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Effect of gravity on the caloric stimulation of the inner ear.

Robert Barany won the 1914 Nobel Prize in medicine for his convection hypothesis for caloric stimulation. Microgravity caloric tests aboard the 1983 SpaceLab 1 mission produced nystagmus results that contradicted the basic premise of Barany's convection theory. In this paper, we present a fluid structural analysis of the caloric stimulation of the lateral semicircular canal. Direct numerical simulations indicate that on earth, natural convection is the dominant mechanism for endolymphatic flow. However, in the microgravity environment of orbiting spacecraft, where buoyancy effects are mitigated, an expansive convection becomes the sole mechanism for producing endolymph motion and cupular displacement. Transient 1 g and microgravity case studies are presented to delineate the different dynamic behaviors of the 1 g and microgravity endolymphatic flows. The associated fluid-structural interactions are also analyzed based on the time evolution of cupular displacements.

Biophysical Phenomena↗

SEE++: a biomechanical model of the oculomotor plant.

The consequences of changes in the oculomotor system on the three-dimensional eye movements are difficult to grasp. Although changes to the rectus muscles can still be approximately understood with simplified geometric models, this approach no longer works with the oblique muscles. It is shown how SEE++, a biomechanical model of the oculomotor plant that was built on the ideas of Miller and Robinson (1984) can improve the understanding of the effects of changes to the oblique eye muscles. By displaying only selected muscles, and by illustrating the relative contribution of these muscles through color-coding the bulb surface, the functional properties of the oblique muscles can be presented in a much clearer way. Investigating the effects of a hyperactive inferior oblique muscle shows that this type of model can help to clarify the functional cause of a pathology, which can otherwise be unclear, even for common pathologies.

Biomechanical Phenomena↗

Basics of cardiac development for the understanding of congenital heart malformations.

Cardiovascular development has become a crucial element of transgene technology in that many transgenic and knockout mice unexpectedly present with a cardiac phenotype, which often turns out to be embryolethal. This demonstrates that formation of the heart and the connecting vessels is essential for the functioning vertebrate organism. The embryonic mesoderm is the source of both the cardiogenic plate, giving rise to the future myocardium as well as the endocardium that will line the system on the inner side. Genetic cascades are unravelled that direct dextral looping and subsequent secondary looping and wedging of the outflow tract of the primitive heart tube. This tube consists of a number of transitional zones and intervening primitive cardiac chambers. After septation and valve formation, the mature two atria and two ventricles still contain elements of the primitive chambers as well as transitional zones. An essential additional element is the contribution of extracardiac cell populations like neural crest cells and epicardium-derived cells. Whereas the neural crest cell is of specific importance for outflow tract septation and formation of the pharyngeal arch arteries, the epicardium-derived cells are essential for proper maturation of the myocardium and coronary vascular formation. Inductive signals, sometimes linked to apoptosis, of the extracardiac cells are thought to be instructive for differentiation of the conduction system. In summary, cardiovascular development is a complex interplay of many cell-cell and cell-matrix interactions. Study of both (transgenic) animal models and human pathology is unravelling the mechanisms underlying congenital cardiac anomalies.

Animals↗

Dendritic processing of excitatory synaptic input in hypothalamic gonadotropin releasing-hormone neurons.

The activity of hypothalamic GnRH neurons results in the intermittent release of GnRH required for reproductive function. This intermittent neurosecretory activity has been proposed to reflect integration of intrinsic properties of and synaptic input to GnRH neurons. Determining the relative impact of synaptic inputs at different locations on the GnRH neuron is difficult, if not impossible, using only experimental approaches. Thus, we used electrophysiological recordings and neuronal reconstructions to generate computer models of GnRH neurons to examine the effects of synaptic inputs at varying distances from the soma along dendrites. The parameters of the models were adjusted to duplicate measured passive and active electrophysiology of cells from mouse brain slices. Our morphological findings reinforce the emerging picture of a complex dendritic structure of GnRH neurons. Furthermore, analysis of reduced morphology models indicated that this population of cells is unlikely to exhibit low-frequency tonic spiking in the absence of synaptic input. Finally, applying realistic patterns of synaptic input to modeled GnRH neurons indicates that synapses located more than about 30% of the average dendrite length from the soma cannot drive firing at frequencies consistent with neuropeptide release. Thus, processing of synaptic input to dendrites of GnRH neurons is probably more complex than simple summation.

Algorithms↗

The receptor-like tyrosine phosphatase lar is required for epithelial planar polarity and for axis determination within drosophila ovarian follicles.

The follicle cell monolayer that encircles each developing Drosophila oocyte contributes actively to egg development and patterning, and also represents a model stem cell-derived epithelium. We have identified mutations in the receptor-like transmembrane tyrosine phosphatase Lar that disorganize follicle formation, block egg chamber elongation and disrupt Oskar localization, which is an indicator of oocyte anterior-posterior polarity. Alterations in actin filament organization correlate with these defects. Actin filaments in the basal follicle cell domain normally become polarized during stage 6 around the anterior-posterior axis defined by the polar cells, but mutations in Lar frequently disrupt polar cell differentiation and actin polarization. Lar function is only needed in somatic cells, and (for Oskar localization) its action is autonomous to posterior follicle cells. Polarity signals may be laid down by these cells within the extracellular matrix (ECM), possibly in the distribution of the candidate Lar ligand Laminin A, and read out at the time Oskar is localized in a Lar-dependent manner. Lar is not required autonomously to polarize somatic cell actin during stages 6. We show that Lar acts somatically early in oogenesis, during follicle formation, and postulate that it functions in germarium intercyst cells that are required for polar cell specification and differentiation. Our studies suggest that positional information can be stored transiently in the ECM. A major function of Lar may be to transduce such signals.

Actins↗

Antennae on transmitters on penguins: balancing energy budgets on the high wire.

The effect of externally mounted antennae on the energetics of penguins was studied by mounting various antennae on a transducer fixed to a model Magellanic penguin Spheniscus magellanicus to determine drag, run at speeds of up to 2 m s(-1) in a swim canal. For rigid antennae set perpendicular to the water flow, measured drag increased with increasing swim speed. Increasing antenna length (for lengths between 100 and 200 mm) or diameter (for diameters between 1 and 4 mm) resulted in accelerating increased drag as a function of both antenna length and diameter. Where antennae were positioned at acute angles to the water flow, drag was markedly reduced, as was drag at higher speeds in flexible antennae. These results were incorporated in a model on the foraging energetics of free-living Magellanic penguins using data (on swim speeds, intervals between prey encounters, amount ingested per patch and dive durations) derived from previously published work and from a field study conducted on birds from a colony at Punta Norte, Argentina, using data loggers. The field work indicated that free-living birds have a foraging efficiency (net energy gain/net energy loss) of about 2.5. The model predicted that birds equipped with the largest rigid external antennae tested (200 mm x 3 mm diameter), set perpendicular to water flow, increased energy expenditure at normal swim speeds of 1.77 m s(-1) by 79% and at prey capture speeds of 2.25 m s(-1) by 147%, and ultimately led to a foraging efficiency that was about 5 times less than that of unequipped birds. Highly flexible antennae were shown to reduce this effect considerably. Deleterious antenna-induced effects are predicted to be particularly critical in penguins that have to travel fast to capture prey. Possible measures taken by the birds to increase foraging efficiency could include reduced travelling speed and selection of smaller prey types. Suggestions are made as to how antenna-induced drag might be minimized for future studies on marine diving animals.

Animals↗

The effect of advance ratio on the aerodynamics of revolving wings.

Recent studies have demonstrated that a quasi-steady model closely matches the instantaneous force produced by an insect wing during hovering flight. It is not clear, however, if such methods extend to forward flight. In this study we use a dynamically scaled robotic model of the fruit fly Drosophila melanogaster to investigate the forces produced by a wing revolving at constant angular velocity while simultaneously translating at velocities appropriate for forward flight. Because the forward and angular velocities were constant wing inertia was negligible, and the measured forces can be attributed to fluid dynamic phenomena. The combined forward and revolving motions of the wing produce a time-dependent free-stream velocity profile, which suggests that added mass forces make a contribution to the measured forces. We find that the forces due added mass make a small, but measurable, component of the total force and are in excellent agreement with theoretical values. Lift and drag coefficients are calculated from the force traces after subtracting the contributions due to added mass. The lift and drag coefficients, for fixed angle of attack, are not constant for non-zero advance ratios, but rather vary in magnitude throughout the stroke. This observation implies that modifications of the quasi-steady model are required in order to predict accurately the instantaneous forces produced during forward flight. We show that the dependence of the lift and drag coefficients upon advance ratio and stroke position can be characterized effectively in terms of the tip velocity ratio--the ratio of the chordwise components of flow velocity at the wing tip due to translation and revolution. On this basis we develop a modified quasi-steady model that can account for the varying magnitudes of the lift and drag coefficients. Our model may also resolve discrepancies in past measurements of wing performance based on translational and revolving motion.

Animals↗

The aerodynamics of hovering flight in Drosophila.

Using 3D infrared high-speed video, we captured the continuous wing and body kinematics of free-flying fruit flies, Drosophila melanogaster, during hovering and slow forward flight. We then 'replayed' the wing kinematics on a dynamically scaled robotic model to measure the aerodynamic forces produced by the wings. Hovering animals generate a U-shaped wing trajectory, in which large drag forces during a downward plunge at the start of each stroke create peak vertical forces. Quasi-steady mechanisms could account for nearly all of the mean measured force required to hover, although temporal discrepancies between instantaneous measured forces and model predictions indicate that unsteady mechanisms also play a significant role. We analyzed the requirements for hovering from an analysis of the time history of forces and moments in all six degrees of freedom. The wing kinematics necessary to generate sufficient lift are highly constrained by the requirement to balance thrust and pitch torque over the stroke cycle. We also compare the wing motion and aerodynamic forces of free and tethered flies. Tethering causes a strong distortion of the stroke pattern that results in a reduction of translational forces and a prominent nose-down pitch moment. The stereotyped distortion under tethered conditions is most likely due to a disruption of sensory feedback. Finally, we calculated flight power based directly on the measurements of wing motion and aerodynamic forces, which yielded a higher estimate of muscle power during free hovering flight than prior estimates based on time-averaged parameters. This discrepancy is mostly due to a two- to threefold underestimate of the mean profile drag coefficient in prior studies. We also compared our values with the predictions of the same time-averaged models using more accurate kinematic and aerodynamic input parameters based on our high-speed videography measurements. In this case, the time-averaged models tended to overestimate flight costs.

Animals↗

A computational study of the aerodynamics and forewing-hindwing interaction of a model dragonfly in forward flight.

The aerodynamics and forewing-hindwing interaction of a model dragonfly in forward flight are studied, using the method of numerically solving the Navier-Stokes equations. Available morphological and stroke-kinematic parameters of dragonfly (Aeshna juncea) are used for the model dragonfly. Six advance ratios (J; ranging from 0 to 0.75) and, at each J, four forewing-hindwing phase angle differences (gamma(d); 180 degrees, 90 degrees, 60 degrees and 0 degree) are considered. The mean vertical force and thrust are made to balance the weight and body-drag, respectively, by adjusting the angles of attack of the wings, so that the flight could better approximate the real flight. At hovering and low J (J=0, 0.15), the model dragonfly uses separated flows or leading-edge vortices (LEV) on both the fore- and hindwing downstrokes; at medium J (J=0.30, 0.45), it uses the LEV on the forewing downstroke and attached flow on the hindwing downstroke; at high J (J=0.6, 0.75), it uses attached flows on both fore- and hindwing downstrokes. (The upstrokes are very lightly loaded and, in general, the flows are attached.) At a given J, at gamma(d)=180 degrees, there are two vertical force peaks in a cycle, one in the first half of the cycle, produced mainly by the hindwing downstroke, and the other in the second half of the cycle, produced mainly by the forewing downstroke; at gamma(d)=90 degrees, 60 degrees and 0 degree, the two force peaks merge into one peak. The vertical force is close to the resultant aerodynamic force [because the thrust (or body-drag) is much smaller than vertical force (or the weight)]. 55-65% of the vertical force is contributed by the drag of the wings. The forewing-hindwing interaction is detrimental to the vertical force (and resultant force) generation. At hovering, the interaction reduces the mean vertical force (and resultant force) by 8-15%, compared with that without interaction; as J increases, the reduction generally decreases (e.g. at J=0.6 and gamma(d)=90 degrees, it becomes 1.6%). A possible reason for the detrimental interaction is as follows: each of the wings produces a mean vertical force coefficient close to half that needed for weight support, and a downward flow is generated in producing the vertical force; thus, in general, a wing moves in the downwash-velocity field induced by the other wing, reducing its aerodynamic forces.

Animals↗

Modeling the electric field of weakly electric fish.

Weakly electric fish characterize the environment in which they live by sensing distortions in their self-generated electric field. These distortions result in electric images forming across their skin. In order to better understand electric field generation and image formation in one particular species of electric fish, Apteronotus leptorhynchus, we have developed three different numerical models of a two-dimensional cross-section of the fish's body and its surroundings. One of these models mimics the real contour of the fish; two other geometrically simple models allow for an independent study of the effects of the fish's body geometry and conductivity on electric field and image formation. Using these models, we show that the fish's tapered body shape is mainly responsible for the smooth, uniform field in the rostral region, where most electroreceptors are located. The fish's narrowing body geometry is also responsible for the relatively large electric potential in the caudal region. Numerical tests also confirm the previous hypothesis that the electric fish body acts approximately like an ideal voltage divider; this is true especially for the tail region. Next, we calculate electric images produced by simple objects and find they vary according to the current density profile assigned to the fish's electric organ. This explains some of the qualitative differences previously reported for different modeling approaches. The variation of the electric image's shape as a function of different object locations is explained in terms of the fish's geometrical and electrical parameters. Lastly, we discuss novel cues for determining an object's rostro-caudal location and lateral distance using these electric images.

Animals↗

Kinematic models of the buccal mass of Aplysia californica.

The feeding behavior of the marine mollusc Aplysia californica is an intensively studied model system for understanding the neural control of behavior. Feeding movements are generated by contractions of the muscles of the buccal mass. These muscles are internal and cannot be visualized during behavior. In order to infer the movements of the muscles of the buccal mass, two kinematic models were constructed. The first kinematic model assumed that the complex consisting of the pincer-like radula and the underlying odontophore was spherical in shape. In this model, the radula/odontophore was moved anteriorly or posteriorly and the more superficial buccal muscles (I1/I3 and I2) were fitted around it. Although the overall buccal mass shapes predicted by this model were similar to those observed in vivo during protraction, the shapes predicted during retraction were very different. We therefore constructed a second kinematic model in which the shape of the radula/odontophore was based on the shapes assumed by those structures in vitro when they were passively forced into protraction, rest or retraction positions. As each of these shapes was rotated, the second kinematic model generated overall shapes of the buccal mass that were similar to those observed in vivo during swallowing and tearing, and made predictions about the antero-posterior length of the buccal mass and the relative location of the lateral groove. These predictions were consistent with observations made in vivo and in vitro. The kinematic patterns of intrinsic buccal muscles I1 and I2 in vivo were estimated using the second model. Both models make testable predictions with regard to the functions and neural control of intrinsic buccal muscles I2 and I3.

Animals↗

Hydrodynamic drag of diving birds: effects of body size, body shape and feathers at steady speeds.

For birds diving to depths where pressure has mostly reduced the buoyancy of air spaces, hydrodynamic drag is the main mechanical cost of steady swimming. Drag is strongly affected by body size and shape, so such differences among species should affect energy costs. Because flow around the body is complicated by the roughness and vibration of feathers, feathers must be considered in evaluating the effects of size and shape on drag. We investigated the effects of size, shape and feathers on the drag of avian divers ranging from wing-propelled auklets weighing 75 g to foot-propelled eiders weighing up to 2060 g. Laser scanning of body surfaces yielded digitized shapes that were averaged over several specimens per species and then used by a milling machine to cut foam models. These models were fitted with casts of the bill area, and their drag was compared with that of frozen specimens. Because of the roughness and vibration of the feathers, the drag of the frozen birds was 2-6 times that of the models. Plots of drag coefficient (C(D)) versus Reynolds number (Re) differed between the model and the frozen birds, with the pattern of difference varying with body shape. Thus, the drag of cast models or similar featherless shapes can differ both quantitatively and qualitatively from that of real birds. On the basis of a new towing method with no posts or stings that alter flow or angles of attack, the dimensionless C(D)/Re curves differed among a size gradient of five auklet species (75-100g) with similar shapes. Thus, extrapolation of C(D)/Re curves among related species must be performed with caution. At lower speeds, the C(D) at a given Re was generally higher for long-necked birds that swim with their neck extended (cormorants, grebes, some ducks) than for birds that swim with their head retracted (penguins, alcids), but this trend was reversed at high speeds. Because swimming birds actually travel at a range of instantaneous speeds during oscillatory strokes, species variations in drag at different speeds must be considered in the context of accelerational stroking.

Animals↗

Injection-moulded models of major and minor arteries: the variability of model wall thickness owing to casting technique.

Cardiovascular disease of major and minor arteries is a common cause of death in Western society. The wall mechanics and haemodynamics within the arteries are considered to be important factors in the disease formation process. This paper is concerned with the development of an efficient computer-integrated technique to manufacture idealized and realistic models of diseased major and minor arteries from radiological images and to address the issue of model wall thickness variability. Variations in wall thickness from the original computer models to the final castings are quantified using a CCD camera. The results found that wall thickness variation from the major and minor idealized artery models to design specification were insignificant, up to a maximum of 16 per cent. In realistic models, however, differences were up to 23 per cent in the major arterial models and 58 per cent in the minor arterial models, but the wall thickness variability remained within the limits of previously reported wall thickness results. It is concluded that the described injection moulding procedure yields idealized and realistic castings suitable for use in experimental investigations, with idealized models giving better agreement with design. Wall thickness is variable and should be assessed after the models are manufactured.

Anatomy, Cross-Sectional↗

Non-linear finite element analysis of formation and treatment of intervertebral disc herniae.

A biomechanical model of the spine motion segment L2/L3 consisting of the truncated vertebrae, endplates, intervertebral disc and pieces of anterior and posterior longitudinal ligaments has been used for a computer simulation study. A non-linear finite element analysis has shown that small loads compressing the spine, not greater than those occurring in everyday life, cause loss of stability of an intervertebral disc, resulting in lateral dislocation of its nucleus pulposus. This could be a potential cause of discopathy. The model indicates that conservative therapy of herniated disc by the traction method may result in retraction of hernia by about 40 per cent.

Biomechanical Phenomena↗

Asperity lubrication in human joints.

The asperity lubrication in human joints is examined in the present paper, with particular reference to the tertiary undulation with wavelengths of around 20-45 microns. It was found that, under dynamic physiological loading conditions, the secondary waviness of the cartilaginous surface (typically 0.5 mm wavelength) could be effectively flattened to sustain a fluid film of 0.1-0.3 micron thick, while the tertiary waviness could be squashed to sustain a much thinner fluid film of 0.01 micron (10 nm) thick with normal synovial fluid as the lubricant. The calculated film thickness for the tertiary undulation was less than 5 nm when the ankle joint was lubricated by Ringer's solution or pathological synovial fluids, or when only quasi-static loading conditions were considered, while a sufficiently thick fluid film could still be formed when the secondary undulations were considered alone. It was thus suggested that the fluid film lubrication mechanism was operative for human joints with normal synovial fluid as the lubricant under physiological dynamic loading conditions and the mixed lubrication mechanism could take over when static loading conditions prevailed or when watery lubricants (eta approximately 0.001 Pa s) were used.

Ankle Joint↗

An experimental investigation of the resolution of tympanometry for micro-perforations of the tympanic membrane.

Tympanometry is well established as a useful tool in the field of otological diagnosis. There have been sporadic reports, however, of bizarre tympanograms in the presence of small perforations of the tympanic membrane. A model of the outer and middle ears was fabricated which was able to produce normal and pathological tympanograms. This model was used to investigate tympanometry in the presence of microperforations. A hypothesis for the mechanism of generation of sawtooth tympanograms in the presence of micro-perforations is postulated.

Acoustic Impedance Tests↗

Can shaking alone cause fatal brain injury? A biomechanical assessment of the Duhaime shaken baby syndrome model.

A biomechanical model of a one-month old baby was designed and tested by Duhaime and co-workers in 1987 in an attempt to assess the biomechanics of the shaken baby syndrome (SBS). The study implied that pure shaking alone cannot cause fatal head injuries, a factor which has been applied in criminal courts. In an attempt to test the validity of the model a preliminary study was undertaken in which a replica was constructed and tested. The broad description of the design and construction of the Duhaime model allowed for variations and therefore uncertainties in its reproduction. It was postulated therefore that differences in certain parameters may increase angular head accelerations. To further investigate this observation, an adjustable replica model was developed and tested. The results indicated that certain parameter changes in the model did in fact lead to an increase in angular head acceleration. When these parameter changes were combined and an injurious shake pattern was employed, using maximum physical effort, the angular head acceleration results exceeded the original Duhaime et al. (1987) results and spanned two scaled tolerance limits for concussion. Additionally, literature suggests that the tolerance limits used to assess the shaking simulation results in the original study may not be reliable. Results from our study were closer to the internal head injury, subdural haematoma, tolerance limits. A series of end point impacts were identified in the shake cycles, therefore, an impact-based head injury measure (Head Injury Criterion - HIC) was utilized to assess their severity. Seven out of ten tests conducted resulted in HIC values exceeding the tolerance limits (critical load value, Stürtz, 1980) suggested for children. At this present stage the authors conclude that it cannot be categorically stated, from a biomechanical perspective, that pure shaking cannot cause fatal head injuries in an infant. Parameters identified in this study require further investigation to assess the accuracy of simulation and increase the biofidelity of the models before further conclusions can be drawn. There must now be sufficient doubt in the reliability of the Duhaime et al. (1987) biomechanical study to warrant the exclusion of such testimony in cases of suspected shaken baby syndrome.

Biomechanical Phenomena↗