Vortex wake,downwash distribution,aerodynamic performance and wingbeat kinematics in slow-flying pied flycatchers

Many small passerines regularly fly slowly when catching prey, flying in cluttered environments or landing on a perch or nest. While flying slowly, passerines generate most of the flight forces during the downstroke, and have a ‘feathered upstroke’ during which they make their wing inactive by retracting it close to the body and by spreading the primary wing feathers. How this flight mode relates aerodynamically to the cruising flight and so-called ‘normal hovering’ as used in hummingbirds is not yet known. Here, we present time-resolved fluid dynamics data in combination with wingbeat kinematics data for three pied flycatchers flying across a range of speeds from near hovering to their calculated minimum power speed. Flycatchers are adapted to low speed flight, which they habitually use when catching insects on the wing. From the wake dynamics data, we constructed average wingbeat wakes and determined the time-resolved flight forces, the time-resolved downwash distributions and the resulting lift-to-drag ratios, span efficiencies and flap efficiencies. During the downstroke, slow-flying flycatchers generate a single-vortex loop wake, which is much more similar to that generated by birds at cruising flight speeds than it is to the double loop vortex wake in hovering hummingbirds. This wake structure results in a relatively high downwash behind the body, which can be explained by the relatively active tail in flycatchers. As a result of this, slow-flying flycatchers have a span efficiency which is similar to that of the birds in cruising flight and which can be assumed to be higher than in hovering hummingbirds. During the upstroke, the wings of slowly flying flycatchers generated no significant forces, but the body–tail configuration added 23 per cent to weight support. This is strikingly similar to the 25 per cent weight support generated by the wing upstroke in hovering hummingbirds. Thus, for slow-flying passerines, the upstroke cannot be regarded as inactive, and the tail may be of importance for flight efficiency and possibly manoeuvrability.     

Comparative aerodynamic performance of flapping flight in two bat species using time-resolved wake visualization

Bats are unique among extant actively flying animals in having very flexible wings, controlled by multi-jointed fingers. This gives the potential for fine-tuned active control to optimize aerodynamic performance throughout the wingbeat and thus a more efficient flight. But how bat wing performance scales with size, morphology and ecology is not yet known. Here, we present time-resolved fluid wake data of two species of bats flying freely across a range of flight speeds using stereoscopic digital particle image velocimetry in a wind tunnel. From these data, we construct an average wake for each bat species and speed combination, which is used to estimate the flight forces throughout the wingbeat and resulting flight performance properties such as lift-to-drag ratio (L/D). The results show that the wake dynamics and flight performance of both bat species are similar, as was expected since both species operate at similar Reynolds numbers (Re) and Strouhal numbers (St). However, maximum L/D is achieved at a significant higher flight speed for the larger, highly mobile and migratory bat species than for the smaller non-migratory species. Although the flight performance of these bats may depend on a range of morphological and ecological factors, the differences in optimal flight speeds between the species could at least partly be explained by differences in their movement ecology.     

Time-varying span efficiency through the wingbeat of desert locusts

The flight performance of animals depends greatly on the efficacy with which they generate aerodynamic forces. Accordingly, maximum range, load-lifting capacity and peak accelerations during manoeuvres are all constrained by the efficiency of momentum transfer to the wake. Here, we use high-speed particle image velocimetry (1 kHz) to record flow velocities in the near wake of desert locusts (Schistocerca gregaria, Forskål). We use the measured flow fields to calculate time-varying span efficiency throughout the wing stroke cycle. The locusts are found to operate at a maximum span efficiency of 79 per cent, typically at a plateau of about 60 per cent for the majority of the downstroke, but at lower values during the upstroke. Moreover, the calculated span efficiencies are highest when the largest lift forces are being generated (90% of the total lift is generated during the plateau of span efficiency) suggesting that the combination of wing kinematics and morphology in locust flight perform most efficiently when doing the most work.     

Time-resolved vortex wake of a common swift flying over a range of flight speeds

The wake of a freely flying common swift (Apus apus L.) is examined in a wind tunnel at three different flight speeds, 5.7, 7.7 and 9.9 m s⁻¹. The wake of the bird is visualized using high-speed stereo digital particle image velocimetry (DPIV). Wake images are recorded in the transverse plane, perpendicular to the airflow. The wake of a swift has been studied previously using DPIV and recording wake images in the longitudinal plane, parallel to the airflow. The high-speed DPIV system allows for time-resolved wake sampling and the result shows features that were not discovered in the previous study, but there was approximately a 40 per cent vertical force deficit. As the earlier study also revealed, a pair of wingtip vortices are trailing behind the wingtips, but in addition, a pair of tail vortices and a pair of ‘wing root vortices’ are found that appear to originate from the wing/body junction. The existence of wing root vortices suggests that the two wings are not acting as a single wing, but are to some extent aerodynamically detached from each other. It is proposed that this is due to the body disrupting the lift distribution over the wing by generating less lift than the wings.     

The influence of sensory delay on the yaw dynamics of a flapping insect

In closed-loop systems, sensor feedback delays may have disastrous implications for performance and stability. Flies have evolved multiple specializations to reduce this latency, but the fastest feedback during flight involves a delay that is still significant on the timescale of body dynamics. We explored the effect of sensor delay on flight stability and performance for yaw turns using a dynamically scaled robotic model of the fruitfly, Drosophila. The robot was equipped with a real-time feedback system that performed active turns in response to measured torque about the functional yaw axis. We performed system response experiments for a proportional controller in yaw velocity for a range of feedback delays, similar in dimensionless timescale to those experienced by a fly. The results show a fundamental trade-off between sensor delay and permissible feedback gain, and suggest that fast mechanosensory feedback in flies, and most probably in other insects, provide a source of active damping which compliments that contributed by passive effects. Presented in the context of these findings, a control architecture whereby a haltere-mediated inner-loop proportional controller provides damping for slower visually mediated feedback is consistent with tethered-flight measurements, free-flight observations and engineering design principles.     

Embodied linearity of speed control in Drosophila melanogaster

Fruitflies regulate flight speed by adjusting their body angle. To understand how low-level posture control serves an overall linear visual speed control strategy, we visually induced free-flight acceleration responses in a wind tunnel and measured the body kinematics using high-speed videography. Subsequently, we reverse engineered the transfer function mapping body pitch angle onto flight speed. A linear model is able to reproduce the behavioural data with good accuracy. Our results show that linearity in speed control is realized already at the level of body posture-mediated speed control and is therefore embodied at the level of the complex aerodynamic mechanisms of body and wings. Together with previous results, this study reveals the existence of a linear hierarchical control strategy, which can provide relevant control principles for biomimetic implementations, such as autonomous flying micro air vehicles.     

圆柱绕流的流场特性及涡脱落规律研究

采用粒子图像测速技术对630、800及950三种雷诺数条件下的圆柱绕流场进行了实验,给出了圆柱下游沿流动方向4倍圆柱直径和垂直方向3倍圆柱直径区域内的速度场、涡量场以及涡脱落现象的时空演化规律.结果表明:圆柱尾流区域位于垂直方向约1.5~2.5倍圆柱直径范围内,随着雷诺数增大,这一范围呈现缩小趋势,而主流对涡的拉伸和输运能力有所增强;涡脱落频率随雷诺数增大而增大,小雷诺数时能够较为完整地捕捉到涡生成、脱落、发展和耗散过程,由于PIV采集频率的限制,大雷诺数条件下涡脱落整个过程不易被完整捕捉到.     

Measurements of the wall shear stress distribution in the outflow tract of an embryonic chicken heart

In order to study the role of blood–tissue interaction in the developing chicken embryo heart, detailed information about the haemodynamic forces is needed. In this study, we present the first in vivo measurements of the three-dimensional distribution of wall shear stress (WSS) in the outflow tract (OFT) of an embryonic chicken heart. The data are obtained in a two-step process: first, the three-dimensional flow fields are measured during the cardiac cycle using scanning microscopic particle image velocimetry; second, the location of the wall and the WSS are determined by post-processing flow velocity data (finding velocity gradients at locations where the flow approaches zero). The results are a three-dimensional reconstruction of the geometry, with a spatial resolution of 15–20 µm, and provides detailed information about the WSS in the OFT. The most significant error is the location of the wall, which results in an estimate of the uncertainty in the WSS values of 20 per cent.     

Reconstructing the flight kinematics of swarming and mating in wild mosquitoes

We describe a novel tracking system for reconstructing three-dimensional tracks of individual mosquitoes in wild swarms and present the results of validating the system by filming swarms and mating events of the malaria mosquito Anopheles gambiae in Mali. The tracking system is designed to address noisy, low frame-rate (25 frames per second) video streams from a stereo camera system. Because flying A. gambiae move at 1–4 m s⁻¹, they appear as faded streaks in the images or sometimes do not appear at all. We provide an adaptive algorithm to search for missing streaks and a likelihood function that uses streak endpoints to extract velocity information. A modified multi-hypothesis tracker probabilistically addresses occlusions and a particle filter estimates the trajectories. The output of the tracking algorithm is a set of track segments with an average length of 0.6–1 s. The segments are verified and combined under human supervision to create individual tracks up to the duration of the video (90 s). We evaluate tracking performance using an established metric for multi-target tracking and validate the accuracy using independent stereo measurements of a single swarm. Three-dimensional reconstructions of A. gambiae swarming and mating events are presented.     

Use of multiple modes of flight subsidy by a soaring terrestrial bird,the golden eagle Aquila chrysaetos,when on migration

Large birds regularly use updrafts to subsidize flight. Although most research on soaring bird flight has focused on use of thermal updrafts, there is evidence suggesting that many species are likely to use multiple modes of subsidy. We tested the degree to which a large soaring species uses multiple modes of subsidy to provide insights into the decision-making that underlies flight behaviour. We statistically classified more than 22 000 global positioning satellite–global system for mobile communications telemetry points collected at 30-s intervals to identify the type of subsidized flight used by 32 migrating golden eagles during spring in eastern North America. Eagles used subsidized flight on 87% of their journey. They spent 41.9% ± 1.5 (, range: 18–56%) of their subsidized northbound migration using thermal soaring, 45.2% ± 2.1 (12–65%) of time gliding between thermals, and 12.9% ± 2.2 (1–55%) of time using orographic updrafts. Golden eagles responded to the variable local-scale meteorological events they encountered by switching flight behaviour to take advantage of multiple modes of subsidy. Orographic soaring occurred more frequently in morning and evening, earlier in the migration season, and when crosswinds and tail winds were greatest. Switching between flight modes allowed migration for relatively longer periods each day and frequent switching behaviour has implications for a better understanding of avian flight behaviour and of the evolution of use of subsidy in flight.     

Understanding macroalgal dispersal in a complex hydrodynamic environment: a combined population genetic and physical modelling approach

Gene flow in macroalgal populations can be strongly influenced by spore or gamete dispersal. This, in turn, is influenced by a convolution of the effects of current flow and specific plant reproductive strategies. Although several studies have demonstrated genetic variability in macroalgal populations over a wide range of spatial scales, the associated current data have generally been poorly resolved spatially and temporally. In this study, we used a combination of population genetic analyses and high-resolution hydrodynamic modelling to investigate potential connectivity between populations of the kelp Laminaria digitata in the Strangford Narrows, a narrow channel characterized by strong currents linking the large semi-enclosed sea lough, Strangford Lough, to the Irish Sea. Levels of genetic structuring based on six microsatellite markers were very low, indicating high levels of gene flow and a pattern of isolation-by-distance, where populations are more likely to exchange migrants with geographically proximal populations, but with occasional long-distance dispersal. This was confirmed by the particle tracking model, which showed that, while the majority of spores settle near the release site, there is potential for dispersal over several kilometres. This combined population genetic and modelling approach suggests that the complex hydrodynamic environment at the entrance to Strangford Lough can facilitate dispersal on a scale exceeding that proposed for L. digitata in particular, and the majority of macroalgae in general. The study demonstrates the potential of integrated physical–biological approaches for the prediction of ecological changes resulting from factors such as anthropogenically induced coastal zone changes.     

The interplay between a self-organized process and an environmental template: corpse clustering under the influence of air currents in ants

Many spatial patterns observed in nature emerge from local processes and their interactions with the local environment. The clustering of objects by social insects represents such a pattern formation process that can be observed at both the individual and the collective level. In this paper, we study the interaction between air currents and clustering behaviour in order to address the coordinating mechanisms at the individual level that underlie the spatial pattern formation process in a heterogeneous environment. We choose the corpse clustering behaviour of the ant Messor sanctus as an experimental paradigm. In a specifically designed experimental set-up with a well-controlled laminar air flow (approx. 1 cm s-1), we first quantify the modulation of the individual corpse aggregation behaviour as a function of corpse density, air flow intensity and the ant's position with respect to corpse piles and air flow direction. We then explore by numerical simulation how the forming corpse piles modify the laminar air flow around them and link this result with the individual behaviour modulation. Finally, we demonstrate on the collective level that this laminar air flow leads to an elongation and a slow displacement of the formed corpse piles in the direction of the air current. Both the individual behaviour modulated by air flow and the air flow modulated by the forming corpse piles can explain the pile patterns observed on the collective level as a stigmergic process. We discuss the generality of this coordinating mechanism to explain the clustering phenomena in heterogeneous environments reported in the literature.     

Population structure in the Neisseria, and the biological significance of fuzzy species

Phenotypic and genetic variation in bacteria can take bewilderingly complex forms even within a single genus. One of the most intriguing examples of this is the genus Neisseria, which comprises both pathogens and commensals colonizing a variety of body sites and host species, and causing a range of disease. Complex relatedness among both named species and previously identified lineages of Neisseria makes it challenging to study their evolution. Using the largest publicly available collection of bacterial sequence data in combination with a population genetic analysis and experiment, we probe the contribution of inter-species recombination to neisserial population structure, and specifically whether it is more common in some strains than others. We identify hybrid groups of strains containing sequences typical of more than one species. These groups of strains, typical of a fuzzy species, appear to have experienced elevated rates of inter-species recombination estimated by population genetic analysis and further supported by transformation experiments. In particular, strains of the pathogen Neisseria meningitidis in the fuzzy species boundary appear to follow a different lifestyle, which may have considerable biological implications concerning distribution of novel resistance elements and meningococcal vaccine development. Despite the strong evidence for negligible geographical barriers to gene flow within the population, exchange of genetic material still shows directionality among named species in a non-uniform manner.     

High-pressure studies on Tc and crystal structure of iron chalcogenide superconductors

The superconducting transition temperature, T_c, in iron-based solids can be enhanced by applied pressure: T_c increases from 8 to 37 K for the 11-type FeSe when the pressure is raised from 0 to 4 GPa. High-pressure studies can elucidate the mechanism of superconductivity in such novel materials. In this paper, we present a high-pressure study of Fe(Se_1−xTe_x) and Fe(Se_1−xS_x). In the case of Fe(Se_1−xTe_x), the maximum T_c under high pressure did not exceed the T_c of FeSe, which can be attributed to the structural transition to the monoclinic phase. For Fe(Se_1−xS_x) (0 < x < 0.3), T_c exhibited a significant increase with pressure; however, the maximum T_c under high pressure did not exceed the T_c of FeSe. This may be due to the disorder induced by substituting S for Se, which is similar to the pressure effect on T_c for the 1111-type superconductor Ca(Fe_1−xCo_x)AsF. The T_c of Fe(Se_1−xS_x) showed a complex behavior below 1 GPa, first decreasing and then increasing with increasing pressure. From high-pressure x-ray diffraction measurements, the T_c (P) curve was correlated with the local structural parameter.     

Campylobacter jejuni colonization and transmission in broiler chickens: a modelling perspective.

Campylobacter jejuni is one of the most common causes of acute enteritis in the developed world. The consumption of contaminated poultry, where C. jejuni is believed to be a commensal organism, is a major risk factor. However, the dynamics of this colonization process in commercially reared chickens is still poorly understood. Quantification of these dynamics of infection at an individual level is vital to understand transmission within populations and formulate new control strategies. There are multiple potential routes of introduction of C. jejuni into a commercial flock. Introduction is followed by a rapid increase in environmental levels of C. jejuni and the level of colonization of individual broilers. Recent experimental and epidemiological evidence suggest that the celerity of this process could be masking a complex pattern of colonization and extinction of bacterial strains within individual hosts. Despite the rapidity of colonization, experimental transmission studies exhibit a highly variable and unexplained delay time in the initial stages of the process. We review past models of transmission of C. jejuni in broilers and consider simple modifications, motivated by the plausible biological mechanisms of clearance and latency, which could account for this delay. We show how simple mathematical models can be used to guide the focus of experimental studies by providing testable predictions based on our hypotheses. We conclude by suggesting that competition experiments could be used to further understand the dynamics and mechanisms underlying the colonization process. The population models for such competition processes have been extensively studied in other ecological and evolutionary contexts. However, C. jejuni can potentially adapt phenotypically through phase variation in gene expression, leading to unification of ecological and evolutionary time-scales. For a theoretician, the colonization dynamics of C. jejuni offer an experimental system to explore these 'phylodynamics', the synthesis of population dynamics and evolutionary biology.     

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.     

Cell wall structure and formation of maturing fibres of moso bamboo (Phyllostachys pubescens) increase buckling resistance

The mechanical stability of the culms of monocotyledonous bamboos is highly attributed to the proper embedding of the stiff fibre caps of the vascular bundles into the soft parenchymatous matrix. Owing to lack of a vascular cambium, bamboos show no secondary thickening growth that impedes geometrical adaptations to mechanical loads and increases the necessity of structural optimization at the material level. Here, we investigate the fine structure and mechanical properties of fibres within a maturing vascular bundle of moso bamboo, Phyllostachys pubescens, with a high spatial resolution. The fibre cell walls were found to show almost axially oriented cellulose fibrils, and the stiffness and hardness of the central part of the cell wall remained basically consistent for the fibres at different regions across the fibre cap. A stiffness gradient across the fibre cap is developed by differential cell wall thickening which affects tissue density and thereby axial tissue stiffness in the different regions of the cap. The almost axially oriented cellulose fibrils in the fibre walls maximize the longitudinal elastic modulus of the fibres and their lignification increases the transverse rigidity. This is interpreted as a structural and mechanical optimization that contributes to the high buckling resistance of the slender bamboo culms.     

Empirical assessment of a threshold model for sylvatic plague.

Plague surveillance programmes established in Kazakhstan, Central Asia, during the previous century, have generated large plague archives that have been used to parameterize an abundance threshold model for sylvatic plague in great gerbil (Rhombomys opimus) populations. Here, we assess the model using additional data from the same archives. Throughout the focus, population levels above the threshold were a necessary condition for an epizootic to occur. However, there were large numbers of occasions when an epizootic was not observed even though great gerbils were, and had been, abundant. We examine six hypotheses that could explain the resulting false positive predictions, namely (i) including end-of-outbreak data erroneously lowers the estimated threshold, (ii) too few gerbils were tested, (iii) plague becomes locally extinct, (iv) the abundance of fleas was too low, (v) the climate was unfavourable, and (vi) a high proportion of gerbils were resistant. Of these, separate thresholds, fleas and climate received some support but accounted for few false positives and can be disregarded as serious omissions from the model. Small sample size and local extinction received strong support and can account for most of the false positives. Host resistance received no support here but should be subject to more direct experimental testing.     

Stochastic extinction and the selection of the transmission mode in microparasites.

Stochastic fluctuations in the transmission process of microparasites generate a risk of parasite extinction that cannot be assessed by deterministic models, especially in host populations of small size. While this risk of extinction represents a strong selection pressure for microparasites, it is usually not clearly separated from the deterministic ones. We suggest here that this stochastic selection pressure can affect the selection of the transmission mode of microparasites. To avoid extinction, parasites should maximize their inter-population transmission to ensure frequent reintroductions. Since the types of contacts may differ if congeners belong to the same or distinct populations, strains that are mainly transmitted through inter-population contacts might be selected. To examine this assumption, we analyse the issue of the competition between two strains differing in their transmission mode using a stochastic metapopulation model in which hosts may display different behaviours inside and outside their populations. We show that stochastic selection pressures may drive parasite evolution towards a transmission mode that maximizes the persistence of the parasite. We study the conditions under which stochastic selection pressures may surpass the deterministic ones. Our results are illustrated by the cases of feline immunodeficiency virus in cats and of sexually transmitted diseases in mammals.     

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.