Downstroke and upstroke conflict during banked turns in butterflies.

For all flyers, aeroplanes or animals, making banked turns involve a rolling motion which, due to higher induced drag on the outer than the inner wing, results in a yawing torque opposite to the turn. This adverse yaw torque can be counteracted using a tail, but how animals that lack tail, e.g. all insects, handle this problem is not fully understood. Here, we quantify the performance of turning take-off flights in butterflies and find that they use force vectoring during banked turns without fully compensating for adverse yaw. This lowers their turning performance, increasing turn radius, since thrust becomes misaligned with the flight path. The separation of function between downstroke (lift production) and upstroke (thrust production) in our butterflies, in combination with a more pronounced adverse yaw during the upstroke increases the misalignment of the thrust. This may be a cost the butterflies pay for the efficient thrust-generating upstroke clap, but also other insects fail to rectify adverse yaw during escape manoeuvres, suggesting a general feature in functionally two-winged insect flight. When lacking tail and left with costly approaches to counteract adverse yaw, costs of flying with adverse yaw may be outweighed by the benefits of maintaining thrust and flight speed.

Butterflies fly using efficient propulsive clap mechanism owing to flexible wings.

Butterflies look like no other flying animal, with unusually short, broad and large wings relative to their body size. Previous studies have suggested butterflies use several unsteady aerodynamic mechanisms to boost force production with upstroke wing clap being a prominent feature. When the wings clap together at the end of upstroke the air between the wings is pressed out, creating a jet, pushing the animal in the opposite direction. Although viewed, for the last 50 years, as a crucial mechanism in insect flight, quantitative aerodynamic measurements of the clap in freely flying animals are lacking. Using quantitative flow measurements behind freely flying butterflies during take-off and a mechanical clapper, we provide aerodynamic performance estimates for the wing clap. We show that flexible butterfly wings, forming a cupped shape during the upstroke and clap, thrust the butterfly forwards, while the downstroke is used for weight support. We further show that flexible wings dramatically increase the useful impulse (+22%) and efficiency (+28%) of the clap compared to rigid wings. Combined, our results suggest butterflies evolved a highly effective clap, which provides a mechanistic hypothesis for their unique wing morphology. Furthermore, our findings could aid the design of man-made flapping drones, boosting propulsive performance.

Power of the wingbeat: modelling the effects of flapping wings in vertebrate flight.

Animal flight performance has been studied using models developed for man-made aircraft. For an aeroplane with fixed wings, the energetic cost as a function of flight speed can be expressed in terms of weight, wing span, wing area and body area, where more details are included in proportionality coefficients. Flying animals flap their wings to produce thrust. Adopting the fixed wing flight model implicitly incorporates the effects of wing flapping in the coefficients. However, in practice, these effects have been ignored. In this paper, the effects of reciprocating wing motion on the coefficients of the fixed wing aerodynamic power model for forward flight are explicitly formulated in terms of thrust requirement, wingbeat frequency and stroke-plane angle, for optimized wingbeat amplitudes. The expressions are obtained by simulating flights over a large parameter range using an optimal vortex wake method combined with a low-level blade element method. The results imply that previously assumed acceptable values for the induced power factor might be strongly underestimated. The results also show the dependence of profile power on wing kinematics. The expressions introduced in this paper can be used to significantly improve animal flight models.

A preliminary investigation into a simple method for the determination of the mean ionisation energy of gas mixtures used in the NPL primary gas counting system.

The activity concentration of gaseous beta-emitting radionuclides such as (3)H, (85)Kr and, more recently, (11)C, is measured at NPL using a set of length-compensated proportional counters. The active gas is mixed with argon-methane (P-10) and passed to the counters. Adding gases to P-10 changes the mean ionisation energy, W, of the gas mixture. Estimation of the counting losses using the Monte Carlo model requires a knowledge of W. Unfortunately, only a limited amount of published data is available. This paper describes the initial experimental studies performed to enable the extension of the MC model based loss correction method to gases other than carbon dioxide in P-10. Preliminary measurements have been made to determine the W value for a gas mixture containing (85)Kr in nitrogen and P-10. The DC current through the counters is measured; the counters are also operated in the normal way with pulse amplifiers, discriminators and scalers. The value of W is derived from a knowledge of activity, counter current and mean beta energy.

A low noise preamplifier with optoelectronic overload protection for radioactivity measurement.

Pulses from detectors used for radioactivity measurement can vary in size by several orders of magnitude. Large pulses will lead to saturation at the preamplifier output and extension of the pulse length. As a consequence, the dead time of the system increases and pulses may be lost. Electronic design techniques employed to protect against overloading tend to increase the amplifier noise level. However, an optoelectronic method of overload protection has been devised which has only a negligible effect on noise. An infrared light emitting diode interfaced to the output of the preamplifier is linked by fibre optic cable to an ultra-low leakage photodiode at the input. The conduction of the photodiode increases with the amplitude of the preamplifier output signal. Excess current is thereby prevented from entering the preamplifier and causing saturation. The preamplifier has been tested on 4π beta-gamma and gas counting systems and found to give good protection against overloading.

Standardisation of 85Kr.

As part of a BIPM key-comparison of (85)Kr, a primary standardisation using internal gas proportional counting was performed. The supplied activity for the comparison was approximately 40 MBq and a two-stage dilution was required to reduce the activity concentration to a level suitable for gas counting. A new gas-handling rig was constructed for performing the dilutions. The dilutions, however, introduced significant uncertainties in the final result, so that additional methods suitable for measurement at higher activity levels were also used. A series of dilution ampoules with activities of 6 and 1 MBq were prepared in the new NPL gas-handling rig using inactive krypton as a diluent. Internal gas proportional counting was performed on each of the 1 MBq dilution ampoules. The proportion of the activity transferred to the counting system was estimated using pressure and volume data and the total ampoule activity calculated. Counting losses below the threshold were assumed to be 2%. The effect of changing the composition of the counting gas by inclusion of krypton was evaluated and found to not significantly change the gas gain, i.e. losses below the noise threshold (approximately 120 eV) remained essentially constant. The proportional counters were assumed to be 100% efficient with an uncertainty of 0.5% (k=1). Both 1 and 6 MBq ampoules were assayed by gamma-spectrometry using HPGe and NaI(Tl) detectors. This method resulted in an activity value with a smaller uncertainty than the primary method. Activity values for the three methods employed were consistent within the uncertainty of measurement.

Validation of a new TDCR system at NPL.

A new triple to double coincidence ratio (TDCR) system has been established at NPL. The system incorporates a spherical optical chamber, a manual sample changing facility and an integral light-tight housing. A 6 in diameter NaI(Tl) detector has been incorporated to allow 4pibeta-gamma coincidence measurements to be performed in parallel. Details of the detectors, the electronics and the overall TDCR system are given. Validation measurements using suitable low-energy beta and electron capture nuclides, i.e. (3)H and (55)Fe have been performed. The highest efficiency achieved with a (3)H solution in Ultima Gold AB in a glass vial was 53% and in a high-efficiency LS cocktail 65%. This indicates that the optical chamber is performing well. After varying the efficiency by de-focusing the PM tubes, the activity of the sources was determined. The activity concentration determined with TDCR agreed with certified values within the range of uncertainties. Further results from validation measurements and the corresponding uncertainty budgets are presented.

The vortex wake of blackcaps (Sylvia atricapilla L.) measured using high-speed digital particle image velocimetry (DPIV).

Reconstructing the vortex wake of freely flying birds is challenging, but in the past few years, direct measurements of the wake circulation have become available for a number of species. Streamwise circulation has been measured at different positions along the span of the birds, but no measurements have been performed in the transverse plane. Recent findings from studies of bat wakes have pointed to the importance of transverse plane data for reconstructing the wake topology because important structures may be missed otherwise. We present results of high-speed DPIV measurements in the transverse plane behind freely flying blackcaps. We found novel wake structures previously not shown in birds, including wing root vortices of opposite as well as the same sign as the wing tip vortices. This suggests a more complex wake structure in birds than previously assumed and calls for more detailed studies of the flow over the wings and body, respectively. Based on measurements on birds with and without a tail we also tested hypotheses regarding the function of the tail during steady flight. We were unable to detect any differences in the wake pattern between birds with and without a tail. We conclude that the birds do not use their tail to exploit vortices shed at the wing root during the downstroke. Neither did we find support for the hypothesis that the tail should reduce the drag of the bird. The function of the tail during steady flight thus remains unclear and calls for further investigation in future studies.

A low-noise current-sensitive amplifier-discriminator system for beta particle counting.

NPL has developed a low-noise current amplifier/discriminator system for radionuclides that emit low-energy electrons and X-rays. The new beta amplifier is based on the low-noise Amptek A-250 operational amplifier. The design has been configured for optimum signal to noise ratio. The new amplifier is described and results obtained using primarily electron-capture decaying radionuclides are presented. The new amplifier gives rise to higher particle detection efficiency than the previously used Atomic Energy of Canada Limited-designed amplifier. This is shown by measurements of (54)Mn and (65)Zn. The counting plateaux are significantly longer and have reduced gradients.

Leading-edge vortex improves lift in slow-flying bats.

Staying aloft when hovering and flying slowly is demanding. According to quasi-steady-state aerodynamic theory, slow-flying vertebrates should not be able to generate enough lift to remain aloft. Therefore, unsteady aerodynamic mechanisms to enhance lift production have been proposed. Using digital particle image velocimetry, we showed that a small nectar-feeding bat is able to increase lift by as much as 40% using attached leading-edge vortices (LEVs) during slow forward flight, resulting in a maximum lift coefficient of 4.8. The airflow passing over the LEV reattaches behind the LEV smoothly to the wing, despite the exceptionally large local angles of attack and wing camber. Our results show that the use of unsteady aerodynamic mechanisms in flapping flight is not limited to insects but is also used by larger and heavier animals.

Bat flight generates complex aerodynamic tracks.

The flapping flight of animals generates an aerodynamic footprint as a time-varying vortex wake in which the rate of momentum change represents the aerodynamic force. We showed that the wakes of a small bat species differ from those of birds in some important respects. In our bats, each wing generated its own vortex loop. Also, at moderate and high flight speeds, the circulation on the outer (hand) wing and the arm wing differed in sign during the upstroke, resulting in negative lift on the hand wing and positive lift on the arm wing. Our interpretations of the unsteady aerodynamic performance and function of membranous-winged, flapping flight should change modeling strategies for the study of equivalent natural and engineered flying devices.

Primary standardisation of 204Tl using the efficiency tracing method.

The electron-capture and beta(-) -emitting radionuclide 204Tl has been known from previous experience to show discrepancies between different methods of standardisation. Source preparation is also difficult due to the complex chemistry of thallium. The Bureau International des Poids et Mesures (BIPM) intercomparison held in 1997, showed discrepancies of up to 10% between measurements of solid sources and liquid scintillation methods. These problems have been the subject of a BIPM CCRI(II) working group since 1999. This paper presents the results from a primary standardisation of a 204Tl solution using 4pibeta-gamma coincidence counting and liquid scintillation counting. The tracer technique was used for the 4pibeta-gamma coincidence counting, where 204Tl was traced with 134Cs and 60Co. The extrapolation to 100% beta-efficiency was performed by three different approaches: foiling, adding carrier and measuring sources of different initial masses. The results showed that tracing with 60Co and using external foiling gave the same result as obtained by liquid scintillation counting. A comparison of the results achieved by the different methods of measurement, tracers and methods for extrapolation is presented and discussed in this paper.

Lift-based paddling in diving grebe.

To examine the hydrodynamic propulsion mechanism of a diving great crested grebe (Podiceps cristatus), the three-dimensional kinematics was determined by digital analysis of sequential video images of dorsal and lateral views. During the acceleration phase of this foot-propelled bird, the feet move through an arc in a plane nearly normal to the bird's line of motion through the water, i.e. the toes move dorsally and medially but not caudally relative to the water. The kinematics of the grebe's lobed feet is different from that in anseriforms, whose feet move in a plane mostly parallel to the bird's line of progress through the water. Our results suggest that the foot-propelled locomotor mechanism of grebes is based primarily on a lift-producing leg and foot stroke, in contrast to the drag-based locomotion assumed previously. We suggest that the lift-based paddling of grebes considerably increases both maximum swimming speed and energetic efficiency over drag-based propulsion. Furthermore, the results implicate a new interpretation of the functional morphology of these birds, with the toes serving as a self-stabilizing multi-slotted hydrofoil during the power phase.