Floquet stability analysis of the longitudinal dynamics of two hovering model insects

Because of the periodically varying aerodynamic and inertial forces of the flapping wings, a hovering or constant-speed flying insect is a cyclically forcing system, and, generally, the flight is not in a fixed-point equilibrium, but in a cyclic-motion equilibrium. Current stability theory of insect flight is based on the averaged model and treats the flight as a fixed-point equilibrium. In the present study, we treated the flight as a cyclic-motion equilibrium and used the Floquet theory to analyse the longitudinal stability of insect flight. Two hovering model insects were considered—a dronefly and a hawkmoth. The former had relatively high wingbeat frequency and small wing-mass to body-mass ratio, and hence very small amplitude of body oscillation; while the latter had relatively low wingbeat frequency and large wing-mass to body-mass ratio, and hence relatively large amplitude of body oscillation. For comparison, analysis using the averaged-model theory (fixed-point stability analysis) was also made. Results of both the cyclic-motion stability analysis and the fixed-point stability analysis were tested by numerical simulation using complete equations of motion coupled with the Navier–Stokes equations. The Floquet theory (cyclic-motion stability analysis) agreed well with the simulation for both the model dronefly and the model hawkmoth; but the averaged-model theory gave good results only for the dronefly. Thus, for an insect with relatively large body oscillation at wingbeat frequency, cyclic-motion stability analysis is required, and for their control analysis, the existing well-developed control theories for systems of fixed-point equilibrium are no longer applicable and new methods that take the cyclic variation of the flight dynamics into account are needed.     

Scaling law and enhancement of lift generation of an insect-size hovering flexible wing

We report a comprehensive scaling law and novel lift generation mechanisms relevant to the aerodynamic functions of structural flexibility in insect flight. Using a Navier–Stokes equation solver, fully coupled to a structural dynamics solver, we consider the hovering motion of a wing of insect size, in which the dynamics of fluid–structure interaction leads to passive wing rotation. Lift generated on the flexible wing scales with the relative shape deformation parameter, whereas the optimal lift is obtained when the wing deformation synchronizes with the imposed translation, consistent with previously reported observations for fruit flies and honeybees. Systematic comparisons with rigid wings illustrate that the nonlinear response in wing motion results in a greater peak angle compared with a simple harmonic motion, yielding higher lift. Moreover, the compliant wing streamlines its shape via camber deformation to mitigate the nonlinear lift-degrading wing–wake interaction to further enhance lift. These bioinspired aeroelastic mechanisms can be used in the development of flapping wing micro-robots.     

Active and passive stabilization of body pitch in insect flight

Flying insects have evolved sophisticated sensory–motor systems, and here we argue that such systems are used to keep upright against intrinsic flight instabilities. We describe a theory that predicts the instability growth rate in body pitch from flapping-wing aerodynamics and reveals two ways of achieving balanced flight: active control with sufficiently rapid reactions and passive stabilization with high body drag. By glueing magnets to fruit flies and perturbing their flight using magnetic impulses, we show that these insects employ active control that is indeed fast relative to the instability. Moreover, we find that fruit flies with their control sensors disabled can keep upright if high-drag fibres are also attached to their bodies, an observation consistent with our prediction for the passive stability condition. Finally, we extend this framework to unify the control strategies used by hovering animals and also furnish criteria for achieving pitch stability in flapping-wing robots.     

Analytical model for instantaneous lift and shape deformation of an insect-scale flapping wing in hover

In the analysis of flexible flapping wings of insects, the aerodynamic outcome depends on the combined structural dynamics and unsteady fluid physics. Because the wing shape and hence the resulting effective angle of attack are a priori unknown, predicting aerodynamic performance is challenging. Here, we show that a coupled aerodynamics/structural dynamics model can be established for hovering, based on a linear beam equation with the Morison equation to account for both added mass and aerodynamic damping effects. Lift strongly depends on the instantaneous angle of attack, resulting from passive pitch associated with wing deformation. We show that both instantaneous wing deformation and lift can be predicted in a much simplified framework. Moreover, our analysis suggests that resulting wing kinematics can be explained by the interplay between acceleration-related and aerodynamic damping forces. Interestingly, while both forces combine to create a high angle of attack resulting in high lift around the midstroke, they offset each other for phase control at the end of the stroke.     

Through the eyes of a bird: modelling visually guided obstacle flight

Various flight navigation strategies for birds have been identified at the large spatial scales of migratory and homing behaviours. However, relatively little is known about close-range obstacle negotiation through cluttered environments. To examine obstacle flight guidance, we tracked pigeons (Columba livia) flying through an artificial forest of vertical poles. Interestingly, pigeons adjusted their flight path only approximately 1.5 m from the forest entry, suggesting a reactive mode of path planning. Combining flight trajectories with obstacle pole positions, we reconstructed the visual experience of the pigeons throughout obstacle flights. Assuming proportional–derivative control with a constant delay, we searched the relevant parameter space of steering gains and visuomotor delays that best explained the observed steering. We found that a pigeon''s steering resembles proportional control driven by the error angle between the flight direction and the desired opening, or gap, between obstacles. Using this pigeon steering controller, we simulated obstacle flights and showed that pigeons do not simply steer to the nearest opening in the direction of flight or destination. Pigeons bias their flight direction towards larger visual gaps when making fast steering decisions. The proposed behavioural modelling method converts the obstacle avoidance behaviour into a (piecewise) target-aiming behaviour, which is better defined and understood. This study demonstrates how such an approach decomposes open-loop free-flight behaviours into components that can be independently evaluated.     

Predictions of tumour morphological stability and evaluation against experimental observations

The hallmark of malignant tumours is their spread into neighbouring tissue and metastasis to distant organs, which can lead to life threatening consequences. One of the defining characteristics of aggressive tumours is an unstable morphology, including the formation of invasive fingers and protrusions observed both in vitro and in vivo. In spite of extensive biological, clinical and modelling study and research at physical scales ranging from the molecular to the tissue, the driving dynamics of tumour invasiveness are not completely understood, partly because it is challenging to observe and study cancer as a multi-scale system. Mathematical modelling has been applied to provide further insights into these complex invasive and metastatic behaviours. Modelling a solid tumour as an incompressible fluid, we consider three possible constitutive relations to describe tumour growth, namely Darcy''s law, Stokes'' law and the combined Darcy–Stokes law. We study the tumour morphological stability described by each model and evaluate the consistency between theoretical model predictions and experimental data from in vitro three-dimensional multicellular tumour spheroids. The analysis reveals that the Stokes model is the most consistent with the experimental observations, and that it predicts our experimental tumour growth is marginally stable. We further show that it is feasible to extract parameter values from a limited set of data and create a self-consistent modelling framework that can be extended to the multi-scale study of cancer.     

Which games are growing bacterial populations playing?

Microbial communities display complex population dynamics, both in frequency and absolute density. Evolutionary game theory provides a natural approach to analyse and model this complexity by studying the detailed interactions among players, including competition and conflict, cooperation and coexistence. Classic evolutionary game theory models typically assume constant population size, which often does not hold for microbial populations. Here, we explicitly take into account population growth with frequency-dependent growth parameters, as observed in our experimental system. We study the in vitro population dynamics of the two commensal bacteria (Curvibacter sp. (AEP1.3) and Duganella sp. (C1.2)) that synergistically protect the metazoan host Hydra vulgaris (AEP) from fungal infection. The frequency-dependent, nonlinear growth rates observed in our experiments indicate that the interactions among bacteria in co-culture are beyond the simple case of direct competition or, equivalently, pairwise games. This is in agreement with the synergistic effect of anti-fungal activity observed in vivo. Our analysis provides new insight into the minimal degree of complexity needed to appropriately understand and predict coexistence or extinction events in this kind of microbial community dynamics. Our approach extends the understanding of microbial communities and points to novel experiments.     

Demographic buffering: titrating the effects of birth rate and imperfect immunity on epidemic dynamics

Host demography can alter the dynamics of infectious disease. In the case of perfectly immunizing infections, observations of strong sensitivity to demographic variation have been mechanistically explained through analysis of the susceptible–infected–recovered (SIR) model that assumes lifelong immunity following recovery from infection. When imperfect immunity is incorporated into this framework via the susceptible–infected–recovered–susceptible (SIRS) model, with individuals regaining full susceptibility following recovery, we show that rapid loss of immunity is predicted to buffer populations against the effects of demographic change. However, this buffering is contrary to the dependence on demography recently observed for partially immunizing infections such as rotavirus and respiratory syncytial virus. We show that this discrepancy arises from a key simplification embedded in the SIR(S) framework, namely that the potential for differential immune responses to repeat exposures is ignored. We explore the minimum additional immunological information that must be included to reflect the range of observed dependencies on demography. We show that including partial protection and lower transmission following primary infection is sufficient to capture more realistic reduced levels of buffering, in addition to changes in epidemic timing, across a range of partially and fully immunizing infections. Furthermore, our results identify key variables in this relationship, including R₀.     

Mesh generation and adaptivity for the solution of compressible viscous high speed flows

An implicit–explicit procedure for the solution of the compressible Navier–Stokes equations on unstructured triangular and tetrahedral meshes is outlined. A procedure for constructing continuous lines, made up of edges in the mesh, is employed and the implicit equation system is solved via line relaxation. The problem of generating, and adapting, unstructured meshes for viscous flow simulations is addressed. A number of examples are included which demonstrate the numerical performance of the proposed procedures.     

The radial growth phase of malignant melanoma: multi-phase modelling,numerical simulations and linear stability analysis

Cutaneous melanoma is disproportionately lethal despite its relatively low incidence and its potential for cure in the early stages. The aim of this study is to foster understanding of the role of microstructure on the occurrence of morphological changes in diseased skin during melanoma evolution. The authors propose a biomechanical analysis of its radial growth phase, investigating the role of intercellular/stromal connections on the initial stages of epidermis invasion. The radial growth phase of a primary melanoma is modelled within the multi-phase theory of mixtures, reproducing the mechanical behaviour of the skin layers and of the epidermal–dermal junction. The theoretical analysis takes into account those cellular processes that have been experimentally observed to disrupt homeostasis in normal epidermis. Numerical simulations demonstrate that the loss of adhesiveness of the melanoma cells both to the basal laminae, caused by deregulation mechanisms of adherent junctions, and to adjacent keratynocytes, consequent to a downregulation of E-cadherin, are the fundamental biomechanical features for promoting tumour initiation. Finally, the authors provide the mathematical proof of a long wavelength instability of the tumour front during the early stages of melanoma invasion. These results open the perspective to correlate the early morphology of a growing melanoma with the biomechanical characteristics of its micro-environment.     

Effects of Mesh Motion on the Stability and Convergence of ALE Based Formulations for Moving Boundary Flows

This paper investigates the effects of mesh motion on the stability of fluid-flow equations when written in an Arbitrary Lagrangian–Eulerian frame for solving moving boundary flow problems. Employing the advection-diffusion equation as a model problem we present a mathematical proof of the destabilizing effects induced by an arbitrary mesh motion on the stability and convergence of an otherwise stable scheme. We show that the satisfaction of the so-called geometric conservation laws is essential to the development of an identity that plays a crucial role in establishing stability. We explicitly show that the advection dominated case is susceptible to growth in error because of the motion of the computational grid. To retain the bound on the growth in error, the mesh motion techniques need to account for a domain based constraint that minimizes the relative mesh velocity. Analysis presented in this work can also be extended to the Navier–Stokes equations when written in an ALE frame for FSI problems.     

Embryonic nodal flow and the dynamics of nodal vesicular parcels

We address with fluid-dynamical simulations using direct numerical techniques three important and fundamental questions with respect to fluid flow within the mouse node and left-right development. First, we consider the differences between what is experimentally observed when assessing cilium-induced fluid flow in the mouse node in vitro and what is to be expected in vivo. The distinction is that in vivo, the leftward fluid flow across the mouse node takes place within a closed system and is consequently confined, while this is no longer the case on removing the covering membrane and immersing the embryo in a fluid-filled volume to perform in vitro experiments. Although there is a central leftward flow in both instances, we elucidate some important distinctions about the closed in vivo situation. Second, we model the movement of the newly discovered nodal vesicular parcels (NVPs) across the node and demonstrate that the flow should indeed cause them to accumulate on the left side of the node, as required for symmetry breaking. Third, we discuss the rupture of NVPs. Based on the biophysical properties of these vesicles, we argue that the morphogens they contain are likely not delivered to the surrounding cells by their mechanical rupture either by the cilia or the flow, and rupture must instead be induced by an as yet undiscovered biochemical mechanism.     

Mathematical modelling of atheroma plaque formation and development in coronary arteries

Atherosclerosis is a vascular disease caused by inflammation of the arterial wall, which results in the accumulation of low-density lipoprotein (LDL) cholesterol, monocytes, macrophages and fat-laden foam cells at the place of the inflammation. This process is commonly referred to as plaque formation. The evolution of the atherosclerosis disease, and in particular the influence of wall shear stress on the growth of atherosclerotic plaques, is still a poorly understood phenomenon. This work presents a mathematical model to reproduce atheroma plaque growth in coronary arteries. This model uses the Navier–Stokes equations and Darcy''s law for fluid dynamics, convection–diffusion–reaction equations for modelling the mass balance in the lumen and intima, and the Kedem–Katchalsky equations for the interfacial coupling at membranes, i.e. endothelium. The volume flux and the solute flux across the interface between the fluid and the porous domains are governed by a three-pore model. The main species and substances which play a role in early atherosclerosis development have been considered in the model, i.e. LDL, oxidized LDL, monocytes, macrophages, foam cells, smooth muscle cells, cytokines and collagen. Furthermore, experimental data taken from the literature have been used in order to physiologically determine model parameters. The mathematical model has been implemented in a representative axisymmetric geometrical coronary artery model. The results show that the mathematical model is able to qualitatively capture the atheroma plaque development observed in the intima layer.     

Integration of intrinsic muscle properties,feed-forward and feedback signals for generating and stabilizing hopping

It was hypothesized that a tight integration of feed-forward and feedback-driven muscle activation with the characteristic intrinsic muscle properties is a key feature of locomotion in challenging environments. In this simulation study it was investigated whether a combination of feed-forward and feedback signals improves hopping stability compared with those simulations with one individual type of activation. In a reduced one-dimensional hopping model with a Hill-type muscle (one contractile element, neither serial nor parallel elastic elements), the level of detail of the muscle''s force–length–velocity relation and the type of activation generation (feed-forward, feedback and combination of both) were varied to test their influence on periodic hopping. The stability of the hopping patterns was evaluated by return map analysis. It was found that the combination of feed-forward and proprioceptive feedback improved hopping stability. Furthermore, the nonlinear Hill-type representation of intrinsic muscle properties led to a faster reduction of perturbations than a linear approximation, independent of the type of activation. The results emphasize the ability of organisms to exploit the stabilizing properties of intrinsic muscle characteristics.     

Modelling the dynamics of viral suppressors of RNA silencing

Virus infection in plants is limited by RNA silencing. In turn, viruses can counter RNA silencing with silencing suppressors. Viral suppressors of RNA silencing have been shown to play a role in symptom development in plants. We here study four different strategies employed by silencing suppressors: small interfering RNA (siRNA) binding, double-strand RNA (dsRNA) binding and degrading or inactivating Argonaute. We study the effect of the suppressors on viral accumulation within the cell as well as its spread on a tissue with mathematical and computational models. We find that suppressors which target Argonaute are very effective in a single cell, but that targeting dsRNA or siRNA is much more effective at the tissue level. Although targeting Argonaute can be beneficial for viral spread, it can also cause hindrance in some cases owing to raised levels of siRNAs that can spread to other cells.     

非线性粘弹性桩耦合运动中的混沌分析

研究轴向周期载荷作用下非线性粘弹性桩纵横向耦合运动中的混沌运动。桩体材料满足Leaderman非线性粘弹性本构关系和近似的非线性几何关系，考虑桩体发生纵横向运动的耦合，得到的方程为耦合的非线性偏微分一积分方程；利用Galerkin方法将方程简化并进行数值计算，揭示非线性粘弹性桩的混沌运动和分岔等动力学行为。     

非线性动力吸振器的动力学分析

运用非线性振动的模态方法对非线性动力吸振器进行了动力学分析,研究了该类吸振器的反共振工作点、工作点的稳定性以及阻尼对反共振工作点的影响.     

To delay once or twice: the effect of hypobiosis and free-living stages on the stability of host-parasite interactions.

The life cycle of many endoparasites can be delayed by free-living infective stages and a developmental arrestment in the host referred to as hypobiosis. We investigated the effects of hypobiosis and its interaction with delay in the free-living stages on host-parasite population dynamics by expanding a previous attempt by Dobson & Hudson. When the parasite life cycle does not include free-living stages, hypobiosis destabilizes the host-parasite interactions, irrespective of the assumptions about the regulation of the host population dynamics. Interestingly, the destabilizing effect varies in a nonlinear way with the duration of hypobiosis, the maximal effect being expected for three to five months delay. When the parasite life cycle involves free-living stages, hypobiosis of short or intermediate duration increases the destabilizing effect of the first time delay. However, hypobiosis of a duration of five months or more can stabilize interactions, irrespective of the regulation of the host population dynamics. Overall, we confirmed that hypobiosis is an unusual time delay as it can stabilize a two-way interaction. Contrary to the previous conclusions, such an atypical effect does not require self-regulation of the host population, but instead depends on the existence of free-living stages.     

Hydrodynamics of diatom chains and semiflexible fibres

Diatoms are non-motile, unicellular phytoplankton that have the ability to form colonies in the form of chains. Depending upon the species of diatoms and the linking structures that hold the cells together, these chains can be quite stiff or very flexible. Recently, the bending rigidities of some species of diatom chains have been quantified. In an effort to understand the role of flexibility in nutrient uptake and aggregate formation, we begin by developing a three-dimensional model of the coupled elastic–hydrodynamic system of a diatom chain moving in an incompressible fluid. We find that simple beam theory does a good job of describing diatom chain deformation in a parabolic flow when its ends are tethered, but does not tell the whole story of chain deformations when they are subjected to compressive stresses in shear. While motivated by the fluid dynamics of diatom chains, our computational model of semiflexible fibres illustrates features that apply widely to other systems. The use of an adaptive immersed boundary framework allows us to capture complicated buckling and recovery dynamics of long, semiflexible fibres in shear.