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. 2024 Jul 8;19(7):e0305084. doi: 10.1371/journal.pone.0305084

Generating controlled gust perturbations using vortex rings

Dipendra Gupta 1,2, Sanjay P Sane 3,*, Jaywant H Arakeri 2,*
Editor: Iman Borazjani4
PMCID: PMC11230548  PMID: 38976706

Abstract

To understand the locomotory mechanisms of flying and swimming animals, it is often necessary to develop assays that enable us to measure their responses to external gust perturbations. Typically, such measurements have been carried out using a variety of gusts which are difficult to control or characterize owing to their inherently turbulent nature. Here, we present a method of generating discrete gusts under controlled laboratory conditions in the form of a vortex rings which are well-characterized and highly controllable. We also provide the theoretical guidelines underlying the design of gust generators for specific applications. As a case study, we tested the efficacy of this method to study the flight response of freely-flying soldier flies Hermetia illucens. The vortex ring based method can be used to generate controlled gusts to study diverse phenomena ranging from a natural flight in insects to the artificial flight of insect-sized drones and micro-aerial vehicles.

Introduction

Flying animals often move in erratic environments facing discrete gusts or continuous turbulence [13]. Such gusts are experienced when there is a sudden and sharp change in wind speed, as typically encountered in the wakes of large objects or at the edge of convective disturbances. Organisms may also encounter continuous turbulence when flying in open environments or at great heights which is usually described using statistical approaches. Despite these unpredictable conditions, flying animals including insects, birds or bats can successfully control their flight in face of strong gusts [410]. For several years, researchers from various fields have tried to quantify the locomotory abilities of diverse animals under challenging conditions. For instance, ecologists are interested in understanding how locomotory ability sets the range over which animals can disperse or migrate [11]. Neurobiologists are interested in the mechanistic details of how animals may sense, process and respond to perturbations to their trajectories [12]. More recently, the field of bioinspired robotics has posed similar questions about the flight ability of their flappers and drones or swimming robots [13].

A key requirement for such studies is the ability to generate a well-characterized gust perturbation which can be used to determine how stable the flying or swimming animals or a robot is in face of a sudden change in ambient conditions. To achieve this, researchers have used several methods to generate turbulence and gusts in context of natural fliers. These include grid-generated turbulence [4, 7, 14], von-Karman vortices [5, 15, 16], compressed air jet [10, 17, 18] etc. These studies have provided keen insights into the general ways in which animals respond to turbulent gusts. Other methods are focused on either flow characterization alone or on the passive response of airfoils or drones [1921].

Here, we present a method that allows the delivery of precise and repeatable gusts, thereby enabling better control on the nature of perturbations encountered by animals in such experiments, and thus gain insight into the nature of their responses. The device presented here is capable of generating discrete, well-characterized, customized gusts using vortex rings. Vortex rings are typically generated by imparting an impulsive motion to a piston [22]. Vortex rings propagate with their own, self-induced velocity due to vorticity concentrated in their core region. A vortex ring is particularly effective in generating precise gusts, because of its sharp impulse and because vorticity is concentrated primarily in its core. This makes it possible to generate a high-speed perturbations while keeping the ring laminar, thereby eliminating any ambiguity in the flow characteristics. When turbulent gusts such as jets, grids, or bluff objects are placed upstream to perturb the flow to measure the response of the animals, their behaviour is a function of both mean flow and turbulent fluctuations making it difficult to interpret their response. In contrast to these methods, vortex rings are structured and relatively free from these effects. The flow physics of such a perturbation method is also well-understood, and hence the flow properties are highly controllable [23]. This method can be readily adapted to a variety of contexts ranging from aerial to aquatic locomotion. In this paper, we specifically use it to measure the responses of a freely flying soldier fly, Hermetia illucens as an example.

Vortex-generated gusts are different from freestream perturbations (e.g. von-Karman vortices) in which gusts superimpose over mean ambient flow, and which are well-suited for hovering insects. In contrast, the method presented here can be used to study the response of both hovering as well as freely-flying insects. Similar methods have been previously employed to study the response of hovering hawkmoths [24]. Here, we first provide a detailed description of the experimental setup to create such vortex rings and related gusts, and the spatio-temporal characterization of their flow properties, in addition to a detailed characterization of the flow properties in application to the animal locomotion which has been typically missing. As a case study, we next test this device on freely-flying soldier flies whose flight was perturbed using the gusts generated by this method. The wing span and the wing-tip speed of the insects dictated the size and velocity of the vortex ring in this study. We also outline the necessary theoretical details to estimate the flow properties of a vortex ring.

Methods and materials

The most common method of generating a vortex ring is using a piston-cylinder arrangement (Fig 1A). The size of the rings depends on the exit diameter (D0) of the cylindrical tube and the extent to which the piston has traversed (referred to as stroke length, L) which can be equal to, smaller or larger than the exit diameter of the tube. The moving piston drives a slug of fluid, causing vorticity to be generated in the boundary layer due to the no-slip condition. As the high-speed slug of fluid emerges from the nozzle, this boundary layer forms a cylindrical vortex sheet that rolls up into a spiral form, thus forming a vortex ring (Fig 1B–1D). In the formative stages, the vortex ring entrains the surrounding fluid as it propagates. This generates a vortex bubble (Dvb), the diameter of which is larger than the ring (Dr) (Fig 1E). Even after the formation process, entrainment of surrounding fluid can occur, followed by subsequent detrainment, the balance between which prevents a substantial change in its diameter.

Fig 1. Geometric details of vortex ring and ring generator.

Fig 1

(A) Piston-nozzle arrangement for generating vortex ring. Dashed line shows the outline of piston after it has moved by ΔX. Grey shaded region is the length of fluid that emerges from the nozzle, called here as slug length (L). (B) Variation of piston velocity/slug velocity with time. D and Up are diameter and velocity of piston, respectively, D0 and Us are exit diameter and velocity at the exit of the nozzle, respectively. Tp is the total time for which piston moves. (C) Side view, (D) isometric view and (E) line diagram of side view of the ring. Black circle denotes the core of the vortex ring, in which the vorticity is concentrated. Dr, Dvb and e denote diameter of the ring (distance between the cores), diameter of the vortex bubble including entrained air, and eccentricity of the ellipsoid, respectively.

The momentum (more precisely, impulse) of the ring is determined by the momentum imparted to the surrounding fluid in the tube by the piston. It depends on the type of motion imposed on the piston, fluid viscosity, piston travel time, its radius, and circulation. During vortex formation, the ring accelerates and then may rise to a constant velocity or slow down. The slowing down may be attributed to the entrainment of surrounding fluid, entrainment followed by detrainment, viscous diffusion of the core or vortex instabilities [25]. The turbulent vortex ring, as compared to the laminar one, is characterized by a rapid growth rate of its diameter and shedding of vorticity to the wake, which causes a rapid decrease in its propagation velocity. Thus, piston movement time (stroke time, Tp) and slug length (L) can be designed to generate the flow properties of the ring (Fig 1B). General working relations based on simple conservation laws to achieve this objective are provided in the Appendix (see S1 Appendix).

Our experimental set-up consisted of a 60 cm long, 30cm square cross-section dismountable clear Perspex chamber, a 40 cm long, 3.7 cm internal diameter (Do) cylindrical PVC nozzle (2mm thick), a 12-inch 100W and 8Ω speaker, a digital-to-analog converter NI-DAQ, a high voltage high current direct coupled (DC) amplifier and a high-speed camera (Miro EX4, Vision Research, Ametek) (Fig 2). The large Perspex chamber served as a closed test section where we generated vortex ring. The larger dimensions of the test chamber compared to the diameter of the vortex ring reduced any effects of ambient air currents on experimental observations, and hence aided in maintaining a still ambient fluid [26]. The experiments were carried out in a closed room with controlled humidity at an ambient temperature of 20°C.

Fig 2. Vortex ring generator system.

Fig 2

The input signal to the DAQ (top) is a trapezoidal wave with voltage amplitude V, rising time t1, constant duration t2-t1 and fall time t3-t2-t1. This signal is converted into analog form, amplified and fed to the speaker for vortex ring generation. Fog particles were injected into nozzle through fog injection port (FIP), and a high speed camera was placed horizontal to record the lateral view of the vortex ring as it propagates. In experiments with flies, we used two high-speed cameras to capture the 3D trajectories for the flies. All dimensions are in cm.

Instead of a piston-cylinder arrangement, we used a speaker to generate the vortex ring. The speaker was enclosed in a 40cm x 40cm x 5cm wooden chamber (driving section) on the diaphragm side, and each side of the chamber was glued with FevicolTM (Pidilite, Mumbai, India) and pin hammered to ensure that it was airtight. A 5cm diameter hole was cut at the centre of the 40cm x 40cm face of the wooden chamber to facilitate attachment of the nozzle via a PVC flange. A rubber gasket placed between the flange and the wooden chamber eliminated any air leakage. The nozzle was sharp chamfered by an angle of 9° at the exit and smooth chamfered at entry. The speaker attached to the nozzle was then fitted to the test chamber through a 4cm hole cut on its longest side (Fig 2).

Input signal

We first synthesized a trapezoidal signal (Fig 1B) using NI-LabVIEW, which consisted of three parts: acceleration (100 μs), constant velocity (30 ms), and deceleration (100 ms), such that it resulted in the desired velocity of the vortex ring. A large deceleration time eliminates the formation of stopping vortex, which results from the abrupt stopping of the piston, due to separation and rolling of the secondary boundary layer induced by the primary vortex ring on the outer surface of the tube [26]. When generating a gust, the formation of stopping vortex must be eliminated, because it alters the strength and size of the vortex ring [26], and also because it may induce a secondary response in insects which are hit first by the vortex ring and next by the stopping vortex. We converted the signal into analog form for physical output using NI–c9263, amplified it using an in-house DC power amplifier, and fed it to the speaker, resulting in the formation of a vortex ring at the exit of the nozzle.

Formation of vortex ring

The signal, when fed to the speaker, displaces the speaker diaphragm which imparts its momentum to the surrounding air, causing an equivalent volume of air being pushed out from the chamber into the nozzle. Unlike conventional piston-cylinder configuration for vortex ring generation [26], in our experimental set-up, we have used a long nozzle (11 times its diameter) so that the exit is far away from the speaker diaphragm. This eliminates the generation of a piston vortex and any disturbances similar to those generated using orifice [26].

Characterization of vortex ring

The actual response of the insects depends on their radial position at the time they intercept the vortex ring (see ref. [27] for instantaneous streamlines). Although the vortex ring has an average self-induced translational velocity, the fluid associated with it has local axial and radial velocities, and these velocities depend on the radial location relative to the centre of the ring. Like translational velocity and diameter, the local velocities are well-documented and repeatable [23]. Here, we measured the gross flow properties of the vortex rings using two techniques: fog visualization and styrofoam bead method. Both methods were carried out separately, and flow properties were characterized using each method (see S1 Video).

Fog visualization

We used fog particles to seed the flow (Antari Fogger, Taiwan) for visualization of the vortex ring. The average particle size was on the order of 1–2 μm. The fog was first filled into a 500ml wash bottle and injected through a fog injection port (FIP) on the nozzle (Fig 2). The port was made on the upper circumference of the nozzle, at 18 cm (4.5 D0) away from its exit plane. The circumferential (lower, upper, or sidewise) position of the port did not affect the visualization. Its longitudinal position, however, determined whether any fog particles were present at the exit of the nozzle before ring formation (i.e. background fog at nozzle exit). Keeping FIP at this distance ensures there is no leakage of fog in the test chamber before the initiation of the diaphragm motion. A 5cm square window was hinged on the longer side opposite to nozzle. We closed the window during visualization to eliminate any effects of external air current on the ring propagation and its trajectory. After the recording, we opened the window to remove the residual fog inside the chamber before starting the next trial.

The vortex ring can be generated under varying Reynolds numbers (= UavgD0/ν) (Fig 3). For example, we generated vortex rings at Reynolds numbers of 4.7×103 (Fig 3B) and 1.6×104 (Fig 3C). The vortex sheet emanates from the edge of the nozzle on triggering the speaker, rolls up to form a spiral and hence, a ring. This rolled up vortex propagates while drawing more fluid from the ambient surrounding until it attains a fully formed size. We did not observe any secondary and piston vortices (also [26]), which makes it well-suited for generating discrete gusts. Although the flow within the core of the ring produced here is laminar, it is also possible to produce turbulent rings by increasing their speed, diameter, or both [27, 28].

Fig 3. Flow visualization and characterization of vortex ring.

Fig 3

(A) White and red circles denote leading edge (LE) and extreme end of the ring, respectively. These were tracked to calculate its axial position and diameter respectively. (B) Flow visualization at different time instances for Uavg = 1.9 m/s. The ring propagates from left to right. Tn = 0 indicates the time instance when ring just starts forming. (C) Effect of gust (Uavg = 6.4 m/s) on a freely hanging Styrofoam bead. Position of bead (i) when there is no gust, (ii) just before the gust, (iii) during gust, and (iv) after gust. The bead moves with the gust (iii) until the thread is taut. Black circle in (iii) shows the position of the bead when it is at the centre of the vortex ring.

Estimation of flow speed using styrofoam bead method

In addition to fog, we used a styrofoam bead to measure flow speeds. Because the density of styrofoam beads (6.52 kg/m3) is of the same order of magnitude as the density of air (for details, see [29]), the bead is expected to attain the same velocity as the gust that it intercepts. This method allowed us to make point measurements of the velocity field created by the gust. We suspended the styrofoam bead using thin sewing thread from the test chamber ceiling, such that it rested on the centreline of the nozzle exit and intercepted the vortex ring. The bead was placed at different axial locations along the centreline of the nozzle exit to measure the velocity of the bead, and hence the gust at various axial locations (Fig 3C).

Using a 12-bit CMOS camera (Phantom Miro EX4, Vision Research, Ametek, New Jersey, USA) fitted with an 18–70 mm focal length lens (Nikon, Tokyo, Japan), we recorded the flow images for both these methods at 1200 fps and 50 μs exposure time. Because of the low exposure time, we additionally illuminated the background using two 1000W halogen lamps. The camera was placed to record a lateral view of vortex ring propagation in a plane perpendicular and vertical to the nozzle exit plane. The external diameter of the nozzle served as a calibration scale for the images.

Based on exit diameter of the nozzle (Do) and the ring average velocity (Uavg), we define non-dimensional time Tn = Uavgt/D0 where t is the measured time. Similarly, the axial distance Xn from the nozzle exit is non-dimensionalized with the exit diameter D0 and given by Xn = X/D0. The dimensionless diameter of the ring is given by Dn = Dvb/D0, where Dvb is the instantaneous diameter of vortex bubble (i.e., diameter of the ring with entrained air; Fig 1), and dimensionless velocity of the ring is given by Un = Uvb/Uavg where Uvb is instantaneous velocity of the vortex bubble.

Experiments with freely-flying soldier flies

We tested this apparatus on soldier flies Hermetia illucens obtained from a culture housed in the National Centre for Biological Sciences campus at Bangalore. Because solider flies are strongly attracted to light, it is possible to exploit this behavior to control the direction of their flight, which was crucial in our experiment. We released a group of 5–6 flies together into a test chamber to increase the probability that at least one of them encountered the gust. We recorded their flight motion at 4000 fps using two synchronized, high-speed cameras (Phantom VEO 640L and Phantom V611, Vision Research) for more than 80 trials, 14 of which were calibrated and digitized using MATLAB-based routines-easywand5 and DLTdv7 [30], respectively to measure their body and wing kinematics in response to the gust. The image data were down-sampled to 1000 fps, and then digitized. We tracked the head, abdomen, each wing base, and tip to get their 3D position in the global reference frame. The Centre of Mass (CoM) was assumed to lie at one-third body length from the abdomen [31] and used to represent the flight trajectory. X is the direction along the nozzle centreline, and Y and Z represent lateral and vertical axes, respectively. Here, the origin (X,Y,Z) = (0,0,0) is placed at the centre of the nozzle exit, and X is positive in forward direction of the ring while Y- and Z axes conform to the conventions of the right-handed coordinate system.

The total velocity (velocity magnitude) of the fly is the resultant of three velocity components. A fourth-order low-pass Butterworth filter with cut-off frequency 200Hz was applied to the corresponding CoM data to minimize digitization error. We next calculated the velocity along each axis using a second-order central difference scheme. We non-dimensionalized velocity by dividing it by the product of the body length of flies and their wingbeat frequency. Similarly, we calculated the body roll angle ) relative to the horizontal plane as the elevation angle of the vector joining the wing base and the CoM. Counter-clockwise rotations with respect to the axial direction of forward flight were treated as positive.

Results and discussion

Characterization of gust pertubations

The discrete gust is characterized by the spatial and temporal evolution of vortex ring and its translational velocity. We observed that the ring velocity varies linearly from 0.4 m/s to 6.4 m/s for input voltages (Vin) ranging from 2.3V to 23V (Uavg = 0.28×Vin, R2 = 0.99) (Fig 4A), with Re ranging from 1×103 to 1.6×104 respectively. Such strong dependence of the translational velocity of the ring on voltage shows that by modulating the amplitude of input signal, vortex rings of different strengths can be generated, thus allowing control over flow properties. The bead velocity and the average velocity of gust measured using flow visualization matched each voltage input, thus providing cross-validation of these methods. Hence, the bead method, which is easy to establish, can be used to measure air flow. The velocity changes in a similar fashion with time and space (distance along axis) for each voltage value.

Fig 4. Flow characteristics of vortex ring.

Fig 4

(A) Average propagation velocity of the ring as a function of input voltage to the speaker. Velocity of the ring obtained using fog visualization (circles) and bead method (triangles) are in good agreement for different input voltages. Dashed line is Uavg = 0.2745 Vin, R2 = 0.99. (B-D) Non-dimensional flow properties of vortex ring with Uavg = 6.4 m/s and vortex bubble diameter 8.6 cm measured using flow visualization and bead method. (B) Xn is the axial distance from the nozzle exit, non-dimensionalized reative to the exit diameter D0 and given by Xn = X/D0. Xn = 0 indicates the centre of nozzle exit. (C) Dn = Dvb/D0, is the dimensionless diameter of the ring, where Dvb is the instantaneous diameter of vortex bubble. (D) Un = Uvb/Uavg is the dimensionless velocity of the ring, where Uvb is instantaneous velocity of the vortex bubble. Values are mean ± SD.

As a special case, we discuss the ring flow properties for Re = 1.6×104 (Uavg = 6.4 m/s) (Fig 4B–4D). The ring propagates as a quadratic function of time until Tn = 4, after which it moves linearly until it reaches near the opposite wall of the test chamber (Fig 4B). It slows down, increases its diameter, and begins to deform as it approaches the opposite wall. The diameter of the ring also grows as a quadratic function of space from 1.25D0 to 2.25D0 up to Xn = 3 (Fig 4C), beyond which it attains space invariant final size of 2.3±0.03 D0. Because the core of the ring was not visible in all images and across all videos, we tracked the point LE to measure its axial position, and lateral extremes of vortex ring to measure its diameter (Fig 3A). The diameter measured here is, thus, the diameter of the vortex bubble and not the ring. Entrained fluid generally constitutes about 20–40% of the total volume of fluid carried by tube generated vortex ring [27]. In the present study, it was experimentally difficult to measure the entrained mass fraction. Instead, we estimated it theoretically based on the eccentricity (e) of the ring. For the present study, e = 0.62 and entrained mass fraction, k = 0.57. Similar values of eccentricity are observed in different studies [22, 27, 32]. k = 0.57 yields Dr = 0.86Dvb.

The non-dimensional ring velocity, calculated here by applying second-order central difference scheme to axial position, becomes uniform after Xn = 3 from the nozzle exit (Fig 4D). The average speed of the ring for Re = 1.6×104 was 6.4 m/s. The average velocity of the ring corresponds to the velocity averaged over Xn≥3 after which it is nearly constant, consistent with previous observations [33]. The maximum standard deviation in ring properties measured for three trials was less than 10% and the average standard deviation was less than 5% of their mean values, implying high repeatability of the measured values. Because the ring attains nearly constant size and constant velocity after a particular time and distance from the nozzle and remains so for a long time and axial distance (Fig 4D), it provides a longer spatial and temporal window to study the gust response of the insects. This method can be used to study perturbation responses in both free-flying insects and hovering insects.

When using a vortex ring as a gust, ideally the gust size should match the characteristic size of the study animal. For instance, in birds, bats and insects, the ring diameter should exceed their wingspan for a head-on gust. For lateral gust perturbations in fish, it should exceed their body length. This allows their flight and swimming to be contained within the ring under controlled laboratory conditions. In our experiments, the ring diameter Dvb (8.5 cm) was about 4 times the fly wingspan (2.2 cm). Because animal trajectories are not under the control of the experimenter, there is some spatial and temporal uncertainty about where the animal intercepts the vortex ring. For instance, in one of case studies depicted in Fig 5, the gusts due to the vortex ring hit only the right wing of one of the flies (shown in black). Furthermore, the translational velocity of the ring should match the order of forward velocity of the subject under consideration to induce an optimal response. Although this gust velocity may hold for birds, bats and fish, in case of insects that flap their wings at higher rates, the ring velocity may need to be of the order of the wing tip velocity. In the present experiments, the ring velocity was chosen to be 6.4 m/s, comparable to the wing-tip velocity of the fly (~4.85 m/s).

Fig 5. Changes in body kinematics of soldier flies (Hermetia illucens) due to the gust induced by the vortex ring in 4 different trials.

Fig 5

(A) Trajectory in Y-Z plane normalized by the average body length (Lavg) of flies. The mean axial distance where the gust hits the insects is X0 = 3.7±0.38D0. Black circle indicates the front view of the vortex ring, and the intersection of vertical and horizontal dashed lines is the centre of the vortex ring. Coloured lines are the trajectories of flies for each trial represented by 1–4, and open circles on each curve denote the position of the flies just before they were hit by the ring. (B) Normalized speed versus non-dimensional time (in wing beats). Forward speed is normalized with the average speed of the flies before being hit by the ring, and time is non-dimensionalized by multiplying with the wing beat frequency. T = 0 indicates the time instance just before flies were just hit by the ring. Time period of gust is indicated in vertical grey strip. (C) Oblique top and the corresponding side views of flight sequences for trial 4 at different time instances showing distinct change in body roll angle. Number on the top row indicates the time instance of fly with respect to gust in terms of wing beats. (D) Change in body roll angle plotted against the wingbeat. Filled circles on black curve (trial 4) denote the time instances of the fly in (C).

Another important consideration when employing vortex ring as a gust is that its formation number should be ideally less than 4 [34]. This ensures that there is no trailing jet behind the ring, and gust is contained within the ring so as not to elicit secondary response in fliers. In all the experiments reported here, the formation number was ≈3.2 (see calculation in S1 Appendix).

Vortex ring based gust perturbation in flying insects

To demonstrate the effectiveness of the vortex ring as a precise gust-imparting device, we subjected freely-flying soldier flies Hermetia illucens with head-on gusts, and measured the influence of these gusts on their flight trajectory, velocity and body angles. As mentioned above, we selected vortex ring velocity of 6.4 m/s (Re = 1.6×104) to perturb the flies. We have chosen results from four experiments to illustrate the types of responses that were obtained.

In these experiments, the flies were perturbed by the gust at mean axial location of X0 = 3.7±0.38 D0 from the exit of the nozzle, and were always contained inside the gust at encounter (Fig 5A, and S2 Video). If the flies flew to the left side of the gust, they continued to fly on the same side even after being hit by the gust. Similarly, they flew downward after the encounter with gust, indicating a possible loss in lift forces. The flies did not recover their initial vertical and lateral position after gust in any trial. Even in trials for which the gust was not visible, the time instant of the gust encounter could be precisely determined by the large displacement of the antennae (S2 Video).

Before being hit by the ring, the forward speed of the flies was near-constant in each case. However, the gust decreased their forward speed by as much as ~70% maximum and ~30% on average (Fig 5B). The deceleration of flies due to gust in the present study is consistent with observation in bumble bees [15, 18], indicating gust as a limiting parameter on the forward speed of insects [4]. On the other hand, recently, researchers have found a increase in airspeed of pigeon when flying in turbulence [35], and golden eagles exploiting gust to accelerate [36], suggesting that birds might enhance their airspeed by harnessing energy from the surroundings.

The flies had near-zero body roll angle (<15°) before intercepting the vortex ring, but changed by as much as (~160°) due to gust (Fig 5C, and S2 Video). Insect such as honeybees [18], bumblebees [8, 15] and orchid bees [4] have also been observed to respond highly to gusts along roll axis. High roll rates of body and tail are also observed in ruby-throated humming birds when flying in turbulent wind conditions rather than undisturbed air [14]. Some insects extend their legs tin response to the perturbation due to gusts [4, 18], whereas ruby-throated hummingbirds compensate for turbulence by modulating their wing kinematics including stroke angle, stroke and asymmetry [14]. In the present study, soldier flies extended their legs as well as changed their wing stroke angles during recovery from the gusts. The observed changes in the flight trajectory, speed and body orientation show that the vortex ring can be used as a precise gust to perturb the trajectories of flies.

Although this method provides a well-characterized and precise gust, the actual response of the insects depends on their radial position at the time they are hit by the vortex ring. These conditions can be well-controlled in robotic flappers or drones, but there is no easy way to ensure that freely-flying insects will have a repeatable position and wing configuration at the time of impact. Because the axial velocity within the ring is non-uniform, it is difficult to elicit repeatability in their response of the insects. As the gust dynamics are predictable, a post-hoc reconstruction of the impact of gust is possible provided the fly does not intercept gust towards the rim of the vortex ring. Thus, this method greatly enhances our ability to determine the magnitude of the gust encounter and interpret the response of the fly relative to this gust. Overall, this method can lend itself to studies on the biomechanics and control and stability of diverse animals and robotic fliers.

Conclusion

We present a method of gust generation via a discrete vortex ring. The ring was generated by an impulsive motion of a diaphragm of an electronic speaker. The flow physics of this perturbation method is well-understood and hence, the flow properties are highly controllable. Besides, this method allows high repeatability and reproducibility, can be implemented at a low-cost, and has a simple mechanical design. As an example, we tested its application to study the impact of gust on insect flight, but as such this method can be used in a variety of situations and habitats. The application of the vortex ring as a gust is not only limited to insects but could potentially be extended to study birds, bats, and micro-aerial vehicles (MAVs) in air, and fish and underwater autonomous vehicles in water. We have provided relevant theoretical relations that will be useful for design of a gust generator for a specific application.

Supporting information

S1 Video. Gust generation and characterization.

(MP4)

Download video file (3.8MB, mp4)
S2 Video. Insect and gust interaction.

(MP4)

Download video file (6.8MB, mp4)
S1 Appendix

(DOCX)

pone.0305084.s003.docx (38.1KB, docx)

Acknowledgments

We thank Dr Toshiyuki Nakata, Chiba University for his helpful feedback on this manuscript.

Data Availability

The data are present at this link: https://drive.ncbs.res.in/index.php/s/Bk7TF2EEjr46YJj.

Funding Statement

Funding for this study was provided by grants from the Air Force Office of Scientific Research (AFOSR) # FA2386-11-1-4057 and # FA9550-16-1-0155, and National Centre for Biological Sciences(Tata Institute of Fundamental Research) to SPS. We also acknowledge the support of the Ministry of Earth Sciences, Government of India, under project no. MESO-0034 and the Department of Atomic Energy, Government of India, under project no. 12-R&D-TFR-5.04-0800.

References

  • 1.Chapman JW, Nilsson C, Lim KS, Bäckman J, Reynolds DR, Alerstam T. Adaptive strategies in nocturnally migrating insects and songbirds: contrasting responses to wind. Journal of Animal Ecology. 2016;85(1):115–24. doi: 10.1111/1365-2656.12420 [DOI] [PubMed] [Google Scholar]
  • 2.Reynolds K V, Thomas ALR, Taylor GK. Wing tucks are a response to atmospheric turbulence in the soaring flight of the steppe eagle Aquila nipalensis. J R Soc Interface. 2014;11(101):20140645. doi: 10.1098/rsif.2014.0645 [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 3.Laurent KM, Fogg B, Ginsburg T, Halverson C, Lanzone MJ, Miller TA, et al. Turbulence explains the accelerations of an eagle in natural flight. Proceedings of the National Academy of Sciences. 2021;118(23):e2102588118. doi: 10.1073/pnas.2102588118 [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 4.Combes SA, Dudley R. Turbulence-driven instabilities limit insect flight performance. Proceedings of the National Academy of Sciences. 2009;106(22):9105–8. doi: 10.1073/pnas.0902186106 [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 5.Ortega-Jimenez VM, Greeter JSM, Mittal R, Hedrick TL. Hawkmoth flight stability in turbulent vortex streets. Journal of Experimental Biology. 2013;216(24):4567–79. doi: 10.1242/jeb.089672 [DOI] [PubMed] [Google Scholar]
  • 6.Ravi S, Garcia JE, Wang C, Dyer AG. The answer is blowing in the wind: free-flying honeybees can integrate visual and mechano-sensory inputs for making complex foraging decisions. Journal of Experimental Biology. 2016;219(21):3465–72. doi: 10.1242/jeb.142679 [DOI] [PubMed] [Google Scholar]
  • 7.Crall JD, Chang JJ, Oppenheimer RL, Combes SA. Foraging in an unsteady world: bumblebee flight performance in field-realistic turbulence. Interface Focus. 2017;7(1):20160086. doi: 10.1098/rsfs.2016.0086 [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 8.Engels T, Kolomenskiy D, Schneider K, Lehmann FO, Sesterhenn J. Bumblebee flight in heavy turbulence. Phys Rev Lett. 2016;116(2):28103. doi: 10.1103/PhysRevLett.116.028103 [DOI] [PubMed] [Google Scholar]
  • 9.Ravi S, Noda R, Gagliardi S, Kolomenskiy D, Combes S, Liu H, et al. Modulation of flight muscle recruitment and wing rotation enables hummingbirds to mitigate aerial roll perturbations. Current Biology. 2020;30(2):187–95. doi: 10.1016/j.cub.2019.11.025 [DOI] [PubMed] [Google Scholar]
  • 10.Boerma DB, Breuer KS, Treskatis TL, Swartz SM. Wings as inertial appendages: how bats recover from aerial stumbles. Journal of Experimental Biology. 2019;222(20):jeb204255. doi: 10.1242/jeb.204255 [DOI] [PubMed] [Google Scholar]
  • 11.Chapman JW, Drake VA, Reynolds DR. Recent insights from radar studies of insect flight. Annu Rev Entomol. 2011;56:337–56. doi: 10.1146/annurev-ento-120709-144820 [DOI] [PubMed] [Google Scholar]
  • 12.Natesan D, Saxena N, Ekeberg Ö, Sane SP. Tuneable reflexes control antennal positioning in flying hawkmoths. Nat Commun. 2019;10(1):5593. doi: 10.1038/s41467-019-13595-3 [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 13.Jafferis NT, Helbling EF, Karpelson M, Wood RJ. Untethered flight of an insect-sized flapping-wing microscale aerial vehicle. Nature. 2019;570(7762):491–5. doi: 10.1038/s41586-019-1322-0 [DOI] [PubMed] [Google Scholar]
  • 14.Ravi S, Crall JD, McNeilly L, Gagliardi SF, Biewener AA, Combes SA. Hummingbird flight stability and control in freestream turbulent winds. J Exp Biol. 2015;218(9):1444–52. doi: 10.1242/jeb.114553 [DOI] [PubMed] [Google Scholar]
  • 15.Ravi S, Crall JD, Fisher A, Combes SA. Rolling with the flow: bumblebees flying in unsteady wakes. Journal of Experimental Biology. 2013;216(22):4299–309. doi: 10.1242/jeb.090845 [DOI] [PubMed] [Google Scholar]
  • 16.Liao JC, Beal DN, Lauder G V, Triantafyllou MS. Fish exploiting vortices decrease muscle activity. Science (1979). 2003;302(5650):1566–9. doi: 10.1126/science.1088295 [DOI] [PubMed] [Google Scholar]
  • 17.Vance JT, Faruque I, Humbert JS. Kinematic strategies for mitigating gust perturbations in insects. Bioinspir Biomim. 2013;8(1):16004. doi: 10.1088/1748-3182/8/1/016004 [DOI] [PubMed] [Google Scholar]
  • 18.Jakobi T, Kolomenskiy D, Ikeda T, Watkins S, Fisher A, Liu H, et al. Bees with attitude: the effects of directed gusts on flight trajectories. Biol Open. 2018;7(10):bio034074. doi: 10.1242/bio.034074 [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 19.Hufstedler EAL, McKeon BJ. Vortical gusts: experimental generation and interaction with wing. AIAA Journal. 2019;57(3):921–31. [Google Scholar]
  • 20.Gupta D, Arakeri JH. Gust enhancement using jet at the trailing edge of a pair of pitching foils. Sadhana. 2021;46(3):140. [Google Scholar]
  • 21.Roadman J, Mohseni K. Gust characterization and generation for wind tunnel testing of micro aerial vehicles. In: 47th AIAA aerospace sciences meeting including the new horizons forum and aerospace exposition. 2009. p. 1290.
  • 22.Didden N. On the formation of vortex rings: rolling-up and production of circulation. Zeitschrift für angewandte Mathematik und Physik ZAMP. 1979;30:101–16. [Google Scholar]
  • 23.Lim TT, Nickels TB. Vortex rings. In: Fluid vortices. Springer; 1995. p. 95–153. [Google Scholar]
  • 24.Zhang C, Hedrick TL, Mittal R. An integrated study of the aeromechanics of hovering flight in perturbed flows. AIAA Journal. 2019;57(9):3753–64. [Google Scholar]
  • 25.Saffman PG. The velocity of viscous vortex rings. Studies in Applied Mathematics. 1970;49(4):371–80. [Google Scholar]
  • 26.Das D, Bansal M, Manghnani A. Generation and characteristics of vortex rings free of piston vortex and stopping vortex effects. J Fluid Mech. 2017;811:138–67. [Google Scholar]
  • 27.Dabiri JO, Gharib M. Fluid entrainment by isolated vortex rings. J Fluid Mech. 2004;511:311–31. [Google Scholar]
  • 28.Sallet DW, RS W. An Experimental Investigation of laminar and Turbulent Vortex Rings in Air. 1974; [Google Scholar]
  • 29.Gupta D. Gust Rejection in Insect Flight [MS Dissertation]. Indian Institute of Science, Bangalore, India; 2020.
  • 30.Hedrick TL. Software techniques for two-and three-dimensional kinematic measurements of biological and biomimetic systems. Bioinspir Biomim. 2008;3(3):34001. doi: 10.1088/1748-3182/3/3/034001 [DOI] [PubMed] [Google Scholar]
  • 31.Ellington C.P. The aerodynamics of hovering insect flight. II. Morphological parameters. Philosophical Transactions of the Royal Society of London B, Biological Sciences. 1984;305(1122). [Google Scholar]
  • 32.Sullivan IS, Niemela JJ, Hershberger RE, Bolster D, Donnelly RJ. Dynamics of thin vortex rings. J Fluid Mech. 2008;609:319–47. [Google Scholar]
  • 33.Hernández RH, Cibert B, Béchet C. Experiments with vortex rings in air. Europhys Lett. 2006;75(5):743. [Google Scholar]
  • 34.Gharib M, Rambod E, Shariff K. A universal time scale for vortex ring formation. J Fluid Mech. 1998;360:121–40. [Google Scholar]
  • 35.Lempidakis E, Ross AN, Quetting M, Krishnan K, Garde B, Wikelski M, et al. Turbulence causes kinematic and behavioural adjustments in a flapping flier. Journal of the Royal Society Interface. 2024;21(212):20230591. doi: 10.1098/rsif.2023.0591 [DOI] [PMC free article] [PubMed] [Google Scholar]
  • 36.Gupta D, Fogg B, Halverson C, Lanzone M, Miller T, Bewley G. Soaring in Turbulence intermittency. Bulletin of the American Physical Society. 2022. Nov 21;67. [Google Scholar]

Decision Letter 0

Iman Borazjani

21 Feb 2024

PONE-D-24-02005Generating controlled gust perturbations using vortex ringsPLOS ONE

Dear Dr. Sane,

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**********

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Reviewer #1: The authors present a study of generating gusts using vortex rings and have applied it to study response of insects to such gusts. In general the paper is written well, however, there are a few points to be addressed

- The vortex characterization is missing any discussion on the formation number of the vortex rings generated. This number is crucial to understand what kind of structure is formed. “A universal time scale for vortex ring formation” (Gharib 1998) is a very important reference regarding vortex ring generation. Given the rings are generated with constant time pulsing and different Reynolds numbers are produced, the vortex rings will have different formation numbers. This difference means the structure of the gust will fundamentally change for each Reynolds number, defeating the whole purpose of the desired well-characterized and repeatable gust.

- Furthermore, If the formation number is larger than 4, then not all of the fluid driven by the piston, or in this case speaker, will roll up into the vortex ring, or bubble as it is called in this paper, so any assumptions like equation 7 and the mass fraction relationship in the appendix are potentially flawed. Since there will be a trailing jet which can make the vortex unstable, it might not present a repeatable case of gust generation using vortex rings at such high formation numbers. Have the authors tried lower formation numbers to ensure better repeatability of the vortex ring formation and hence the associated gust?

- Could the authors comment on the efficacy of using a bead to estimate the velocity of the gust, perhaps there are better ways of achieving this end outcome?

- The authors could take this opportunity to discuss some more about the flight dynamics of the insect under such perturbations. Perhaps estimate some aerodynamics on the flapping wing and comment on the aerodynamic loading and associated responses of the insect. Also comparing these responses to other natural flyers such as birds (across several flapping time scales - eagles/owls to hummingbirds) during gust encounters.

Additional specific comments are below

- Lines 67-70: Suggest rewording with Vortex ring as the subject as that is what is important, not the impulsive flow. The sentence reads awkwardly with that as the subject. “Vortex rings are generated from…”

- Lines 94-95: Saying the piston movement generates the layer of vorticity is awkward. The piston drives a slug of fluid, and it is the motion of this slug combining with the no-slip condition to form a boundary layer. This is important especially in some designs when the piston is far upstream of the nozzle.

- Line 96: As this is a general description of a vortex generation, I would say fluid instead of air.

- Lines 134-135: Many vortex ring studies are not concerned with the stopping vortex. If it is important to prevent from forming in this application, then provide explanation as to why it is important.

- Line 144: missing “a” before “long nozzle”

- Lines 145-146: missing “a” or “the” before piston vortex

- Line 146: I believe the antecedent of “that” is the plural “disturbances,” so replace with “to those generated using an orifice.”

- Line 149: Why have the authors chosen to put “e.g.” in their citation just this one time? Is it more applicable than the other citations?

- Line 170: What is meant by “at different points in time?” Either delete or provide more detail.

- Line 171: The faster vortex ring is referenced to Figure 3B, and the slower vortex ring is referenced to Figure 3C, but the caption is backwards. Please make the caption and reference agree.

- Lines 131-177

- Lines 410-411 The wording defining =0 as the “time instance when no ring is formed” is awkward. Defining that time as the last frame where there is no flow or the instant the vortex generation process begins is clearer.

Reviewer #2: This work introduces a system to generate controllable gusts relying on vortex rings for studying animal locomotion. The authors provided detailed design and characterization of the gust generator, and its application for a case study on free-flying soldier flies. In general, I find the paper is very interesting, very well written, and easy to follow. It also provide a useful and easy-to-build method for studying insect and insect-inspired flight in gusty conditions. I have several minor comments for the authors:

1) It is still unclear what is the limitations of existing methods of gust generation in the Introduction (lines 59-64)

2) I recommend to add a sub-title for the result on gust characterization, for example “Generation of gust perturbations”

3) Line 262: “When using a vortex ring as a gust, it should fully encompass the subject.” Why is this a condition? What happen if it cannot cover the subject? For example, the flight trial 4 of the fly in Movie 2 may represent this case as the gust hit the right wing only (or impact stronger on the right wing), causing rolling response of the fly.

**********

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Reviewer #2: No

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PLoS One. 2024 Jul 8;19(7):e0305084. doi: 10.1371/journal.pone.0305084.r002

Author response to Decision Letter 0


15 Apr 2024

General comments:

We thank both reviewers for their engagement with the manuscript and for providing detailed and helpful comments, which we have tried to address below. The suggested changes add much value and rigor to the manuscript, and also help streamline it a little better. In the responses of reviewers below, the referee’s comments are in bold, and our responses are in regular font. The changes made in the manuscript have been highlighted in yellow.

Response to Reviewer #1:

-The vortex characterization is missing any discussion on the formation number of the vortex rings generated. This number is crucial to understand what kind of structure is formed. “A universal time scale for vortex ring formation” (Gharib 1998) is a very important reference regarding vortex ring generation. Given the rings are generated with constant time pulsing and different Reynolds numbers are produced, the vortex rings will have different formation numbers. This difference means the structure of the gust will fundamentally change for each Reynolds number, defeating the whole purpose of the desired well-characterized and repeatable gust.

We agree that the paper would benefit by the inclusion of some discussion of the formation number, and have therefore added text to clarify the role of formation number in our method. We have also included the citation of Gharib (1998) in this context. Because we used a speaker to generate vortex rings, the calculation of a formation number was somewhat trickier, but we were able to do so using measured characteristics of the vortex rings. We elaborate on this point below:

For small formation numbers (<4), the vortex ring is formed with all the vorticity and mass being entrained from the slug of fluid exiting the nozzle, and the ring achieves its maximum size. As formation number increases, the size and strength of the ring do not increase. The additional fluid mass and vorticity is contained in the trailing jet, giving rise to “pinched-off” state, thus altering the structure of the flow.

In the present study, the experiments with soldier flies were conducted at the same Reynolds number (Re=(U_avg D_0)/ν=1.6 × 〖10〗^4), and hence also same formation number. The speed or diameter of the vortex rings was left unaltered. Because the experiments were carried out in a closed room at ambient temperature of 〖20〗^∘ C, i.e. at constant kinematic viscosity, Re based on the speed and size of the ring and the kinematic viscosity of air remained constant in all the experiments with soldier flies.

During characterization of the rings, we generated vortex rings with various speeds, and hence, different Re (see fig 4A). Fig. 3B is one such case showing the formation of the ring at Re=4.7×〖10〗^3 but did not use the low Re vortex ring to probe the response of soldier flies.

All the experiments with soldier flies were conducted at formation number ≈ 3.2 (as estimated below) which was the maximum formation number for the vortex rings in our experiments. This is consistent with the fact that, qualitatively, we did not observe a trailing edge pinch-off in the flow visualization of the ring (See SI video1). Moreover, a secondary response would be expected from the flies if they were hit first by the ring and then again by a trailing jet formed at formation numbers > 4, but we never observed such a response. These observations strongly suggest that our formation number does not exceed 4.

We estimated the formation numbers in our setup in the following way:

Estimation of formation number:

In the experiments reported here, Dr = 0.86Dvb=7.3 cm, e =0.62, and D0 =3.7 cm. Inserting these values into equation 4,

L=(2D_r^3 e)/(3D_o^2 )=11.75 cm

And the formation number, L/D_(0 )=3.17

This estimate is based on the diameters of the ring and the nozzle exit, which are measured quantities in this study, and additionally assumes volume conservation (i.e., all the fluid ejected from the nozzle gets rolled into the vortex ring) based on flow visualization and SI Video1.

- Furthermore, If the formation number is larger than 4, then not all of the fluid driven by the piston, or in this case speaker, will roll up into the vortex ring, or bubble as it is called in this paper, so any assumptions like equation 7 and the mass fraction relationship in the appendix are potentially flawed. Since there will be a trailing jet which can make the vortex unstable, it might not present a repeatable case of gust generation using vortex rings at such high formation numbers. Have the authors tried lower formation numbers to ensure better repeatability of the vortex ring formation and hence the associated gust?

We agree that equation 7 will not hold if the formation number exceeds 4, and have added this cautionary statement in line 381 -382. In our case, the vortex rings were generated at lower formation numbers (see Fig. 4A), and each case was repeatable. Fig. 4 B-D shows mean values ± standard deviation (which is less than 5% the mean values) for the vortex ring used in the present study, and serves a good check for repeatability of the vortex ring and the associated gust.

- Could the authors comment on the efficacy of using a bead to estimate the velocity of the gust, perhaps there are better ways of achieving this end outcome?

This study uses two methods to estimate the gust velocity. First, we use a Styrofoam bead because its density is of the same order as that of air, i.e., it is close to neutrally buoyant. This means that it instantaneously responds to the flow, and can be safely used as a tracer to measure the average gust velocity. Second, we used the fog visualization method to corroborate the bead measurement. There is an excellent agreement between the velocity estimate obtained using the bead and that calculated using fog visualization method for different speed of gusts (Fig 4A, D). An added advantage of using bead is that it is economical and readily available in the local market, which allowed us to get a rough estimate of the gust propagation speed quickly, especially we hope that this method will be readily accessible.

- The authors could take this opportunity to discuss some more about the flight dynamics of the insect under such perturbations. Perhaps estimate some aerodynamics on the flapping wing and comment on the aerodynamic loading and associated responses of the insect. Also comparing these responses to other natural flyers such as birds (across several flapping time scales - eagles/owls to hummingbirds) during gust encounters.

As suggested by the reviewer, we have added a brief discussion in the main text (line 308 to 321) on comparison of the observed responses in soldier flies to other natural flyers.

Additional specific comments are below

- Lines 67-70: Suggest rewording with Vortex ring as the subject as that is what is important, not the impulsive flow. The sentence reads awkwardly with that as the subject. “Vortex rings are generated from…”

Agreed. We have made the suggested changes.

- Lines 94-95: Saying the piston movement generates the layer of vorticity is awkward. The piston drives a slug of fluid, and it is the motion of this slug combining with the no-slip condition to form a boundary layer. This is important especially in some designs when the piston is far upstream of the nozzle.

Agreed. We have made the suggested changes.

- Line 96: As this is a general description of a vortex generation, I would say fluid instead of air.

Agreed. We have made the suggested changes.

- Lines 134-135: Many vortex ring studies are not concerned with the stopping vortex. If it is important to prevent from forming in this application, then provide explanation as to why it is important.

We have made the suggested changes.

- Line 144: missing “a” before “long nozzle”

We have made the suggested changes.

- Lines 145-146: missing “a” or “the” before piston vortex

We have made the suggested changes.

- Line 146: I believe the antecedent of “that” is the plural “disturbances,” so replace with “to those generated using an orifice.”

We have replaced “that” with “those”, as suggested.

- Line 149: Why have the authors chosen to put “e.g.” in their citation just this one time? Is it more applicable than the other citations?

We have removed ‘e.g.’ in the citation, and added more details. In the previous version, we had added e.g., because this point has been made by several other authors as well.

- Line 170: What is meant by “at different points in time?” Either delete or provide more detail.

We thank the reviewer for pointing out this error, and have deleted this.

- Line 171: The faster vortex ring is referenced to Figure 3B, and the slower vortex ring is referenced to Figure 3C, but the caption is backwards. Please make the caption and reference agree.

Thanks much for pointing out this error. We have corrected it.

- Lines 410-411 The wording defining =0 as the “time instance when no ring is formed” is awkward. Defining that time as the last frame where there is no flow or the instant the vortex generation process begins is clearer.

We have made the changes, as suggested.

Response to Reviewer #2:

1) It is still unclear what is the limitations of existing methods of gust generation in the Introduction (lines 59-64)

One advantage of using vortex ring as a gust is that it is possible to generate a high-speed perturbation while keeping the ring laminar. Laminarity is important to eliminate any ambiguity in the flow characteristics, and consequently, to better understand the response of insects. On the contrary, most of the existing methods are turbulent gusts. For example, jets and grid-generated turbulence are turbulent, and therefore, the response of the insects could be a function of both mean flow and turbulent fluctuations, thereby making it difficult to decouple their effects on the insects’ response. Karman-vortices, on the other hand, are generated by cylinders placed in the upstream, which lead to wake formation, and hence, lack of neat and laminar flow in the region where insect experiences gusts. Vortex ring is, however, free from these effects. We have added these points in the main text (line 73-79).

2) I recommend to add a sub-title for the result on gust characterization, for example “Generation of gust perturbations”

We have added the suggestion sub-title.

3) Line 262: “When using a vortex ring as a gust, it should fully encompass the subject.” Why is this a condition? What happen if it cannot cover the subject? For example, the flight trial 4 of the fly in Movie 2 may represent this case as the gust hit the right wing only (or impact stronger on the right wing), causing rolling response of the fly.

We thank the reviewer for this question, and have, accordingly, made the changes (line 277-280).

Attachment

Submitted filename: Response_to_Reviewers_ver3.docx

pone.0305084.s004.docx (22.8KB, docx)

Decision Letter 1

Iman Borazjani

24 May 2024

Generating controlled gust perturbations using vortex rings

PONE-D-24-02005R1

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Kind regards,

Iman Borazjani, Ph.D.

Academic Editor

PLOS ONE

Additional Editor Comments (optional): Reviewer 1 has a minor comment, which can be addressed when submitting all the finals or during the proofing. 

Reviewers' comments:

Reviewer's Responses to Questions

Comments to the Author

1. If the authors have adequately addressed your comments raised in a previous round of review and you feel that this manuscript is now acceptable for publication, you may indicate that here to bypass the “Comments to the Author” section, enter your conflict of interest statement in the “Confidential to Editor” section, and submit your "Accept" recommendation.

Reviewer #1: (No Response)

Reviewer #2: All comments have been addressed

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2. Is the manuscript technically sound, and do the data support the conclusions?

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Reviewer #1: Yes

Reviewer #2: Yes

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3. Has the statistical analysis been performed appropriately and rigorously?

Reviewer #1: I Don't Know

Reviewer #2: (No Response)

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Reviewer #1: No

Reviewer #2: Yes

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5. Is the manuscript presented in an intelligible fashion and written in standard English?

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Reviewer #1: Yes

Reviewer #2: Yes

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6. Review Comments to the Author

Please use the space provided to explain your answers to the questions above. You may also include additional comments for the author, including concerns about dual publication, research ethics, or publication ethics. (Please upload your review as an attachment if it exceeds 20,000 characters)

Reviewer #1: - Appendix line 384 - "... if the formation number (L/D0) is approximately 4 (34)".. should this be "... (L/D0) is limited to approximately 4 (34)" ?

Reviewer #2: The authors have reflected all my concerns. The reviewer has no further comments and recommends acceptance.

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Reviewer #1: No

Reviewer #2: No

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Acceptance letter

Iman Borazjani

31 May 2024

PONE-D-24-02005R1

PLOS ONE

Dear Dr. Sane,

I'm pleased to inform you that your manuscript has been deemed suitable for publication in PLOS ONE. Congratulations! Your manuscript is now being handed over to our production team.

At this stage, our production department will prepare your paper for publication. This includes ensuring the following:

* All references, tables, and figures are properly cited

* All relevant supporting information is included in the manuscript submission,

* There are no issues that prevent the paper from being properly typeset

If revisions are needed, the production department will contact you directly to resolve them. If no revisions are needed, you will receive an email when the publication date has been set. At this time, we do not offer pre-publication proofs to authors during production of the accepted work. Please keep in mind that we are working through a large volume of accepted articles, so please give us a few weeks to review your paper and let you know the next and final steps.

Lastly, if your institution or institutions have a press office, please let them know about your upcoming paper now to help maximize its impact. If they'll be preparing press materials, please inform our press team within the next 48 hours. Your manuscript will remain under strict press embargo until 2 pm Eastern Time on the date of publication. For more information, please contact onepress@plos.org.

If we can help with anything else, please email us at customercare@plos.org.

Thank you for submitting your work to PLOS ONE and supporting open access.

Kind regards,

PLOS ONE Editorial Office Staff

on behalf of

Dr. Iman Borazjani

Academic Editor

PLOS ONE

Associated Data

    This section collects any data citations, data availability statements, or supplementary materials included in this article.

    Supplementary Materials

    S1 Video. Gust generation and characterization.

    (MP4)

    Download video file (3.8MB, mp4)
    S2 Video. Insect and gust interaction.

    (MP4)

    Download video file (6.8MB, mp4)
    S1 Appendix

    (DOCX)

    pone.0305084.s003.docx (38.1KB, docx)
    Attachment

    Submitted filename: Response_to_Reviewers_ver3.docx

    pone.0305084.s004.docx (22.8KB, docx)

    Data Availability Statement

    The data are present at this link: https://drive.ncbs.res.in/index.php/s/Bk7TF2EEjr46YJj.


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