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T L Daniel

Publications and source records attributed to T L Daniel.

18 recordsLinked to original sources

Flight control in the hawkmoth Manduca sexta: the inverse problem of hovering.

The inverse problem of hovering flight, that is, the range of wing movements appropriate for sustained flight at a fixed position and orientation, was examined by developing a simulation of the hawkmoth Manduca sexta. Inverse problems arise when one is seeking the parameters that are required to achieve a specified model outcome. In contrast, forward problems explore the outcomes given a specified set of input parameters. The simulation was coupled to a microgenetic algorithm that found specific sequences of wing and body motions, encoded by ten independent kinematic parameters, capable of generating the fixed body position and orientation characteristic of hovering flight. Additionally, we explored the consequences of restricting the number of free kinematic parameters and used this information to assess the importance to flight control of individual parameters and various combinations of them. Output from the simulated moth was compared to kinematic recordings of hovering flight in real hawkmoths; the real and simulated moths performed similarly with respect to their range of variation in position and orientation. The simulated moth also used average wingbeat kinematics (amplitude, stroke plane orientation, etc) similar to those of the real moths. However, many different subsets of the available kinematic were sufficient for hovering flight and available kinematic data from real moths does not include sufficient detail to assess which, if any, of these was consistent with the real moth. This general result, the multiplicity of possible hovering kinematics, shows that the means by which Manduca sexta actually maintains position and orientation may have considerable freedom and therefore may be influenced by many other factors beyond the physical and aerodynamic requirements of hovering flight.

Algorithms↗

Into thin air: Contributions of aerodynamic and inertial-elastic forces to wing bending in the hawkmoth Manduca sexta.

During flapping flight, insect wings must withstand not only fluid-dynamic forces, but also inertial-elastic forces generated by the rapid acceleration and deceleration of their own mass. Estimates of overall aerodynamic and inertial forces vary widely, and the relative importance of these forces in determining passive wing deformations remains unknown. If aeroelastic interactions between a wing and the fluid-dynamic forces it generates are minor compared to the effects of wing inertia, models of insect flight that account for passive wing flexibility would be far simpler to develop. We used an experimental approach to examine the contributions of aerodynamic and inertial-elastic forces to wing bending in the hawkmoth Manduca sexta. We attached fresh Manduca wings to a motor and flapped them at a realistic wing-beat frequency and stroke amplitude. We compared wing bending in normal air versus helium (approx. 15% air density), in which the contribution of fluid-dynamic forces to wing deformations is significantly reduced. This 85% reduction in air density produced only slight changes in the pattern of Manduca wing deformations, suggesting that fluid-dynamic forces have a minimal effect on wing bending. We used a simplified finite element model of a wing to show that the differences observed between wings flapped in air versus helium are most likely due to fluid damping, rather than to aerodynamic forces. This suggests that damped finite element models of insect wings (with no fluid-dynamic forces included) may be able to predict overall patterns of wing deformation prior to calculations of aerodynamic force production, facilitating integrative models of insect flight.

Animals↗

Flexural stiffness in insect wings. I. Scaling and the influence of wing venation.

During flight, many insect wings undergo dramatic deformations that are controlled largely by the architecture of the wing. The pattern of supporting veins in wings varies widely among insect orders and families, but the functional significance of phylogenetic trends in wing venation remains unknown, and measurements of the mechanical properties of wings are rare. In this study, we address the relationship between venation pattern and wing flexibility by measuring the flexural stiffness of wings (in both the spanwise and chordwise directions) and quantifying wing venation in 16 insect species from six orders. These measurements show that spanwise flexural stiffness scales strongly with the cube of wing span, whereas chordwise flexural stiffness scales with the square of chord length. Wing size accounts for over 95% of the variability in measured flexural stiffness; the residuals of this relationship are small and uncorrelated with standardized independent contrasts of wing venation characters. In all species tested, spanwise flexural stiffness is 1-2 orders of magnitude larger than chordwise flexural stiffness. A finite element model of an insect wing demonstrates that leading edge veins are crucial in generating this spanwise-chordwise anisotropy.

Animals↗

Flexural stiffness in insect wings. II. Spatial distribution and dynamic wing bending.

The dynamic, three-dimensional shape of flapping insect wings may influence many aspects of flight performance. Insect wing deformations during flight are largely passive, and are controlled primarily by the architecture and material properties of the wing. Although many details of wing structure are well understood, the distribution of flexural stiffness in insect wings and its effects on wing bending are unknown. In this study, we developed a method of estimating spatial variation in flexural stiffness in both the spanwise and chordwise direction of insect wings. We measured displacement along the wing in response to a point force, and modeled flexural stiffness variation as a simple mathematical function capable of approximating this measured displacement. We used this method to estimate flexural stiffness variation in the hawkmoth Manduca sexta, and the dragonfly Aeshna multicolor. In both species, flexural stiffness declines sharply from the wing base to the tip, and from the leading edge to the trailing edge; this variation can be approximated by an exponential decline. The wings of M. sexta also display dorsal/ventral asymmetry in flexural stiffness and significant differences between males and females. Finite element models based on M. sexta forewings demonstrate that the measured spatial variation in flexural stiffness preserves rigidity in proximal regions of the wing, while transferring bending to the edges, where aerodynamic force production is most sensitive to subtle changes in shape.

Animals↗

Shape, flapping and flexion: wing and fin design for forward flight.

Both kinematics and morphology are critical determinants of performance in flapping flight. However, the functional consequences of changes in these traits are not yet well understood. Traditional aerodynamic studies of planform wing shape have suggested that high-aspect-ratio wings generate more force per area and perform more efficiently than low-aspect-ratio wings, but these analyses may neglect critical components of flapping flight such as unsteady fluid dynamics and wing or fin flexion. In this paper, we use an unsteady potential flow analysis that incorporates wing flexion to test predictions of optimal wing shape under varying degrees of unsteady motion and wing flexion. We focus on forward flapping flight and examine the effects of wing/fin morphology and movements on thrust generation and efficiency. We test the model by comparing our predictions with kinematic data derived from the aquatic flight of the ratfish Hydrolagus colliei. Our analyses show that aspect ratio and the proportion of area in the outer one-fifth of the wing can characterize wing shape in terms of aero- or hydrodynamic performance. By comparing the performance of wings that vary in these two parameters, we find that traditional predictions of optimal wing shape are valid only under limited circumstances (when flapping frequency is low, wings are stiff or wings are tapered at the tips). This indicates a complex relationship between locomotor traits and performance and helps explain the diversity of wing kinematics and morphologies observed in nature.

Animals↗

Viscosity and solute dependence of F-actin translocation by rabbit skeletal heavy meromyosin.

We tested the hypothesis that solvent viscosity affects translocation of rhodamine phalloidin-labeled F-actin by rabbit skeletal heavy meromyosin (HMM). When viscosity was increased using either glycerol, fructose, sucrose, or dextran (1.5, 6.0, or 15-20 kDa mol mass), there was little or no effect on the fraction of moving filaments, whereas sliding speed decreased in inverse proportion to viscosity. The results could be explained neither by an effect of osmotic pressure at high solute concentrations nor by altered solvent drag on the actin filament. Elevated viscosity inhibited HMM ATPase activity in solution, but only at much higher viscosities than were needed to reduce sliding speed. Polyethylene glycols (300, 1,000, or 3,000 mol wt) also inhibited speed via elevated viscosity but secondarily inhibited by enhancing electrostatic interactions. These results demonstrate that a diffusion-controlled process intrinsic to cross-bridge cycling can be limiting to actomyosin function.

Actins↗

Animal movement, mechanical tuning and coupled systems.

Over the past two decades, there has been a growing interest in developing predictive models of animal movement and force generation in fluids. In a departure from past studies that have asked how prescribed motions of a propulsor (wing or fin) generate lift and thrust during swimming and flying, we are increasingly interested in predicting the propulsor's movement as well as the forces generated by it. This interest, motivated by a need to understand the control and dynamics of locomotion and its applications to robotics and animal physiology, requires that we develop integrative models and analyses of swimming and flying that incorporate neural control and muscle physiology into more traditional biomechanical studies of locomotion in fluids. This approach extends from whole-animal studies to the molecular basis of force generation. In this paper, we explore mechanical tuning from the level of the whole animal to the proteins driving force generation in muscle.

Animals↗

Compliant realignment of binding sites in muscle: transient behavior and mechanical tuning.

The presence of compliance in the lattice of filaments in muscle raises a number of concerns about how one accounts for force generation in the context of the cross-bridge cycle--binding site motions and coupling between cross-bridges confound more traditional analyses. To explore these issues, we developed a spatially explicit, mechanochemical model of skeletal muscle contraction. With a simple three-state model of the cross-bridge cycle, we used a Monte Carlo simulation to compute the instantaneous balance of forces throughout the filament lattice, accounting for both thin and thick filament distortions in response to cross-bridge forces. This approach is compared to more traditional mass action kinetic models (in the form of coupled partial differential equations) that assume filament inextensibility. We also monitored instantaneous force generation, ATP utilization, and the dynamics of the cross-bridge cycle in simulations of step changes in length and variations in shortening velocity. Three critical results emerge from our analyses: 1) there is a significant realignment of actin-binding sites in response to cross-bridge forces, 2) this realignment recruits additional cross-bridge binding, and 3) we predict mechanical behaviors that are consistent with experimental results for velocity and length transients. Binding site realignment depends on the relative compliance of the filament lattice and cross-bridges, and within the measured range of these parameters, gives rise to a sharply tuned peak for force generation. Such mechanical tuning at the molecular level is the result of mechanical coupling between individual cross-bridges, mediated by thick filament deformations, and the resultant realignment of binding sites on the thin filament.

Actins↗

Invertebrate swimming: integrating internal and external mechanics.

Challenges in understanding design for locomotion in any swimming animal revolve around the complex interactions between the mechanics of fluid motions around an animal and the mechanics of internal force production. The former is governed by force-velocity relationships that arise from fluid motion, whereas the latter is governed by force-velocity relationships of both active (e.g. muscle) and passive tissues. In reality, all such relationships must be satisfied simultaneously in any swimming creature. Towards this end, I combine traditional analyses of muscle contractility, soft tissue mechanics and hydrodynamic models to examine how organism morphology, muscle physiology and mode of propulsion interact to affect swimming performance. This combination involves a solution to a set of equations that describe the relevant internal and external mechanics. The solution has the unique benefit of providing predictions for both body and propulsor kinematics. Such predictions are examined for a variety swimming animals and are compared with existing kinematic data. Armed with such an approach, I re-examine traditional scaling arguments to show that classical approaches that neglect internal mechanics predict swimming performance relationships that may not be physiologically feasible.

Animals↗

Myotendinous junctions of tonic muscle cells: structure and loading.

Regions within frog semitendinosus muscle that are rich in tonic muscle cells were identified histochemically by myosin adenosine triphosphatase- and succinic dehydrogenase-staining procedures. Bundles of cells still attached to tendinous insertions were removed from those sites, prepared for electron microscopy and sectioned longitudinally through their myotendinous junctions. Tonic cells were identified by electron-microscopic criteria and their myotendinous junctions' morphology evaluated by morphometry. Although junctional components appear identical to those in twitch cells, the degree of membrane folding increases tonic junction area by a factor of 50.2 whereas twitch cells' junctional area is increased 22.2 times by folding relative to cells terminating as right circular cylinders. Calculations show that the tonic cell junction bears average loads of 3.4 X 10(3) N X m-2 during maximum force generation and that nearly all of the load is borne as shear stress at the junction. The junctions of twitch cells bear average loads of 1.6 X 10(4) N X m-2 during peak tension. The findings indicate that the magnitude of loading does not alone determine the degree of junctional membrane folding. Interpretation of the data in view of viscoelastic behavior of membranes indicates that duration of loading may be a functionally important correlate to degree of membrane folding at myotendinous junctions.

Animals↗

Elastic energy storage in rigored skeletal muscle cells under physiological loading conditions.

The capability of heavy meromyosin (HMM) to store energy in reversible deformations has been investigated previously; yet, whether HMM is the site of most elastic energy storage in skeletal muscle cells has not been established. We conducted dynamic loading tests on single rigored muscle cells over the physiological range of sarcomere lengths. These tests enabled us to calculate the energy stored in reversible deformations or dissipated in the cell during each cycle of oscillation. Our findings show that these cells are capable of storing approximately 0.5 J . kg-1 of elastic energy during the last 50 ms of passive extension in vivo by agonists and before their own active contraction. Possible sites of this energy storage are HMM subunit 2, the proximal portion of HMM subunit 1, and parallel structures. However, energy storage increases monotonically as myofilament overlap decreases in the physiological range. This negative correlation suggests that HMM subunits are not the primary sites of elastic energy storage. Our electron-microscopic observations show that collagen fibrils at the cell's surface become oriented parallel to the cell's long axis over the range of sarcomere lengths where energy storage increases. This provides a mechanism for the observed increases in elastic energy storage.

Animals↗

Swyer-James syndrome--unilateral hyperlucent lung syndrome. A case report and review.

Swyer-James syndrome is a pulmonary condition acquired following bronchiolitis obliterans early in life. Clinically characterized by repeated bouts of pulmonary infections, Swyer-James syndrome is characterized radiographically by a unilateral small, hyperlucent lung that demonstrates diminished arterial supply to the involved lung, air-trapping, and bronchiectasis. Radioimaging procedures may reveal otherwise unsuspected bilateral involvement. A 14-year-old Caucasian female with classic findings of Swyer-James syndrome is presented. The differential diagnosis of conditions that may present with a unilateral hyperlucent lung is discussed. Therapy is aimed primarily at control of the intercurrent episodes of bacterial pneumonia.

Adolescent↗

Feeding strategy and the mechanics of blood sucking in insects.

As a means of exploring foraging strategies of blood-feeding insects, we studied the mechanics of blood feeding. We develop a mechanistic model for the dynamics of non-Newtonian fluid flow to describe the feeding process for blood feeders. Using available feeding and morphological data, we examine the relationship of feeding time to proboscis design, and consider optimal foraging strategies for blood feeders. Because of the flow rates typical of many blood feeders, the non-Newtonian nature of blood is of little importance for flow dynamics. Observed feeding times and flow rates do not necessarily reflect the energy requirements for feeding. The radius of the food canal is the major morphological determinant of flow dynamics. Feeding time is a monotonically increasing function of blood hematocrit. There is an optimal blood hematocrit of 0.3 which maximizes the rate of total protein intake for blood feeders, regardless of the energy output or proboscis design. This hematocrit level is typical of humans with blood parasite infections. In contrast, the rate of red blood cell intake is maximized at a hematocrit of 0.4. We argue that the existence of such optima may be a general consequence of the mechanics of feeding on nutrients dissolved or suspended in a fluid medium. Results are discussed in relation to foraging strategy, proboscis design, and the coevolution among host, vector, and parasite in blood feeding insects.

Aedes↗

Evaluation of traumatic atlantooccipital dislocations.

The diagnosis of traumatic atlantooccipital dislocation (AOD) from the cross-table lateral radiograph is difficult because of problems in demonstrating the complex anatomy of this area and the intricate radiographic methods used to diagnose AOD. Although CT or polytomography seem to be the most accurate diagnostic methods, it is often the lateral radiograph from which the diagnosis and further decisions are made. To determine both the best radiographic method for diagnosing AOD from the lateral radiograph and the role of CT and tomography in the diagnosis of AOD, the literature was reviewed concerning how the diagnosis of AOD has been obtained. In addition, the Wholey dens-basion line, the Powers ratio, the Dublin method of diagnosing AOD, and measurement of the atlantooccipital joint width were applied to 12 cases of traumatic AOD; and the Wholey dens-basion line and the Powers ratio were determined in 100 normal adults and 50 normal children. An alternative plain radiographic method for diagnosing AOD was developed, called the X-line method. This was the most accurate of the methods tested, correctly diagnosing AOD in 75% of cases. The Wholey dens-basion line and direct measurement of the atlantooccipital joint width were each correct in 50% of cases, the Powers ratio in 33% of cases, and the Dublin method in only 25% of cases. Ultimately, either CT or polytomography should provide the definitive diagnosis. In this regard high-resolution CT with reformatted coronal and sagittal images generated from 2-mm thin axial slices appeared to have the most promise as the first study of choice.

Adult↗