Experiments and theory of undulatory locomotion in a simple structured medium

Undulatory locomotion of micro-organisms through geometrically complex, fluidic environments is ubiquitous in nature and requires the organism to negotiate both hydrodynamic effects and geometrical constraints. To understand locomotion through such media, we experimentally investigate swimming of the nematode Caenorhabditis elegans through fluid-filled arrays of micro-pillars and conduct numerical simulations based on a mechanical model of the worm that incorporates hydrodynamic and contact interactions with the lattice. We show that the nematode''s path, speed and gait are significantly altered by the presence of the obstacles and depend strongly on lattice spacing. These changes and their dependence on lattice spacing are captured, both qualitatively and quantitatively, by our purely mechanical model. Using the model, we demonstrate that purely mechanical interactions between the swimmer and obstacles can produce complex trajectories, gait changes and velocity fluctuations, yielding some of the life-like dynamics exhibited by the real nematode. Our results show that mechanics, rather than biological sensing and behaviour, can explain some of the observed changes in the worm''s locomotory dynamics.     

A hydrodynamic mechanism for attraction of undulatory microswimmers to surfaces (bordertaxis)

Although small nematodes significantly impact human and animal health, agriculture, and ecology, little is known about the role of hydrodynamics in their life cycles. Using the nematode Caenorhabditis elegans as a model undulatory microswimmer, we have observed that animals are attracted to and swim along surfaces. The attraction to surfaces does not require mechanosensory neuron function. In dilute swarms, swimmers aggregate near surfaces. Using resistive force-based theory, symmetry arguments, and direct hydrodynamic simulations, we demonstrate that forces resulting from the interaction between the swimmer-induced flow field and a nearby surface cause a short-range hydrodynamic torque that stirs the swimmers towards the surface. When combined with steric forces, this causes locomotion along the surface. This surface attraction may affect nematode mate and food finding behaviour and, in the case of parasitic nematodes, may facilitate host penetration. Surface attraction must be accounted for when studying animals'' responses to various stimuli, and suggests means of controlling undulatory microswimmers.     

Remote automated multi-generational growth and observation of an animal in low Earth orbit

The ultimate survival of humanity is dependent upon colonization of other planetary bodies. Key challenges to such habitation are (patho)physiologic changes induced by known, and unknown, factors associated with long-duration and distance space exploration. However, we currently lack biological models for detecting and studying these changes. Here, we use a remote automated culture system to successfully grow an animal in low Earth orbit for six months. Our observations, over 12 generations, demonstrate that the multi-cellular soil worm Caenorhabditis elegans develops from egg to adulthood and produces progeny with identical timings in space as on the Earth. Additionally, these animals display normal rates of movement when fully fed, comparable declines in movement when starved, and appropriate growth arrest upon starvation and recovery upon re-feeding. These observations establish C. elegans as a biological model that can be used to detect changes in animal growth, development, reproduction and behaviour in response to environmental conditions during long-duration spaceflight. This experimental system is ready to be incorporated on future, unmanned interplanetary missions and could be used to study cost-effectively the effects of such missions on these biological processes and the efficacy of new life support systems and radiation shielding technologies.     

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.     

Fractal analysis of behaviour in a wild primate: behavioural complexity in health and disease

Parasitism and other stressors are ubiquitous in nature but their effects on animal behaviour can be difficult to identify. We investigated the effects of nematode parasitism and other indicators of physiological impairment on the sequential complexity of foraging and locomotion behaviour among wild Japanese macaques (Macaca fuscata yakui). We observed all sexually mature individuals (n = 28) in one macaque study group between October 2007 and August 2008, and collected two faecal samples/month/individual (n = 362) for parasitological examination. We used detrended fluctuation analysis (DFA) to investigate long-range autocorrelation in separate, binary sequences of foraging (n = 459) and locomotion (n = 446) behaviour collected via focal sampling. All behavioural sequences exhibited long-range autocorrelation, and linear mixed-effects models suggest that increasing infection with the nodular worm Oesophagostomum aculeatum, clinically impaired health, reproductive activity, ageing and low dominance status were associated with reductions in the complexity of locomotion, and to a lesser extent foraging, behaviour. Furthermore, the sequential complexity of behaviour increased with environmental complexity. We argue that a reduction in complexity in animal behaviour characterizes individuals in impaired or ‘stressed’ states, and may have consequences if animals cannot cope with heterogeneity in their natural habitats.     

The importance of accurate muscle modelling for biomechanical analyses: a case study with a lizard skull

Computer-based simulation techniques such as multi-body dynamics analysis are becoming increasingly popular in the field of skull mechanics. Multi-body models can be used for studying the relationships between skull architecture, muscle morphology and feeding performance. However, to be confident in the modelling results, models need to be validated against experimental data, and the effects of uncertainties or inaccuracies in the chosen model attributes need to be assessed with sensitivity analyses. Here, we compare the bite forces predicted by a multi-body model of a lizard (Tupinambis merianae) with in vivo measurements, using anatomical data collected from the same specimen. This subject-specific model predicts bite forces that are very close to the in vivo measurements and also shows a consistent increase in bite force as the bite position is moved posteriorly on the jaw. However, the model is very sensitive to changes in muscle attributes such as fibre length, intrinsic muscle strength and force orientation, with bite force predictions varying considerably when these three variables are altered. We conclude that accurate muscle measurements are crucial to building realistic multi-body models and that subject-specific data should be used whenever possible.     

Mechanistic analysis of the search behaviour of Caenorhabditis elegans

A central question in movement research is how animals use information and movement to promote encounter success. Current random search theory identifies reorientation patterns as key to the compromise between optimizing encounters for both nearby and faraway targets, but how the balance between intrinsic motor programmes and previous environmental experience determines the occurrence of these reorientation behaviours remains unknown. We used high-resolution tracking and imaging data to describe the complete motor behaviour of Caenorhabditis elegans when placed in a novel environment (one in which food is absent). Movement in C. elegans is structured around different reorientation behaviours, and we measured how these contributed to changing search strategies as worms became familiar with their new environment. This behavioural transition shows that different reorientation behaviours are governed by two processes: (i) an environmentally informed ‘extrinsic’ strategy that is influenced by recent experience and that controls for area-restricted search behaviour, and (ii) a time-independent, ‘intrinsic’ strategy that reduces spatial oversampling and improves random encounter success. Our results show how movement strategies arise from a balance between intrinsic and extrinsic mechanisms, that search behaviour in C. elegans is initially determined by expectations developed from previous environmental experiences, and which reorientation behaviours are modified as information is acquired from new environments.     

Disparity and convergence in bipedal archosaur locomotion

This study aims to investigate functional disparity in the locomotor apparatus of bipedal archosaurs. We use reconstructions of hindlimb myology of extant and extinct archosaurs to generate musculoskeletal biomechanical models to test hypothesized convergence between bipedal crocodile-line archosaurs and dinosaurs. Quantitative comparison of muscle leverage supports the inference that bipedal crocodile-line archosaurs and non-avian theropods had highly convergent hindlimb myology, suggesting similar muscular mechanics and neuromuscular control of locomotion. While these groups independently evolved similar musculoskeletal solutions to the challenges of parasagittally erect bipedalism, differences also clearly exist, particularly the distinct hip and crurotarsal ankle morphology characteristic of many pseudosuchian archosaurs. Furthermore, comparative analyses of muscle design in extant archosaurs reveal that muscular parameters such as size and architecture are more highly adapted or optimized for habitual locomotion than moment arms. The importance of these aspects of muscle design, which are not directly retrievable from fossils, warns against over-extrapolating the functional significance of anatomical convergences. Nevertheless, links identified between posture, muscle moments and neural control in archosaur locomotion suggest that functional interpretations of osteological changes in limb anatomy traditionally linked to postural evolution in Late Triassic archosaurs could be constrained through musculoskeletal modelling.     

仿鱼鳍推进水下机器人流体动力测试系统

以基于仿生柔性长鳍波动推进的水下机器人试验模型为背景,主要研究如何构建满足多种水下机器人试验模型在各种姿态下流体动力/力矩的静态与动态测量要求的测试系统.文章首先提出流体动力测试系统的功能需求与设计准则,并按照模块化设计思想设计其体系结构,重点研究了6分力测量原理、传感器阵列配置结构以及测试流程,最后对整个系统进行了误差源分析.论文研究的测试系统具有6分力测量、2自由度运动导向以及载体运动参数测量等功能,是支撑柔性长鳍波动推进水下机器人研究的重要试验平台.     

Mapping the stereotyped behaviour of freely moving fruit flies

A frequent assumption in behavioural science is that most of an animal''s activities can be described in terms of a small set of stereotyped motifs. Here, we introduce a method for mapping an animal''s actions, relying only upon the underlying structure of postural movement data to organize and classify behaviours. Applying this method to the ground-based behaviour of the fruit fly, Drosophila melanogaster, we find that flies perform stereotyped actions roughly 50% of the time, discovering over 100 distinguishable, stereotyped behavioural states. These include multiple modes of locomotion and grooming. We use the resulting measurements as the basis for identifying subtle sex-specific behavioural differences and revealing the low-dimensional nature of animal motions.     

Magnetically Driven Undulatory Microswimmers Integrating Multiple Rigid Segments

Mimicking biological locomotion strategies offers important possibilities and motivations for robot design and control methods. Among bioinspired microrobots, flexible microrobots exhibit remarkable efficiency and agility. These microrobots traditionally rely on soft material components to achieve undulatory propulsion, which may encounter challenges in design and manufacture including the complex fabrication processes and the interfacing of rigid and soft components. Herein, a bioinspired magnetically driven microswimmer that mimics the undulatory propulsive mechanism is proposed. The designed microswimmer consists of four rigid segments, and each segment is connected to the succeeding segment by joints. The microswimmer is fabricated integrally by 3D laser lithography without further assembly, thereby simplifying microrobot fabrication while enhancing structural integrity. Experimental results show that the microswimmer can successfully swim forward along guided directions via undulatory locomotion in the low Reynolds number (Re) regime. This work demonstrates for the first time that the flexible characteristic of microswimmers can be emulated by 3D structures with multiple rigid segments, which broadens possibilities in microrobot design. The proposed magnetically driven microswimmer can potentially be used in biomedical applications, such as medical diagnosis and treatment in precision medicine.     

Logic programming to predict cell fate patterns and retrodict genotypes in organogenesis

Caenorhabditis elegans vulval development is a paradigm system for understanding cell differentiation in the process of organogenesis. Through temporal and spatial controls, the fate pattern of six cells is determined by the competition of the LET-23 and the Notch signalling pathways. Modelling cell fate determination in vulval development using state-based models, coupled with formal analysis techniques, has been established as a powerful approach in predicting the outcome of combinations of mutations. However, computing the outcomes of complex and highly concurrent models can become prohibitive. Here, we show how logic programs derived from state machines describing the differentiation of C. elegans vulval precursor cells can increase the speed of prediction by four orders of magnitude relative to previous approaches. Moreover, this increase in speed allows us to infer, or ‘retrodict’, compatible genomes from cell fate patterns. We exploit this technique to predict highly variable cell fate patterns resulting from dig-1 reduced-function mutations and let-23 mosaics. In addition to the new insights offered, we propose our technique as a platform for aiding the design and analysis of experimental data.     

Finite-element modelling reveals force modulation of jaw adductors in stag beetles

Male stag beetles carry large and heavy mandibles that arose through sexual selection over mating rights. Although the mandibles of Cyclommatus metallifer males are used in pugnacious fights, they are surprisingly slender. Our bite force measurements show a muscle force reduction of 18% for tip biting when compared with bites with the teeth located halfway along the mandibles. This suggests a behavioural adaptation to prevent failure. We confirmed this by constructing finite-element (FE) models that mimic both natural bite situations as well as the hypothetical situation of tip biting without muscle force modulation. These models, based on micro-CT images, investigate the material stresses in the mandibles for different combinations of bite location and muscle force. Young''s modulus of the cuticle was experimentally determined to be 5.1 GPa with the double indentation method, and the model was validated by digital image correlation on living beetles. FE analysis proves to be a valuable tool in the investigation of the trade-offs of (animal) weapon morphology and usage. Furthermore, the demonstrated bite force modulation in male stag beetles suggests the presence of mechanosensors inside the armature.     

Fabrication and actuation of ionic polymer metal composites patterned by combining electroplating with electroless plating

The novel fabrication technique that patterns the multiple electrodes of the ionic polymer metal composite actuators was developed to mimic the swimming and flapping locomotion of a living thing. The developed method is to combine electroplating with the electroless chemical reduction using the patterned mask. The advantages of this fabrication method are that the initial compositing between the polymer and platinum particles can be assured by the chemical reduction method, and the thickness of each electrode can be controlled easily and rapidly by electroplating. By using the fabricated actuator with a multiple degree of freedom, the oscillatory and undulatory waves of the flexible membrane actuator was generated and a twisting motion was also realized to verify the possibility of mimicking the fish-like locomotion. Present results show that this novel method combining electroplating with electroless plating can be a promising technique to easily pattern multiple electrodes and to implement the biomimetic motion of the polymer actuators with good mechanical bending performance.     

Numerical study of a sphere descending along an inclined slope in a liquid

The descending process of a sphere rolling and/or sliding along an inclined slope in a liquid involves interactions between the hydrodynamic forces on the sphere and the contact forces between the sphere and the plane. In this study, the descending process of sphere in a liquid was examined using coupled LBM–DEM technique. The effects of slope angle, viscosity and friction coefficient on the movement of a sphere were investigated. Two distinct descending patterns were observed: (a) a stable rolling/sliding movement along the slope, and (b) a fluctuating pattern along the slope. Five dimensionless coefficients (Reynolds number (Re), drag coefficient, lift coefficient, moment coefficient and rolling coefficient) were used to analyze the observed processes. The vortex structure in the wake of the sphere gives a lift force to the sphere, which in turn controls the different descending patterns. It is found that the generation of a vortex is not only governed by Re, but also by particle rotation. Relationships between the forces/moments and the dimensionless coefficients are established.     

Curvature interference characteristic of conical surface enveloping conical worm

The theory to distinguish whether the curvature interference happens or not is well established for the enveloping conical worm. Some important results, such as the meshing function, the equation of the enveloping conical worm helicoid, and the curvature interference limit function, are all obtained. The foundation of computing the curvature interference limit line is to solve the system of nonlinear equations. A technique based on the elimination method and the geometric construction is proposed, which can be employed to study the existence of the solution of a system of nonlinear equations and to solve all the solutions of this system within the given solving domain. By means of the theory and technique brought forward, it is discovered that there usually are two curvature interference limit lines separately on each side of a tooth of an enveloping conical worm. When the number of the enveloping conical worm threads is smaller, the curvature interference in general does not happen on the both sides of a tooth. The avoiding mechanism of the curvature interference is that the limit line does not exist during the real cutting mesh because the limit line is inside the entity of the enveloping conical worm and its conjugate line is outside the entity of the grinding wheel. The numerical outcome shows that, the toe on the \(e\) flank has the greatest potential risk to be subjected to the undercutting.     

Muscle performance in a soft-bodied terrestrial crawler: constitutive modelling of strain-rate dependency.

Experimental data on the passive mechanical properties of the ventral interior lateral muscle of the tobacco hornworm caterpillar, Manduca sexta, are reported. The stress-deformation response of the Manduca muscle is shown to be nonlinear pseudo-elastic, capable of large deformations and subject to stress softening during initial loading cycles. The muscle passive mechanical properties also depend on multiple time-dependent processes. In particular, we show new experimental data from cyclic loading tests of an unstimulated muscle with constant maximum stretch and different, constant engineering strain rates. Then, on the basis of these data a constitutive model is derived to reproduce the main characteristics of this behaviour. In formulating the constitutive model, we consider the muscle as a complex macromolecular structure with fibrous components at numerous size scales. The model uses a phenomenological approach to account for different mechanisms by which passive force changes during applied deformation and how the muscle properties recover after unloading.     

基液黏度对草酸氧钛电流变液性能的影响

以草酸氧钛纳米颗粒为分散相,以不同黏度的二甲基硅油(运动学黏度为10 cSt、50 cSt、100 cSt、500 cSt和1000 cSt)为基液制备电流变液,测试其剪切屈服强度、零场黏度、电流变效率、响应时间及沉降稳定性,研究了基液黏度对草酸氧钛电流变液性能的影响。结果表明,以50 cSt二甲基硅油为基液的电流变液具有最佳的电流变效率,以100 cSt二甲基硅油为基液的电流变液具有最短的响应时间和较高的沉降稳定性。其机制是,较高的基液黏度可提供较大的黏滞阻力,但易引起颗粒的团聚。     

Stability of Plane-Parallel Pulsational Flow of Two Miscible Fluids under High Frequency Horizontal Vibrations

Linear stability of plane-parallel pulsational flow of two miscible fluids in a horizontal layer subjected to high frequency horizontal vibrations is investigated neglecting viscosity and diffusion. Long-wave instability is studied analytically and instability to the perturbations with finite wavelength—numerically.     

Intrinsic foot muscles have the capacity to control deformation of the longitudinal arch

The human foot is characterized by a pronounced longitudinal arch (LA) that compresses and recoils in response to external load during locomotion, allowing for storage and return of elastic energy within the passive structures of the arch and contributing to metabolic energy savings. Here, we examine the potential for active muscular contribution to the biomechanics of arch deformation and recoil. We test the hypotheses that activation of the three largest plantar intrinsic foot muscles, abductor hallucis, flexor digitorum and quadratus plantae is associated with muscle stretch in response to external load on the foot and that activation of these muscles (via electrical stimulation) will generate sufficient force to counter the deformation of LA caused by the external load. We found that recruitment of the intrinsic foot muscles increased with increasing load, beyond specific load thresholds. Interestingly, LA deformation and muscle stretch plateaued towards the maximum load of 150% body weight, when muscle activity was greatest. Electrical stimulation of the plantar intrinsic muscles countered the deformation that occurred owing to the application of external load by reducing the length and increasing the height of the LA. These findings demonstrate that these muscles have the capacity to control foot posture and LA stiffness and may provide a buttressing effect during foot loading. This active arch stiffening mechanism may have important implications for how forces are transmitted during locomotion and postural activities as well as consequences for metabolic energy saving.