Lentink's lab studies every aspect of biological flight as an inspiration for designing aerial robots. We focus on key biological questions which we probe with new engineering methods to find inspiration for aerial robots that can fly like animals. Our comparative biological flight research ranges from maple seeds, insects and bats to birds such as swifts, parrotlets, lovebirds, doves and a wide range of hummingbirds. For in-depth comparative biomechanics research we focus on bird flight. We use biofluid dynamics and other quantitative engineering disciplines including robotics as research tools to mechanistically understand and embody animal flight performance. We translate our integrative and comparative biological research driven by scientific curiosity to aerial robot design to solve the engineering challenge of autonomous flight in complex cluttered environments and turbulent atmospheric conditions.

Academic Appointments

  • Sr Research Engineer, Mechanical Engineering
  • Member, Bio-X

Honors & Awards

  • Inaugural Steven Vogel Young Investigator Award, Journal Bioinspiration & Biomimetics (2018)
  • Gilbreth Lecturer, National Academy of Engineering (2017)
  • CAREER Award, National Science Foundation (2016)
  • Recognized as one of 40 scientists under 40 by the World Economic Forum, Annual Meeting of the New Champions, Dalian, China. (2013)
  • 100kE Dutch Academic Year Prize, 100kE (2010)
  • Biophysics thesis award, Dutch Society for Biophysics and Biomedical Technology (2009)
  • Zoology Award, Royal Dutch Zoological Society (2009)
  • Bolk Prize, Netherlands Society for Anatomy (2008)
  • Ritsema van Eck Award, Delft University of Technology (2006)
  • Most Exotic Micro Air Vehicle (MAV) Award, First American-European MAV contest (2005)
  • Elsevier Young Scientist Award, Society for Experimental Biology (2005)
  • AIAA best Fluid Dynamics conference paper, AIAA (2003)
  • Dobbinga Award, Delft University of Technology (2003)

Boards, Advisory Committees, Professional Organizations

  • Editorial Board Member, Journal of Bioinspiration & Biomimetics (2010 - Present)
  • Reviewing Editorial Board member, eLife (2016 - Present)
  • Alumnus, Young Academy of the Royal Netherlands Academy of Arts and Sciences (2011 - 2016)

Professional Education

  • PhD, cum laude, Wageningen University, The Netherlands, Experimental Zoology (2008)
  • MS, BS, Delft University of Technology, The Netherlands, Aerospace Engineering (2003)

2020-21 Courses

Stanford Advisees

  • Doctoral Dissertation Reader (AC)
    Sandra Schachat
  • Doctoral Dissertation Advisor (AC)
    Kenneth Hoffmann
  • Doctoral Dissertation Co-Advisor (AC)
    Trevor Halsted
  • Doctoral (Program)
    Elliot Weiss

All Publications

  • Soft biohybrid morphing wings with feathers underactuated by wrist and finger motion SCIENCE ROBOTICS Chang, E., Matloff, L. Y., Stowers, A. K., Lentink, D. 2020; 5 (38)
  • How flight feathers stick together to form a continuous morphing wing. Science (New York, N.Y.) Matloff, L. Y., Chang, E., Feo, T. J., Jeffries, L., Stowers, A. K., Thomson, C., Lentink, D. 2020; 367 (6475): 293–97


    Variable feather overlap enables birds to morph their wings, unlike aircraft. They accomplish this feat by means of elastic compliance of connective tissue, which passively redistributes the overlapping flight feathers when the skeleton moves to morph the wing planform. Distinctive microstructures form "directional Velcro," such that when adjacent feathers slide apart during extension, thousands of lobate cilia on the underlapping feathers lock probabilistically with hooked rami of overlapping feathers to prevent gaps. These structures unlock automatically during flexion. Using a feathered biohybrid aerial robot, we demonstrate how both passive mechanisms make morphing wings robust to turbulence. We found that the hooked microstructures fasten feathers across bird species except silent fliers, whose feathers also lack the associated Velcro-like noise. These findings could inspire innovative directional fasteners and morphing aircraft.

    View details for DOI 10.1126/science.aaz3358

    View details for PubMedID 31949079

  • Birds repurpose the role of drag and lift to take off and land. Nature communications Chin, D. D., Lentink, D. 2019; 10 (1): 5354


    The lift that animal wings generate to fly is typically considered a vertical force that supports weight, while drag is considered a horizontal force that opposes thrust. To determine how birds use lift and drag, here we report aerodynamic forces and kinematics of Pacific parrotlets (Forpus coelestis) during short, foraging flights. At takeoff they incline their wing stroke plane, which orients lift forward to accelerate and drag upward to support nearly half of their bodyweight. Upon landing, lift is oriented backward to contribute a quarter of the braking force, which reduces the aerodynamic power required to land. Wingbeat power requirements are dominated by downstrokes, while relatively inactive upstrokes cost almost no aerodynamic power. The parrotlets repurpose lift and drag during these flights with lift-to-drag ratios below two. Such low ratios are within range of proto-wings, showing how avian precursors may have relied on drag to take off with flapping wings.

    View details for DOI 10.1038/s41467-019-13347-3

    View details for PubMedID 31767856

  • How lovebirds maneuver through lateral gusts with minimal visual information. Proceedings of the National Academy of Sciences of the United States of America Quinn, D., Kress, D., Chang, E., Stein, A., Wegrzynski, M., Lentink, D. 2019


    Flying birds maneuver effectively through lateral gusts, even when gust speeds are as high as flight speeds. What information birds use to sense gusts and how they compensate is largely unknown. We found that lovebirds can maneuver through 45° lateral gusts similarly well in forest-, lake-, and cave-like visual environments. Despite being diurnal and raised in captivity, the birds fly to their goal perch with only a dim point light source as a beacon, showing that they do not need optic flow or a visual horizon to maneuver. To accomplish this feat, lovebirds primarily yaw their bodies into the gust while fixating their head on the goal using neck angles of up to 30°. Our corroborated model for proportional yaw reorientation and speed control shows how lovebirds can compensate for lateral gusts informed by muscle proprioceptive cues from neck twist. The neck muscles not only stabilize the lovebirds' visual and inertial head orientations by compensating low-frequency body maneuvers, but also attenuate faster 3D wingbeat-induced perturbations. This head stabilization enables the vestibular system to sense the direction of gravity. Apparently, the visual horizon can be replaced by a gravitational horizon to inform the observed horizontal gust compensation maneuvers in the dark. Our scaling analysis shows how this minimal sensorimotor solution scales favorably for bigger birds, offering local wind angle feedback within a wingbeat. The way lovebirds glean wind orientation may thus inform minimal control algorithms that enable aerial robots to maneuver in similar windy and dark environments.

    View details for DOI 10.1073/pnas.1903422116

    View details for PubMedID 31289235

  • Biomechanics of hover performance in Neotropical hummingbirds versus bats. Science advances Ingersoll, R., Haizmann, L., Lentink, D. 2018; 4 (9): eaat2980


    Hummingbirds and nectar bats are the only vertebrates that are specialized for hovering in front of flowers to forage nectar. How their aerodynamic performance compares is, however, unclear. To hover, hummingbirds consistently generate about a quarter of the vertical aerodynamic force required to support their body weight during the upstroke. In contrast, generalist birds in slow hovering flight generate little upstroke weight support. We report that nectar bats also generate elevated weight support during the upstroke compared to generalist bats. Comparing 20 Neotropical species, we show how nectarivorous birds and bats converged on this ability by inverting their respective feathered and membrane wings more than species with other diets. However, while hummingbirds converged on an efficient horizontal wingbeat to mostly generate lift, bats rely on lift and drag during the downstroke to fully support their body weight. Furthermore, whereas the ability of nectar bats to aerodynamically support their body weight during the upstroke is elevated, it is much smaller than that of hummingbirds. Bats compensate by generating more aerodynamic weight support during their extended downstroke. Although, in principle, it requires more aerodynamic power to hover using this method, bats have adapted by evolving much larger wings for their body weight. Therefore, the net aerodynamic induced power required to hover is similar among hummingbirds and bats per unit body mass. This mechanistic insight into how feathered wings and membrane wings ultimately require similar aerodynamic power to hover may inform analogous design trade-offs in aerial robots.

    View details for PubMedID 30263957

  • How birds direct impulse to minimize the energetic cost of foraging flight. Science advances Chin, D. D., Lentink, D. 2017; 3 (5)


    Birds frequently hop and fly between tree branches to forage. To determine the mechanical energy trade-offs of their bimodal locomotion, we rewarded four Pacific parrotlets with a seed for flying voluntarily between instrumented perches inside a new aerodynamic force platform. By integrating direct measurements of both leg and wing forces with kinematics in a bimodal long jump and flight model, we discovered that parrotlets direct their leg impulse to minimize the mechanical energy needed to forage over different distances and inclinations. The bimodal locomotion model further shows how even a small lift contribution from a single proto-wingbeat would have significantly lengthened the long jump of foraging arboreal dinosaurs. These avian bimodal locomotion strategies can also help robots traverse cluttered environments more effectively.

    View details for DOI 10.1126/sciadv.1603041

    View details for PubMedID 28560342

  • In vivo recording of aerodynamic force with an aerodynamic force platform: from drones to birds. Journal of the Royal Society, Interface / the Royal Society Lentink, D., Haselsteiner, A. F., Ingersoll, R. 2015; 12 (104)


    Flapping wings enable flying animals and biomimetic robots to generate elevated aerodynamic forces. Measurements that demonstrate this capability are based on experiments with tethered robots and animals, and indirect force calculations based on measured kinematics or airflow during free flight. Remarkably, there exists no method to measure these forces directly during free flight. Such in vivo recordings in freely behaving animals are essential to better understand the precise aerodynamic function of their flapping wings, in particular during the downstroke versus upstroke. Here, we demonstrate a new aerodynamic force platform (AFP) for non-intrusive aerodynamic force measurement in freely flying animals and robots. The platform encloses the animal or object that generates fluid force with a physical control surface, which mechanically integrates the net aerodynamic force that is transferred to the earth. Using a straightforward analytical solution of the Navier-Stokes equation, we verified that the method is accurate. We subsequently validated the method with a quadcopter that is suspended in the AFP and generates unsteady thrust profiles. These independent measurements confirm that the AFP is indeed accurate. We demonstrate the effectiveness of the AFP by studying aerodynamic weight support of a freely flying bird in vivo. These measurements confirm earlier findings based on kinematics and flow measurements, which suggest that the avian downstroke, not the upstroke, is primarily responsible for body weight support during take-off and landing.

    View details for DOI 10.1098/rsif.2014.1283

    View details for PubMedID 25589565

  • Leading-edge vortices elevate lift of autorotating plant seeds. Science Lentink, D., van Dickson, W., B., van Leeuwen, J., L., Dickinson, M., H. 2009; 324: 1438 – 1440
  • Rotational accelerations stabilize leading edge vortices on revolving fly wings. J. Exp. Biol. Lentink, D., Dickinson, M., H. 2009; 212: 2705 – 2719
  • How swifts control their glide performance with morphing wings. Nature Lentink, D., Müller, U., K., Stamhuis, E., J., de Kat, R., van Gestel, W., Veldhuis, L., L.M. 2007; 446: 1082 – 1085
  • Design Principles and Function of Mechanical Fasteners in Nature and Technology APPLIED MECHANICS REVIEWS Jeffries, L., Lentink, D. 2020; 72 (5)

    View details for DOI 10.1115/1.4048448

    View details for Web of Science ID 000578498200002

  • The aerodynamic force platform as an ergometer. The Journal of experimental biology Deetjen, M. E., Chin, D. D., Lentink, D. 2020


    Animal flight requires aerodynamic power, which is challenging to determine accurately in vivo Existing methods rely on approximate calculations based on wake flow field measurements, inverse dynamics approaches, or invasive muscle physiological recordings. In contrast, the external mechanical work required for terrestrial locomotion can be determined more directly by using a force platform as an ergometer. Based on an extension of the recent invention of the aerodynamic force platform, we now present a more direct method to determine the in vivo aerodynamic power by taking the dot product of the aerodynamic force vector on the wing with the representative wing velocity vector based on kinematics and morphology. We demonstrate this new method by studying a slowly flying dove, but it can be applied more generally across flying and swimming animals as well as animals that locomote over water surfaces. Finally, our mathematical framework also works for power analyses based on flow field measurements.

    View details for DOI 10.1242/jeb.220475

    View details for PubMedID 32253285

  • Soft biohybrid morphing wings with feathers underactuated by wrist and finger motion. Science robotics Chang, E., Matloff, L. Y., Stowers, A. K., Lentink, D. 2020; 5 (38)


    Since the Wright Flyer, engineers have strived to develop flying machines with morphing wings that can control flight as deftly as birds. Birds morph their wing planform parameters simultaneously-including sweep, span, and area-in a way that has proven to be particularly challenging to embody robotically. Previous solutions have primarily centered around the classical aerospace paradigm of controlling every degree of freedom to ensure predictable performance, but underperform compared with birds. To understand how birds accomplish wing morphing, we measured the kinematics of wing flexion and extension in common pigeons, Columba livia The skeletal and feather kinematics show that the 20 primary and 20 secondary feathers are coordinated via approximately linear transfer functions controlled by wrist and finger motion. To replicate this control principle in a robot, we developed a biohybrid morphing wing with real feathers to understand the underlying design principles. The outcome, PigeonBot, embodies 42 degrees of freedom that control the position of 40 elastically connected feathers via four servo-actuated wrist and finger joints. Our flight tests demonstrate that the soft feathered wings morph rapidly and robustly under aerodynamic loading. They not only enable wing morphing but also make robot interactions safer, the wing more robust to crashing, and the wing reparable via "preening." In flight tests, we found that both asymmetric wrist and finger motion can initiate turn maneuvers-evidence that birds may use their fingers to steer in flight.

    View details for DOI 10.1126/scirobotics.aay1246

    View details for PubMedID 33022590

  • Fluid moment and force measurement based on control surface integration EXPERIMENTS IN FLUIDS Chin, D. D., Lentink, D. 2019; 61 (1)
  • Birds land reliably on complex surfaces by adapting their foot-surface interactions upon contact. eLife Roderick, W. R., Chin, D. D., Cutkosky, M. R., Lentink, D. 2019; 8


    Birds land on a wide range of complex surfaces, yet it is unclear how they grasp a perch reliably. Here, we show how Pacific parrotlets exhibit stereotyped leg and wing dynamics regardless of perch diameter and texture, but foot, toe, and claw kinematics become surface-specific upon touchdown. A new dynamic grasping model, which integrates our detailed measurements, reveals how birds stabilize their grasp. They combine predictable toe pad friction with probabilistic friction from their claws, which they drag to find surface asperities-dragging further when they can squeeze less. Remarkably, parrotlet claws can undergo superfast movements, within 1-2 ms, on moderately slippery surfaces to find more secure asperities when necessary. With this strategy, they first ramp up safety margins by squeezing before relaxing their grasp. The model further shows it is advantageous to be small for stable perching when high friction relative to normal force is required because claws can find more usable surface, but this trend reverses when required friction shrinks. This explains how many animals and robots may grasp complex surfaces reliably.

    View details for DOI 10.7554/eLife.46415

    View details for PubMedID 31385573

  • How Hummingbirds Reorient Forces During Maneuvering Flight Hightower, B., Ingersoll, R., Shorr, D., Chin, D., Lentink, D., Hightower, B. OXFORD UNIV PRESS INC. 2019: E100
  • A Bird's-Eye View of Regulatory, Animal Care, and Training Considerations Regarding Avian Flight Research. Comparative medicine Baker, S. W., Tucci, E. R., Felt, S. A., Zehnder, A., Lentink, D., Vilches-Moure, J. G. 2019


    A thorough understanding of how animals fly is a central goal of many scientific disciplines. Birds are a commonly usedmodel organism for flight research. The success of this model requires studying healthy and naturally flying birds in a laboratory setting. This use of a nontraditional laboratory animal species presents unique challenges to animal care staff and researchers alike. Here we review regulatory, animal care, and training considerations associated with avian flight research.

    View details for DOI 10.30802/AALAS-CM-18-000033

    View details for PubMedID 30764892

  • How the hummingbird wingbeat is tuned for efficient hovering. The Journal of experimental biology Ingersoll, R., Lentink, D. 2018; 221 (Pt 20)


    Both hummingbirds and insects flap their wings to hover. Some insects, like fruit flies, improve efficiency by lifting their body weight equally over the upstroke and downstroke, while utilizing elastic recoil during stroke reversal. It is unclear whether hummingbirds converged on a similar elastic storage solution, because of asymmetries in their lift generation and specialized flight muscle apparatus. The muscles are activated a quarter of a stroke earlier than in larger birds, and contract superfast, which cannot be explained by previous stroke-averaged analyses. We measured the aerodynamic force and kinematics of Anna's hummingbirds to resolve wing torque and power within the wingbeat. Comparing these wingbeat-resolved aerodynamic weight support measurements with those of fruit flies, hawk moths and a generalist bird, the parrotlet, we found that hummingbirds have about the same low induced power losses as the two insects, lower than that of the generalist bird in slow hovering flight. Previous analyses emphasized how bird flight muscles have to overcome wing drag midstroke. We found that high wing inertia revises this for hummingbirds - the pectoralis has to coordinate upstroke to downstroke reversal while the supracoracoideus coordinates downstroke to upstroke reversal. Our mechanistic analysis aligns with all previous muscle recordings and shows how early activation helps furnish elastic recoil through stroke reversal to stay within the physiological limits of muscles. Our findings thus support Weis-Fogh's hypothesis that flies and hummingbirds have converged on a mechanically efficient wingbeat to meet the high energetic demands of hovering flight. These insights can help improve the efficiency of flapping robots.

    View details for PubMedID 30323114

  • Accurate fluid force measurement based on control surface integration EXPERIMENTS IN FLUIDS Lentink, D. 2018; 59 (1)
  • Adaptive control of turbulence intensity is accelerated by frugal flow sampling. Journal of the Royal Society, Interface Quinn, D. B., van Halder, Y., Lentink, D. 2017; 14 (136)


    The aerodynamic performance of vehicles and animals, as well as the productivity of turbines and energy harvesters, depends on the turbulence intensity of the incoming flow. Previous studies have pointed at the potential benefits of active closed-loop turbulence control. However, it is unclear what the minimal sensory and algorithmic requirements are for realizing this control. Here we show that very low-bandwidth anemometers record sufficient information for an adaptive control algorithm to converge quickly. Our online Newton-Raphson algorithm tunes the turbulence in a recirculating wind tunnel by taking readings from an anemometer in the test section. After starting at 9% turbulence intensity, the algorithm converges on values ranging from 10% to 45% in less than 12 iterations within 1% accuracy. By down-sampling our measurements, we show that very-low-bandwidth anemometers record sufficient information for convergence. Furthermore, down-sampling accelerates convergence by smoothing gradients in turbulence intensity. Our results explain why low-bandwidth anemometers in engineering and mechanoreceptors in biology may be sufficient for adaptive control of turbulence intensity. Finally, our analysis suggests that, if certain turbulent eddy sizes are more important to control than others, frugal adaptive control schemes can be particularly computationally effective for improving performance.

    View details for PubMedID 29118116

  • The biomechanical origin of extreme wing allometry in hummingbirds NATURE COMMUNICATIONS Skandalis, D. A., Segre, P. S., Bahlman, J. W., Groom, D. E., Welch, K. C., Witt, C. C., McGuire, J. A., Dudley, R., Lentink, D., Altshuler, D. L. 2017; 8: 1047


    Flying animals of different masses vary widely in body proportions, but the functional implications of this variation are often unclear. We address this ambiguity by developing an integrative allometric approach, which we apply here to hummingbirds to examine how the physical environment, wing morphology and stroke kinematics have contributed to the evolution of their highly specialised flight. Surprisingly, hummingbirds maintain constant wing velocity despite an order of magnitude variation in body weight; increased weight is supported solely through disproportionate increases in wing area. Conversely, wing velocity increases with body weight within species, compensating for lower relative wing area in larger individuals. By comparing inter- and intraspecific allometries, we find that the extreme wing area allometry of hummingbirds is likely an adaptation to maintain constant burst flight capacity and induced power requirements with increasing weight. Selection for relatively large wings simultaneously maximises aerial performance and minimises flight costs, which are essential elements of humming bird life history.

    View details for PubMedID 29051535

  • How pigeons couple three-dimensional elbow and wrist motion to morph their wings JOURNAL OF THE ROYAL SOCIETY INTERFACE Stowers, A. K., Matloff, L. Y., Lentink, D. 2017; 14 (133)


    Birds change the shape and area of their wings to an exceptional degree, surpassing insects, bats and aircraft in their ability to morph their wings for a variety of tasks. This morphing is governed by a musculoskeletal system, which couples elbow and wrist motion. Since the discovery of this effect in 1839, the planar 'drawing parallels' mechanism has been used to explain the coupling. Remarkably, this mechanism has never been corroborated from quantitative motion data. Therefore, we measured how the wing skeleton of a pigeon (Columba livia) moves during morphing. Despite earlier planar assumptions, we found that the skeletal motion paths are highly three-dimensional and do not lie in the anatomical plane, ruling out the 'drawing parallels' mechanism. Furthermore, micro-computed tomography scans in seven consecutive poses show how the two wrist bones contribute to morphing, particularly the sliding ulnare. From these data, we infer the joint types for all six bones that form the wing morphing mechanism and corroborate the most parsimonious mechanism based on least-squares error minimization. Remarkably, the algorithm shows that all optimal four-bar mechanisms either lock, are unable to track the highly three-dimensional bone motion paths, or require the radius and ulna to cross for accuracy, which is anatomically unrealistic. In contrast, the algorithm finds that a six-bar mechanism recreates the measured motion accurately with a parallel radius and ulna and a sliding ulnare. This revises our mechanistic understanding of how birds morph their wings, and offers quantitative inspiration for engineering morphing wings.

    View details for PubMedID 28794161

    View details for PubMedCentralID PMC5582118

  • Inspiration for wing design: how forelimb specialization enables active flight in modern vertebrates. Journal of the Royal Society, Interface Chin, D. D., Matloff, L. Y., Stowers, A. K., Tucci, E. R., Lentink, D. 2017; 14 (131)


    Harnessing flight strategies refined by millions of years of evolution can help expedite the design of more efficient, manoeuvrable and robust flying robots. This review synthesizes recent advances and highlights remaining gaps in our understanding of how bird and bat wing adaptations enable effective flight. Included in this discussion is an evaluation of how current robotic analogues measure up to their biological sources of inspiration. Studies of vertebrate wings have revealed skeletal systems well suited for enduring the loads required during flight, but the mechanisms that drive coordinated motions between bones and connected integuments remain ill-described. Similarly, vertebrate flight muscles have adapted to sustain increased wing loading, but a lack of in vivo studies limits our understanding of specific muscular functions. Forelimb adaptations diverge at the integument level, but both bird feathers and bat membranes yield aerodynamic surfaces with a level of robustness unparalleled by engineered wings. These morphological adaptations enable a diverse range of kinematics tuned for different flight speeds and manoeuvres. By integrating vertebrate flight specializations-particularly those that enable greater robustness and adaptability-into the design and control of robotic wings, engineers can begin narrowing the wide margin that currently exists between flying robots and vertebrates. In turn, these robotic wings can help biologists create experiments that would be impossible in vivo.

    View details for DOI 10.1098/rsif.2017.0240

    View details for PubMedID 28592663

  • High-speed surface reconstruction of a flying bird using structured light. journal of experimental biology Deetjen, M. E., Biewener, A. A., Lentink, D. 2017; 220: 1956-1961


    Birds fly effectively and maneuver nimbly by dynamically changing the shape of their wings during each wingbeat. These shape changes have yet to be quantified automatically at high temporal and spatial resolution. Therefore, we developed a custom 3D surface reconstruction method, which uses a high-speed camera to identify spatially encoded binary striped patterns that are projected on a flying bird. This non-invasive structured-light method allows automated 3D reconstruction of each stand-alone frame and can be extended to multiple views. We demonstrate this new technique by automatically reconstructing the dorsal surface of a parrotlet wing at 3200 frames s(-1) during flapping flight. From this shape we analyze key parameters such as wing twist and angle of attack distribution. While our binary 'single-shot' algorithm is demonstrated by quantifying dynamic shape changes of a flying bird, it is generally applicable to moving animals, plants and deforming objects.

    View details for DOI 10.1242/jeb.149708

    View details for PubMedID 28348041

  • A new low-turbulence wind tunnel for animal and small vehicle flight experiments ROYAL SOCIETY OPEN SCIENCE Quinn, D. B., Watts, A., Nagle, T., Lentink, D. 2017; 4 (3)


    Our understanding of animal flight benefits greatly from specialized wind tunnels designed for flying animals. Existing facilities can simulate laminar flow during straight, ascending and descending flight, as well as at different altitudes. However, the atmosphere in which animals fly is even more complex. Flow can be laminar and quiet at high altitudes but highly turbulent near the ground, and gusts can rapidly change wind speed. To study flight in both laminar and turbulent environments, a multi-purpose wind tunnel for studying animal and small vehicle flight was built at Stanford University. The tunnel is closed-circuit and can produce airspeeds up to 50 m s(-1) in a rectangular test section that is 1.0 m wide, 0.82 m tall and 1.73 m long. Seamless honeycomb and screens in the airline together with a carefully designed contraction reduce centreline turbulence intensities to less than or equal to 0.030% at all operating speeds. A large diameter fan and specialized acoustic treatment allow the tunnel to operate at low noise levels of 76.4 dB at 20 m s(-1). To simulate high turbulence, an active turbulence grid can increase turbulence intensities up to 45%. Finally, an open jet configuration enables stereo high-speed fluoroscopy for studying musculoskeletal control in turbulent flow.

    View details for DOI 10.1098/rsos.160960

    View details for Web of Science ID 000398107700035

    View details for PubMedID 28405384

  • Lift calculations based on accepted wake models for animal flight are inconsistent and sensitive to vortex dynamics BIOINSPIRATION & BIOMIMETICS Gutierrez, E., Quinn, D. B., Chin, D. D., Lentink, D. 2017; 12 (1)
  • Touchdown to take-off: at the interface of flight and surface locomotion INTERFACE FOCUS Roderick, W. R., Cutkosky, M. R., Lentink, D. 2017; 7 (1)


    Small aerial robots are limited to short mission times because aerodynamic and energy conversion efficiency diminish with scale. One way to extend mission times is to perch, as biological flyers do. Beyond perching, small robot flyers benefit from manoeuvring on surfaces for a diverse set of tasks, including exploration, inspection and collection of samples. These opportunities have prompted an interest in bimodal aerial and surface locomotion on both engineered and natural surfaces. To accomplish such novel robot behaviours, recent efforts have included advancing our understanding of the aerodynamics of surface approach and take-off, the contact dynamics of perching and attachment and making surface locomotion more efficient and robust. While current aerial robots show promise, flying animals, including insects, bats and birds, far surpass them in versatility, reliability and robustness. The maximal size of both perching animals and robots is limited by scaling laws for both adhesion and claw-based surface attachment. Biomechanists can use the current variety of specialized robots as inspiration for probing unknown aspects of bimodal animal locomotion. Similarly, the pitch-up landing manoeuvres and surface attachment techniques of animals can offer an evolutionary design guide for developing robots that perch on more diverse and complex surfaces.

    View details for DOI 10.1098/rsfs.2016.0094

    View details for Web of Science ID 000391694100017

    View details for PubMedID 28163884

    View details for PubMedCentralID PMC5206611

  • Lift calculations based on accepted wake models for animal flight are inconsistent and sensitive to vortex dynamics. Bioinspiration & biomimetics Gutierrez, E., Quinn, D. B., Chin, D. D., Lentink, D. 2016; 12 (1): 016004-?


    There are three common methods for calculating the lift generated by a flying animal based on the measured airflow in the wake. However, these methods might not be accurate according to computational and robot-based studies of flapping wings. Here we test this hypothesis for the first time for a slowly flying Pacific parrotlet in still air using stereo particle image velocimetry recorded at 1000 Hz. The bird was trained to fly between two perches through a laser sheet wearing laser safety goggles. We found that the wingtip vortices generated during mid-downstroke advected down and broke up quickly, contradicting the frozen turbulence hypothesis typically assumed in animal flight experiments. The quasi-steady lift at mid-downstroke was estimated based on the velocity field by applying the widely used Kutta-Joukowski theorem, vortex ring model, and actuator disk model. The calculated lift was found to be sensitive to the applied model and its different parameters, including vortex span and distance between the bird and laser sheet-rendering these three accepted ways of calculating weight support inconsistent. The three models predict different aerodynamic force values mid-downstroke compared to independent direct measurements with an aerodynamic force platform that we had available for the same species flying over a similar distance. Whereas the lift predictions of the Kutta-Joukowski theorem and the vortex ring model stayed relatively constant despite vortex breakdown, their values were too low. In contrast, the actuator disk model predicted lift reasonably accurately before vortex breakdown, but predicted almost no lift during and after vortex breakdown. Some of these limitations might be better understood, and partially reconciled, if future animal flight studies report lift calculations based on all three quasi-steady lift models instead. This would also enable much needed meta studies of animal flight to derive bioinspired design principles for quasi-steady lift generation with flapping wings.

    View details for PubMedID 27921999

  • Fruit fly scale robots can hover longer with flapping wings than with spinning wings. Journal of the Royal Society, Interface Hawkes, E. W., Lentink, D. 2016; 13 (123)


    Hovering flies generate exceptionally high lift, because their wings generate a stable leading edge vortex. Micro flying robots with a similar wing design can generate similar high lift by either flapping or spinning their wings. While it requires less power to spin a wing, the overall efficiency depends also on the actuator system driving the wing. Here, we present the first holistic analysis to calculate how long a fly-inspired micro robot can hover with flapping versus spinning wings across scales. We integrate aerodynamic data with data-driven scaling laws for actuator, electronics and mechanism performance from fruit fly to hummingbird scales. Our analysis finds that spinning wings driven by rotary actuators are superior for robots with wingspans similar to hummingbirds, yet flapping wings driven by oscillatory actuators are superior at fruit fly scale. This crossover is driven by the reduction in performance of rotary compared with oscillatory actuators at smaller scale. Our calculations emphasize that a systems-level analysis is essential for trading-off flapping versus spinning wings for micro flying robots.

    View details for PubMedID 27707903

  • Flapping wing aerodynamics: from insects to vertebrates JOURNAL OF EXPERIMENTAL BIOLOGY Chin, D. D., Lentink, D. 2016; 219 (7): 920-932

    View details for DOI 10.1242/jeb.042317

    View details for Web of Science ID 000373212600011

  • Flapping wing aerodynamics: from insects to vertebrates. journal of experimental biology Chin, D. D., Lentink, D. 2016; 219: 920-932


    More than a million insects and approximately 11,000 vertebrates utilize flapping wings to fly. However, flapping flight has only been studied in a few of these species, so many challenges remain in understanding this form of locomotion. Five key aerodynamic mechanisms have been identified for insect flight. Among these is the leading edge vortex, which is a convergent solution to avoid stall for insects, bats and birds. The roles of the other mechanisms - added mass, clap and fling, rotational circulation and wing-wake interactions - have not yet been thoroughly studied in the context of vertebrate flight. Further challenges to understanding bat and bird flight are posed by the complex, dynamic wing morphologies of these species and the more turbulent airflow generated by their wings compared with that observed during insect flight. Nevertheless, three dimensionless numbers that combine key flow, morphological and kinematic parameters - the Reynolds number, Rossby number and advance ratio - govern flapping wing aerodynamics for both insects and vertebrates. These numbers can thus be used to organize an integrative framework for studying and comparing animal flapping flight. Here, we provide a roadmap for developing such a framework, highlighting the aerodynamic mechanisms that remain to be quantified and compared across species. Ultimately, incorporating complex flight maneuvers, environmental effects and developmental stages into this framework will also be essential to advancing our understanding of the biomechanics, movement ecology and evolution of animal flight.

    View details for DOI 10.1242/jeb.042317

    View details for PubMedID 27030773

  • The biophysics of bird flight: functional relationships integrate aerodynamics, morphology, kinematics, muscles, and sensors CANADIAN JOURNAL OF ZOOLOGY Altshuler, D. L., Bahlman, J. W., Dakin, R., Gaede, A. H., Goller, B., Lentink, D., Segre, P. S., Skandalis, D. A. 2015; 93 (12): 961-975
  • Feather roughness reduces flow separation during low Reynolds number glides of swifts JOURNAL OF EXPERIMENTAL BIOLOGY van Bokhorst, E., de Kat, R., Elsinga, G. E., Lentink, D. 2015; 218 (20): 3179-3191

    View details for DOI 10.1242/jeb.121426

    View details for PubMedID 26347563

  • The role of passive avian head stabilization in flapping flight. Journal of the Royal Society, Interface / the Royal Society Pete, A. E., Kress, D., Dimitrov, M. A., Lentink, D. 2015; 12 (110)

    View details for DOI 10.1098/rsif.2015.0508

    View details for PubMedID 26311316

  • The role of passive avian head stabilization in flapping flight. Journal of the Royal Society, Interface / the Royal Society Pete, A. E., Kress, D., Dimitrov, M. A., Lentink, D. 2015; 12 (110)


    Birds improve vision by stabilizing head position relative to their surroundings, while their body is forced up and down during flapping flight. Stabilization is facilitated by compensatory motion of the sophisticated avian head-neck system. While relative head motion has been studied in stationary and walking birds, little is known about how birds accomplish head stabilization during flapping flight. To unravel this, we approximate the avian neck with a linear mass-spring-damper system for vertical displacements, analogous to proven head stabilization models for walking humans. We corroborate the model's dimensionless natural frequency and damping ratios from high-speed video recordings of whooper swans (Cygnus cygnus) flying over a lake. The data show that flap-induced body oscillations can be passively attenuated through the neck. We find that the passive model robustly attenuates large body oscillations, even in response to head mass and gust perturbations. Our proof of principle shows that bird-inspired drones with flapping wings could record better images with a swan-inspired passive camera suspension.

    View details for DOI 10.1098/rsif.2015.0508

    View details for PubMedID 26311316

  • Power reduction and the radial limit of stall delay in revolving wings of different aspect ratio JOURNAL OF THE ROYAL SOCIETY INTERFACE Kruyt, J. W., Van Heijst, G. F., Altshuler, D. L., Lentink, D. 2015; 12 (105)


    Airplanes and helicopters use high aspect ratio wings to reduce the power required to fly, but must operate at low angle of attack to prevent flow separation and stall. Animals capable of slow sustained flight, such as hummingbirds, have low aspect ratio wings and flap their wings at high angle of attack without stalling. Instead, they generate an attached vortex along the leading edge of the wing that elevates lift. Previous studies have demonstrated that this vortex and high lift can be reproduced by revolving the animal wing at the same angle of attack. How do flapping and revolving animal wings delay stall and reduce power? It has been hypothesized that stall delay derives from having a short radial distance between the shoulder joint and wing tip, measured in chord lengths. This non-dimensional measure of wing length represents the relative magnitude of inertial forces versus rotational accelerations operating in the boundary layer of revolving and flapping wings. Here we show for a suite of aspect ratios, which represent both animal and aircraft wings, that the attachment of the leading edge vortex on a revolving wing is determined by wing aspect ratio, defined with respect to the centre of revolution. At high angle of attack, the vortex remains attached when the local radius is shorter than four chord lengths and separates outboard on higher aspect ratio wings. This radial stall limit explains why revolving high aspect ratio wings (of helicopters) require less power compared with low aspect ratio wings (of hummingbirds) at low angle of attack and vice versa at high angle of attack.

    View details for DOI 10.1098/rsif.2015.0051

    View details for Web of Science ID 000351230700018

    View details for PubMedID 25788539

    View details for PubMedCentralID PMC4387534

  • Folding in and out: passive morphing in flapping wings. Bioinspiration & biomimetics Stowers, A. K., Lentink, D. 2015; 10 (2): 025001-?


    We present a new mechanism for passive wing morphing of flapping wings inspired by bat and bird wing morphology. The mechanism consists of an unactuated hand wing connected to the arm wing with a wrist joint. Flapping motion generates centrifugal accelerations in the hand wing, forcing it to unfold passively. Using a robotic model in hover, we made kinematic measurements of unfolding kinematics as functions of the non-dimensional wingspan fold ratio (2-2.5) and flapping frequency (5-17 Hz) using stereo high-speed cameras. We find that the wings unfold passively within one to two flaps and remain unfolded with only small amplitude oscillations. To better understand the passive dynamics, we constructed a computer model of the unfolding process based on rigid body dynamics, contact models, and aerodynamic correlations. This model predicts the measured passive unfolding within about one flap and shows that unfolding is driven by centrifugal acceleration induced by flapping. The simulations also predict that relative unfolding time only weakly depends on flapping frequency and can be reduced to less than half a wingbeat by increasing flapping amplitude. Subsequent dimensional analysis shows that the time required to unfold passively is of the same order of magnitude as the flapping period. This suggests that centrifugal acceleration can drive passive unfolding within approximately one wingbeat in small and large wings. Finally, we show experimentally that passive unfolding wings can withstand impact with a branch, by first folding and then unfolding passively. This mechanism enables flapping robots to squeeze through clutter without sophisticated control. Passive unfolding also provides a new avenue in morphing wing design that makes future flapping morphing wings possibly more energy efficient and light-weight. Simultaneously these results point to possible inertia driven, and therefore metabolically efficient, control strategies in bats and birds to morph or recover within a beat.

    View details for DOI 10.1088/1748-3190/10/2/025001

    View details for PubMedID 25807583

  • Folding in and out: passive morphing in flapping wings BIOINSPIRATION & BIOMIMETICS Stowers, A. K., Lentink, D. 2015; 10 (2)
  • How Lovebirds Maneuver Rapidly Using Super-Fast Head Saccades and Image Feature Stabilization. PloS one Kress, D., van Bokhorst, E., Lentink, D. 2015; 10 (6)


    Diurnal flying animals such as birds depend primarily on vision to coordinate their flight path during goal-directed flight tasks. To extract the spatial structure of the surrounding environment, birds are thought to use retinal image motion (optical flow) that is primarily induced by motion of their head. It is unclear what gaze behaviors birds perform to support visuomotor control during rapid maneuvering flight in which they continuously switch between flight modes. To analyze this, we measured the gaze behavior of rapidly turning lovebirds in a goal-directed task: take-off and fly away from a perch, turn on a dime, and fly back and land on the same perch. High-speed flight recordings revealed that rapidly turning lovebirds perform a remarkable stereotypical gaze behavior with peak saccadic head turns up to 2700 degrees per second, as fast as insects, enabled by fast neck muscles. In between saccades, gaze orientation is held constant. By comparing saccade and wingbeat phase, we find that these super-fast saccades are coordinated with the downstroke when the lateral visual field is occluded by the wings. Lovebirds thus maximize visual perception by overlying behaviors that impair vision, which helps coordinate maneuvers. Before the turn, lovebirds keep a high contrast edge in their visual midline. Similarly, before landing, the lovebirds stabilize the center of the perch in their visual midline. The perch on which the birds land swings, like a branch in the wind, and we find that retinal size of the perch is the most parsimonious visual cue to initiate landing. Our observations show that rapidly maneuvering birds use precisely timed stereotypic gaze behaviors consisting of rapid head turns and frontal feature stabilization, which facilitates optical flow based flight control. Similar gaze behaviors have been reported for visually navigating humans. This finding can inspire more effective vision-based autopilots for drones.

    View details for DOI 10.1371/journal.pone.0129287

    View details for PubMedID 26107413

    View details for PubMedCentralID PMC4481315

  • How Lovebirds Maneuver Rapidly Using Super-Fast Head Saccades and Image Feature Stabilization. PloS one Kress, D., van Bokhorst, E., Lentink, D. 2015; 10 (6)

    View details for DOI 10.1371/journal.pone.0129287

    View details for PubMedID 26107413

  • Hummingbird wing efficacy depends on aspect ratio and compares with helicopter rotors. Journal of the Royal Society, Interface / the Royal Society Kruyt, J. W., Quicazán-Rubio, E. M., Van Heijst, G. F., Altshuler, D. L., Lentink, D. 2014; 11 (99)


    Hummingbirds are the only birds that can sustain hovering. This unique flight behaviour comes, however, at high energetic cost. Based on helicopter and aeroplane design theory, we expect that hummingbird wing aspect ratio (AR), which ranges from about 3.0 to 4.5, determines aerodynamic efficacy. Previous quasi-steady experiments with a wing spinner set-up provide no support for this prediction. To test this more carefully, we compare the quasi-steady hover performance of 26 wings, from 12 hummingbird taxa. We spun the wings at angular velocities and angles of attack that are representative for every species and measured lift and torque more precisely. The power (aerodynamic torque × angular velocity) required to lift weight depends on aerodynamic efficacy, which is measured by the power factor. Our comparative analysis shows that AR has a modest influence on lift and drag forces, as reported earlier, but interspecific differences in power factor are large. During the downstroke, the power required to hover decreases for larger AR wings at the angles of attack at which hummingbirds flap their wings (p < 0.05). Quantitative flow visualization demonstrates that variation in hover power among hummingbird wings is driven by similar stable leading edge vortices that delay stall during the down- and upstroke. A side-by-side aerodynamic performance comparison of hummingbird wings and an advanced micro helicopter rotor shows that they are remarkably similar.

    View details for DOI 10.1098/rsif.2014.0585

    View details for PubMedID 25079868

  • Gliding Swifts Attain Laminar Flow over Rough Wings PLOS ONE Lentink, D., de Kat, R. 2014; 9 (6)
  • Bioinspired flight control. Bioinspiration & biomimetics Lentink, D. 2014; 9 (2): 020301-?

    View details for DOI 10.1088/1748-3182/9/2/020301

    View details for PubMedID 24854957

  • Gliding Swifts Attain Laminar Flow over Rough Wings. PloS one Lentink, D., de Kat, R. 2014; 9 (6)


    Swifts are among the most aerodynamically refined gliding birds. However, the overlapping vanes and protruding shafts of their primary feathers make swift wings remarkably rough for their size. Wing roughness height is 1-2% of chord length on the upper surface--10,000 times rougher than sailplane wings. Sailplanes depend on extreme wing smoothness to increase the area of laminar flow on the wing surface and minimize drag for extended glides. To understand why the swift does not rely on smooth wings, we used a stethoscope to map laminar flow over preserved wings in a low-turbulence wind tunnel. By combining laminar area, lift, and drag measurements, we show that average area of laminar flow on swift wings is 69% (n = 3; std 13%) of their total area during glides that maximize flight distance and duration--similar to high-performance sailplanes. Our aerodynamic analysis indicates that swifts attain laminar flow over their rough wings because their wing size is comparable to the distance the air travels (after a roughness-induced perturbation) before it transitions from laminar to turbulent. To interpret the function of swift wing roughness, we simulated its effect on smooth model wings using physical models. This manipulation shows that laminar flow is reduced and drag increased at high speeds. At the speeds at which swifts cruise, however, swift-like roughness prolongs laminar flow and reduces drag. This feature gives small birds with rudimentary wings an edge during the evolution of glide performance.

    View details for DOI 10.1371/journal.pone.0099901

    View details for PubMedID 24964089

    View details for PubMedCentralID PMC4070913

  • Small aspect ratio differences impact hover efficacy among 12 hummingbird species Annual Meeting of the Society-for-Integrative-and-Comparative-Biology (SICB) Kruyt, J. W., Quicazan-Rubio, E. M., Van Heijst, G. J., Altshuler, D. L., Lentink, D. OXFORD UNIV PRESS INC. 2013: E118–E118
  • Flight Artists: An outreach project that enables the general public to film natural flight using the worlds most advanced high-speed camera Annual Meeting of the Society-for-Integrative-and-Comparative-Biology (SICB) Lentink, D., Fiaz, A. W. OXFORD UNIV PRESS INC. 2013: E124–E124
  • Flying like a fly. Nature Lentink, D. 2013

    View details for DOI 10.1038/nature12258

  • Vortex interactions with flapping wings and fins can be unpredictable. Biol. Lett. Lentink, D., van Heijst, G., J.F., Muijres, F., T., van Leeuwen, J., L. 2010

    View details for DOI 10.1098/rsbl.2009.0806.

  • Nature inspired flight – beyond the leap. Bioinspir. Biomim. Lentink, D., Biewener, A., A. 2010; 5
  • Structural analysis of a dragonfly wing. J. Exp. Mech. Jongerius, S., R., Lentink, D. 2010; 50: 1323-1334
  • Biofluiddynamic scaling of flapping, spinning and translating fins and wings. J. Exp. Biol. Lentink, D., Dickinson, M., H. 2009; 212: 2691 – 2704
  • Automated visual tracking for studying the ontogeny of zebrafish swimming. J. Exp. Biology. Fontaine, E., Lentink, D., Kranenbarg, S., Müller, U., K., van Leeuwen, J., L., Barr, A., H. 2008; 211: 1305 – 1316
  • Vortex-wake interactions of a flapping foil that models animal swimming and flight. J. Exp. Biology. Lentink, D., Muijres, F., T., Donker-Duyvis, F., J., van Leeuwen, J., L. 2008; 211: 267 – 273
  • Wake visualization of a heaving and pitching foil in a soap film. Exp. Fluids Muijres, F., T., Lentink, D. 2007; 43: 665 – 673
  • Turning on a Dime. Science Müller, U., K., Lentink, D. 2004; 306: 1899 – 1900