Insect wings are deformable structures that change shape passively and dynamically owing to inertial and aerodynamic forces during flight. It is still unclear how the three-dimensional and passive change of wing kinematics owing to inherent wing flexibility contributes to unsteady aerodynamics and energetics in insect flapping flight. Here, we perform a systematic fluid-structure interaction based analysis on the aerodynamic performance of a hovering hawkmoth,Manduca, with an integrated computational model of a hovering insect with rigid and flexible wings. Aerodynamic performance of flapping wings with passive deformation or prescribed deformation is evaluated in terms of aerodynamic force, power and efficiency. Our results reveal that wing flexibility can increase downwash in wake and hence aerodynamic force: first, a dynamic wing bending is observed, which delays the breakdown of leading edge vortex near the wing tip, responsible for augmenting the aerodynamic force-production; second, a combination of the dynamic change of wing bending and twist favourably modifies the wing kinematics in the distal area, which leads to the aerodynamic force enhancement immediately before stroke reversal. Moreover, an increase in hovering efficiency of the flexible wing is achieved as a result of the wing twist. An extensive study of wing stiffness effect on aerodynamic performance is further conducted through a tuning of Young's modulus and thickness, indicating that insect wing structures may be optimized not only in terms of aerodynamic performance but also dependent on many factors, such as the wing strength, the circulation capability of wing veins and the control of wing movements.

There are nearly a million known species of flying insects and 13 000 species of flying warm-blooded vertebrates, including mammals, birds and bats. While in flight, their wings not only move forward relative to the air, they also flap up and down, plunge and sweep, so that both lift and thrust can be generated and balanced, accommodate uncertain surrounding environment, with superior flight stability and dynamics with highly varied speeds and missions. As the size of a flyer is reduced, the wing-to-body mass ratio tends to decrease as well. Furthermore, these flyers use integrated system consisting of wings to generate aerodynamic forces, muscles to move the wings, and sensing and control systems to guide and manoeuvre. In this article, recent advances in insect-scale flapping-wing aerodynamics, flexible wing structures, unsteady flight environment, sensing, stability and control are reviewed with perspective offered. In particular, the special features of the low Reynolds number flyers associated with small sizes, thin and light structures, slow flight with comparable wind gust speeds, bioinspired fabrication of wing structures, neuron-based sensing and adaptive control are highlighted.

Insect- and bird-size drones—micro air vehicles (MAV) that can perform autonomous flight in natural and man-made environments are now an active and well-integrated research area. MAVs normally operate at a low speed in a Reynolds number regime of 10 ⁴ –10 ⁵ or lower, in which most flying animals of insects, birds and bats fly, and encounter unconventional challenges in generating sufficient aerodynamic forces to stay airborne and in controlling flight autonomy to achieve complex manoeuvres. Flying insects that power and control flight by flapping wings are capable of sophisticated aerodynamic force production and precise, agile manoeuvring, through an integrated system consisting of wings to generate aerodynamic force, muscles to move the wings and a control system to modulate power output from the muscles. In this article, we give a selective review on the state of the art of biomechanics in bioinspired flight systems in terms of flapping and flexible wing aerodynamics, flight dynamics and stability, passive and active mechanisms in stabilization and control, as well as flapping flight in unsteady environments. We further highlight recent advances in biomimetics of flapping-wing MAVs with a specific focus on insect-inspired wing design and fabrication, as well as sensing systems. This article is part of the themed issue ‘Moving in a moving medium: new perspectives on flight’.

Cerebral hyperperfusion syndrome develops in a small subset of patients following carotid artery surgery (CAS) performed to treat severe carotid artery stenosis. This syndrome has been found to have a close correlation with cerebral hyperperfusion occurring after CAS. The purpose of this study is to investigate whether and how the anatomy of the Circle of Willis (CoW) of the cerebral circulation influences post-CAS cerebral hyperperfusion. A computational model of the cerebral circulation coupled with the global cardiovascular system has been developed to investigate hemodynamic events associated with CAS. Nine topological structures of the CoW were investigated in combination with various distribution patterns of stenosis in the feeding arteries of the cerebral circulation. The occurrence of post-CAS cerebral hyperperfusion was predicted for the CoW structures that have poor collateral pathways between the stenosed cerebral feeding arteries and the remaining normal feeding arteries. The risk and the localization of post-CAS hyperperfusion were determined jointly by the anatomy of the CoW and the distribution pattern of stenosis in the cerebral feeding arteries. The presence of basilar artery stenosis or contralateral ICA stenosis increased the risk of post-CAS hyperperfusion and enlarged the cerebral region affected by hyperperfusion. For a certain CoW structure, the diameters of the cerebral communicating arteries and the severity of carotid artery stenosis both had a significant influence on the computed post-CAS cerebral hyperperfusion rates. Moreover, post-CAS cerebral hyperperfusion was predicted to be accompanied with an excessively high capillary transmural pressure. This study demonstrated the importance of considering the anatomy of the CoW in assessing the risk of post-CAS cerebral hyperperfusion. Particularly, since the anatomy of the CoW and the distribution pattern of stenosis in the cerebral feeding arteries jointly determine the risk and localization of post-CAS cerebral hyperperfusion, a patient-specific hemodynamic analysis aimed to help physicians identify patients at high risk of cerebral hyperperfusion should account for the combined effect of the anatomy of cerebral arteries and cerebral feeding artery stenoses on cerebral hemodynamics.

Flight speed is positively correlated with body size in animals¹. However, miniature featherwing beetles can fly at speeds and accelerations of insects three times their size². Here we show that this performance results from a reduced wing mass and a previously unknown type of wing-motion cycle. Our experiment combines three-dimensional reconstructions of morphology and kinematics in one of the smallest insects, the beetle Paratuposa placentis (body length 395 μm). The flapping bristled wings follow a pronounced figure-of-eight loop that consists of subperpendicular up and down strokes followed by claps at stroke reversals above and below the body. The elytra act as inertial brakes that prevent excessive body oscillation. Computational analyses suggest functional decomposition of the wingbeat cycle into two power half strokes, which produce a large upward force, and two down-dragging recovery half strokes. In contrast to heavier membranous wings, the motion of bristled wings of the same size requires little inertial power. Muscle mechanical power requirements thus remain positive throughout the wingbeat cycle, making elastic energy storage obsolete. These adaptations help to explain how extremely small insects have preserved good aerial performance during miniaturization, one of the factors of their evolutionary success.