness along the length of the test specimens. Different effective moduli of elasticity are required to yield correct absolute values for the frequencies which correspond to experimental findings, thus implying the presence of some inhomogeneities in material properties around the bone cross-section and/or along its length.     

Heterochrony and tooth evolution in hyperodapedontine rhynchosaurs (Reptilia, Diapsida)

The tooth arrangement of hyperodapedontine rhynchosaurs shows clear patterns of morphological derivation, which can be summarized as three main apomorphic trends: the increase in the number of tooth rows lateral to the main maxillary groove, the loss of dental structures (medial groove and lingual teeth) medial to the main maxillary groove, and the loss of dental structures (medial crest and lingual teeth) medial to the main dentary crest. The analysis of these trends from a heterochronic viewpoint reveals that acceleration was the most probable process involved in the increase in number of the lateral maxillary tooth rows, while the loss of the medial structures of the maxilla and dentary seem to be related respectively to neoteny and post-displacement. Both peramorphic and paedomorphic processes are, therefore, thought to have directed the main modifications seen in the tooth arrangement of the hyperodapedontine rhynchosaurs. Heterochrony plays an important role in the evolution of the Late Triassic rhynchosaurs, which are differentiated mainly on the basis of their dental morphology.     

Shear deformation kinematics during cartilage articulation: effect of lubrication, degeneration, and stress relaxation

During joint articulation, the biomechanical behavior of cartilage not only facilitates load-bearing and low-friction, but also provides regulatory cues to chondrocytes. Elucidation of cartilage kinematics under combined compression and shearing conditions clarifies these cues in health and disease. The objectives of this study were to elucidate the effects of lubricant, tissue degeneration, and stress relaxation duration on cartilage shear kinematics during articulation. Human osteochondral cores with normal and mildly degenerate surface structures were isolated. Paired blocks from each core were apposed, compressed, allowed to stress relax for 5 or 60 min, and shear tested with a micro-scale video microscopy system using phosphate-buffered saline (PBS) or synovial fluid as lubricant. During applied lateral motion, local and overall shear strain (Exz) of articular cartilage were determined. The applied lateral displacement at which Exz reached 50% of the peak (Deltax(1/2)) was also determined. Quantitatively, surface Exz increased at the onset of lateral motion and peaked just as surfaces detached and slid. With continued lateral motion, surface Exz was maintained. After short stress relaxation, effects of lubrication on Exz and Deltax(1/2) were not apparent. With prolonged stress relaxation, Exz and Deltax(1/2) near the articular surface increased markedly when PBS was used as lubricant. Similar patterns were observed for overall Exz and Deltax(1/2). With degeneration, surface Exz was consistently higher for all cases after the onset of lateral motion. Thus, cartilage shear kinematics is markedly affected by lubricant, cartilage degeneration, and loading duration. Changes in these factors may be involved in the pathogenesis of osteoarthritis.     

In vivo shear stress response

EC (endothelial cell) responses to shear stress generated by vascular perfusion play an important role in circulatory homoeostasis, whereas abnormal responses are implicated in vascular diseases such as hypertension and atherosclerosis. ECs subjected to high shear stress in vitro alter their morphology, function and gene expression. The molecular basis for mechanotransduction of a shear stress signal, and the identity of the sensing mechanisms, remain unclear with many candidates under investigation. Translating these findings in vivo has proved difficult. The role of VEGF (vascular endothelial growth factor) flow-dependent nitric oxide release in remodelling skeletal muscle microcirculation is established for elevated (activity, dilatation) and reduced (overload, ischaemia) shear stress, although their temporal relationship to angiogenesis varies. It is clear that growth factor levels may offer only a permissive environment, and alteration of receptor levels may be a viable therapeutic target. Angiogenesis in vivo appears to be a graded phenomenon, and capillary regression on withdrawal of stimulus may be rapid. Combinations of physiological angiogenic stimuli appear not to be additive.     

Cover Image,Volume 116, Number 5, May 2019

Human pluripotent stem cell-derived endothelial cells (hPSC-ECs) present an attractive alternative to primary EC sources for vascular grafting. However, there is a need to mature them towards either an arterial or venous subtype. A vital environmental factor involved in the arteriovenous specification of ECs during early embryonic development is fluid shear stress; therefore, there have been attempts to employ adult arterial shear stress conditions to mature hPSC-ECs. However, hPSC-ECs are naïve to fluid shear stress, and their shear responses are still not well understood. Here, we used a multiplex microfluidic platform to systematically investigate the dose-time shear responses on hPSC-EC morphology and arterial-venous phenotypes over a range of magnitudes coincidental with physiological levels of embryonic and adult vasculatures. The device comprised of six parallel cell culture chambers that were individually linked to flow-setting resistance channels, allowing us to simultaneously apply shear stress ranging from 0.4 to 15 dyne/cm ². We found that hPSC-ECs required up to 40 hr of shear exposure to elicit a stable phenotypic change. Cell alignment was visible at shear stress <1 dyne/cm ², which was independent of shear stress magnitude and duration of exposure. We discovered that the arterial markers NOTCH1 and EphrinB2 exhibited a dose-dependent increase in a similar manner beyond a threshold level of 3.8 dyne/cm ², whereas the venous markers COUP-TFII and EphB4 expression remained relatively constant across different magnitudes. These findings indicated that hPSC-ECs were sensitive to relatively low magnitudes of shear stress, and a critical level of ~4 dyne/cm ² was sufficient to preferentially enhance their maturation into an arterial phenotype for future vascular tissue engineering applications.     

Estructuras de escape y equilibrio asociadas a Conichnus conicus como indicadores de tasas de sedimentación variables en medios litorales tortonienses del SO de España

Numerous and large vertically extensive specimens of Conichnus conicus are described from the Tortonian deposits of the central Guadalquivir Basin (SW Spain). These burrowing structures form a low bioturbation index of the ichnofabric, which allows one to infer the original depositional setting. Both ichnological and sedimentological features suggest a coastal to shallow marine environment dominated by high sediment supply and high-energy sedimentary processes. Conichnus exhibits a characteristic internal fabric that reflects changes in sedimentation rates linked to various processes: (1) initial substrate penetration by the tracemaker; (2) gradual retrusive movement in response to rapid but continuous sediment aggradation; 3) moderate to high rates of retrusive migration as a consequence of rapid to very rapid sediment aggradation and tracemaker escape; and, finally, (4) abandonment of the burrow or burial of the tracemaker subsequently in response to extremely high sediment accumulation rates. Conichnus has been also described from other areas with similar deposits of different ages, where its important value as a paleoenvironmental indicator similarly has been indicated.     

Responses of benthic algal communities and their traits to experimental changes in fine sediments,nutrients and flow

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