Lift Production Research Articles

The novel aerodynamics of insect flight: applications to micro-air vehicles.

The wing motion in free flight has been described for insects ranging from 1 to 100 mm in wingspan. To support the body weight, the wings typically produce 2-3 times more lift than can be accounted for by conventional aerodynamics. Some insects use the fling mechanism: the wings are clapped together and then flung open before the start of the downstroke, creating a lift-enhancing vortex around each wing. Most insects, however, rely on a leading-edge vortex (LEV) created by dynamic stall during flapping; a strong spanwise flow is also generated by the pressure gradients on the flapping wing, causing the LEV to spiral out to the wingtip. Technical applications of the fling are limited by the mechanical damage that accompanies repeated clapping of the wings, but the spiral LEV can be used to augment the lift production of propellers, rotors and micro-air vehicles (MAVs). Design characteristics of insect-based flying machines are presented, along with estimates of the mass supported, the mechanical power requirement and maximum flight speeds over a wide range of sizes and frequencies. To support a given mass, larger machines need less power, but smaller ones operating at higher frequencies will reach faster speeds.

Discrimination of flying mimetic, passion–vine butterfliesHeliconius

Wing–beat frequency and the degree of asymmetry in wing motion were more similar among mimics than among sister species of passion–vine butterflies in the genus Heliconius . Asymmetry in wing motion is not attributed to lift production, and serves as the first clear example of a mimetic behavioural signal for a flying organism. Because the similarities in wing motion are too subtle for humans to observe with the naked eye, they serve as a previously unexplored mimetic signal.

Kinematics of flap-bounding flight in the zebra finch over a wide range of speeds

ABSTRACTIt has been proposed elsewhere that flap-bounding, an intermittent flight style consisting of flapping phases interspersed with flexed-wing bounds, should offer no savings in average mechanical power relative to continuous flapping unless a bird flies 1.2 times faster than its maximum range speed (Vmr). Why do some species use intermittent bounds at speeds slower than 1.2Vmr The ‘fixed-gear hypothesis’ suggests that flap-bounding is used to vary mean power output in small birds that are otherwise constrained by muscle physiology and wing anatomy to use a fixed muscle shortening velocity and pattern of wing motion at all flight speeds; the ‘body-lift hypothesis’ suggests that some weight support during bounds could make flap-bounding flight aerodynamically advantageous in comparison with continuous flapping over most forward flight speeds. To test these predictions, we studied high-speed film recordings (300 Hz) of wing and body motion in zebra finches (Taenopygia guttata, mean mass 13.2 g, N=4) taken as the birds flew in a variable-speed wind tunnel (0–14 m s−1). The zebra finches used flap-bounding flight at all speeds, so their flight style was unique compared with that of birds that facultatively shift from continuous flapping or flap-gliding at slow speeds to flap-bounding at fast speeds. There was a significant effect of flight speed on all measured aspects of wing motion except percentage of the wingbeat spent in downstroke. Changes in angular velocity of the wing indicated that contractile velocity in the pectoralis muscle changed with flight speed, which is not consistent with the fixed-gear hypothesis. Although variation in stroke-plane angle relative to the body, pronation angle of the wing and wing span at mid-upstroke showed that the zebra finch changed within-wingbeat geometries according to speed, a vortex-ring gait with a feathered upstroke appeared to be the only gait used during flapping. In contrast, two small species that use continuous flapping during slow flight (0–4 m s−1) either change wingbeat gait according to flight speed or exhibit more variation in stroke-plane and pronation angles relative to the body. Differences in kinematics among species appear to be related to wing design (aspect ratio, skeletal proportions) rather than to pectoralis muscle fiber composition, indicating that the fixed-gear hypothesis should perhaps be modified to exclude muscle physiology and to emphasize constraints due to wing anatomy. Body lift was produced during bounds at speeds from 4 to 14m s−1. Maximum body lift was 0.0206 N (15.9 % of body weight) at 10 m s−1; body lift:drag ratio declined with increasing air speed. The aerodynamic function of bounds differed with increasing speed from an emphasis on lift production (4–10 m s−1) to an emphasis on drag reduction with a slight loss in lift (12 and 14 m s−1). From a mathematical model of aerodynamic costs, it appeared that flap-bounding offered the zebra finch an aerodynamic advantage relative to continuous flapping at moderate and fast flight speeds (6–14 m s−1), with body lift augmenting any savings offered solely by flap-bounding at speeds faster than 7.1 m s−1. The percentage of time spent flapping during an intermittent flight cycle decreased with increasing speed, so the mechanical cost of transport was likely to be lowest at faster flight speeds (10–14 m s−1).

Utilisation of substrates during tethered flight with and without lift generation in the African fruit beetle Pachnoda sinuata (Cetoniinae).

We have investigated the pattern of metabolic changes during tethered flight with and without lift generation in the African fruit beetle Pachnoda sinuata. Two distinct metabolic phases occur during lift-generating flight. The first phase is characterised by a high rate of oxygen consumption and a rapid change in proline and alanine levels in the haemolymph and flight muscles and in glycogen level in the flight muscles. Carbohydrates are released from the fat body into the haemolymph. These carbohydrates are oxidised during the second phase. Changes in proline and alanine levels in the haemolymph and flight muscles and in glycogen level in the flight muscles are minor during the second phase and the rate of oxygen consumption is reduced. During lift-generating flight, metabolic changes are rapid. Proline concentrations in the haemolymph and flight muscles fall dramatically during the first 30 s of flight, while alanine concentrations rise concomitantly. While haemolymph concentrations of proline and alanine remain virtually unchanged thereafter, further changes in the levels of these amino acids occur in the flight muscles during 5 min of flight. The initial levels of the two amino acids in the flight muscles are re-established over 1 h of rest following a 5 min flight, while this process takes longer in the haemolymph. The concentration of haemolymph carbohydrates increases during the first 30 s of flight and declines thereafter during 5 min of flight. The pre-flight levels are restored after 1 h of subsequent rest. The stores of glycogen in the flight muscles are rapidly diminished during the first 10 s of flight and decrease at a lower rate during further flight lasting up to 5 min. A subsequent 1 h of rest is sufficient almost to restore pre-flight levels. Haemolymph lipid levels are slightly but significantly increased during 11 min of flight and after 1 h of subsequent rest. During flight without lift production, the metabolic changes are considerably slower and beetles fly approximately seven times longer than during lift-generating flight. Resting basalar (BM), dorso-ventral (DVM) and dorso-longitudinal (DLM) flight muscles show no differences in levels of proline, alanine and glycogen. After different periods of flight, during which lift and wing loading were minimised, the DVM was found to have the highest level of proline after 5 min of flight. Levels of alanine in the DVM were lower than in the DLM. There was no evidence to suggest that different flight muscles are specialised for either proline or carbohydrate utilisation. Proline and carbohydrates participate equally in supplying energy to the flight muscles during lift-generating flight. The contribution to the energy supply by the flight muscles is 54 %, while that of the haemolymph is 46 %.

A computational fluid dynamic study of hawkmoth hovering

A computational fluid dynamic (CFD) modelling approach is used to study the unsteady aerodynamics of the flapping wing of a hovering hawkmoth. We use the geometry of a Manduca sexta-based robotic wing to define the shape of a three-dimensional 'virtual' wing model and 'hover' this wing, mimicking accurately the three-dimensional movements of the wing of a hovering hawkmoth. Our CFD analysis has established an overall understanding of the viscous and unsteady flow around the flapping wing and of the time course of instantaneous force production, which reveals that hovering flight is dominated by the unsteady aerodynamics of both the instantaneous dynamics and also the past history of the wing. <P> A coherent leading-edge vortex with axial flow was detected during translational motions of both the up- and downstrokes. The attached leading-edge vortex causes a negative pressure region and, hence, is responsible for enhancing lift production. The axial flow, which is derived from the spanwise pressure gradient, stabilises the vortex and gives it a characteristic spiral conical shape. <P> The leading-edge vortex created during previous translational motion remains attached during the rotational motions of pronation and supination. This vortex, however, is substantially deformed due to coupling between the translational and rotational motions, develops into a complex structure, and is eventually shed before the subsequent translational motion. <P> Estimation of the forces during one complete flapping cycle shows that lift is produced mainly during the downstroke and the latter half of the upstroke, with little force generated during pronation and supination. The stroke plane angle that satisfies the horizontal force balance of hovering is 23.6 degrees , which shows excellent agreement with observed angles of approximately 20-25 degrees . The time-averaged vertical force is 40 % greater than that needed to support the weight of the hawkmoth.

Maturational changes in troponin T expression, Ca2+-sensitivity and twitch contraction kinetics in dragonfly flight muscle

Maximum lift production and the thermal sensitivity of lift production increase dramatically during adult maturation of Libellula pulchella dragonflies. Here, we report that the mechanistic basis for this transition appears to involve a developmental change in protein expression, which alters the Ca2+-sensitivity of muscle activation and twitch contraction kinetics. The alternatively spliced Ca2+ regulatory protein troponin T (TnT) undergoes an isoform shift during adult maturation. Skinned (demembranated) fibers of mature flight muscle are up to 13 times more sensitive to activation by Ca2+ than skinned fibers from teneral (newly emerged adult) flight muscle, and their Ca2+-sensitivity is more strongly affected by temperature. Intact muscle from mature individuals has a shorter time to peak tension and longer time to half-relaxation during twitch contractions, which is consistent with a greater Ca2+-sensitivity of mature muscle. Because it becomes activated more quickly and relaxes more slowly, mature flight muscle is able to generate, with each twitch, more force per unit area than teneral muscle; this difference in force becomes greater at high temperatures. There do not appear to be any age-related differences in actomyosin crossbridge properties, since teneral and mature flight muscles do not differ in shortening velocity, tetanic tension or instantaneous power output during isotonic contraction. Thus, variation in TnT expression appears to affect the temperature-dependent Ca2+-sensitivity of muscle activation, which in turn affects the kinetics and force production of the twitch contractions used by dragonflies during flight. This cascade of effects suggests that maturational changes in the expression of TnT isoforms may be a key determinant of overall muscle and organismal performance.

The changes in power requirements and muscle efficiency during elevated force production in the fruit fly Drosophila melanogaster.

The limits of flight performance have been estimated in tethered Drosophila melanogaster by modulating power requirements in a 'virtual reality' flight arena. At peak capacity, the flight muscles can sustain a mechanical power output of nearly 80 W kg-1 muscle mass at 24 degrees C, which is sufficient to generate forces of approximately 150% of the animal's weight. The increase in flight force above that required to support body weight is accompanied by a rise in wing velocity, brought about by an increase in stroke amplitude and a decrease in stroke frequency. Inertial costs, although greater than either profile or induced power, would be minimal with even modest amounts of elastic storage, and total mechanical power energy should be equivalent to aerodynamic power alone. Because of the large profile drag expected at low Reynolds numbers, the profile power was approximately twice the induced power at all levels of force generation. Thus, it is the cost of overcoming drag, and not the production of lift, that is the primary requirement for flight in Drosophila melanogaster. By comparing the estimated mechanical power output with respirometrically measured total power input, we determined that muscle efficiency rises with increasing force production to a maximum of 10%. This change in efficiency may reflect either increased crossbridge activation or a favorable strain regime during the production of peak forces.

The three–dimensional leading–edge vortex of a ‘hovering’ model hawkmoth

Recent flow visualisation experiments with the hawkmoth, Manduca sexta , revealed small but clear leading–edge vortex and a pronounced three–dimensional flow. Details of this flow pattern were studied with a scaled–up, robotic insect (‘the flapper’) that accurately mimicked the wing movements of a hovering hawkmoth. Smoke released from the leading edge of the flapper wing confirmed the existence of a small, strong and stable leading–edge vortex, increasing in size from wingbase to wingtip. Between 25 and 75 % of the wing length, its diameter increased approximately from 10 to 50 % of the wing chord. The leading–edge vortex had a strong axial flow veolocity, which stabilized it and reduced its diamater. The vortex separated from the wing at approximately 75 % of the wing length and thus fed vorticity into a large, tangled tip vortex. If the circulation of the leading–edge vortex were fully used for lift generation, it could support up to two–thirds of the hawkmoth's weight during the downstroke. The growth of this circulation with time and spanwise position clearly identify dynamic stall as the unsteady aerodynamic mechanism responsible for high lift production by hovering hawkmoths and possibly also by many other insect species.

Contributions of the propatagium to avian flight

Through flight experiments with live birds and computer modeling we define the aerodynamic contributions of the propatagium in avian flight. From flight trials we found that in House Sparrows, with all flight feathers removed except for the distal six primaries, the loss of approximately 50% of the propatagium's projected area and its cambered profile produced a significant reduction in the distance a bird was able to cover in flight. Removal of the secondary feathers, leaving six distal primaries and an intact propatagium, did not have a noticeable affect upon flight. From the computer model which is representative of the bird wing's mid-antebrachial chord (cambered propatagium, symmetrical musculoskeletal elements, and flat secondary flight feathers), we found that the propatagium: (1) produced the majority of the lift; (2) had a higher (relative to secondary feathers) production of lift in relation to its angle of attack, i.e., steeper lift-curve slope; and (3) produced more lift with a chord only 1/5 that of the feather subsection. We conclude that the cambered propatagium is the major lift generating component of the wing proximal to the wrist. © 1996 Wiley-Liss, Inc.

Flight kinematics of black-billed magpies and pigeons over a wide range of speeds

To investigate how birds that differ in morphology change their wing and body movements while flying at a range of speeds, we analyzed high-speed (60 Hz) video tapes of black-billed magpies (Pica pica) flying at speeds of 4-14 m s-1 and pigeons (Columba livia) flying at 6-20 m s-1 in a wind-tunnel. Pigeons had higher wing loading and higher-aspect-ratio wings compared with magpies. Both species alternated phases of steady-speed flight with phases of acceleration and deceleration, particularly at intermediate flight speeds. The birds modulated their wingbeat kinematics among these phases and frequently exhibited non-flapping phases while decelerating. Such modulation in kinematics during forward flight is typical of magpies but not of pigeons in the wild. The behavior of the pigeons may have been a response to the reduced power costs for flight in the closed wind-tunnel relative to those for free flight at similar speeds. During steady-speed flight, wingbeat frequency did not change appreciably with increasing flight speed. Body angle relative to the horizontal, the stroke-plane angles of the wingtip and wrist relative to the horizontal and the angle describing tail spread at mid-downstroke all decreased with increasing flight speed, thereby illustrating a shift in the dominant function of wing flapping from weight support at slow speeds to positive thrust at fast speeds. Using wingbeat kinematics to infer lift production, it appeared that magpies used a vortex-ring gait during steady-speed flight at all speeds whereas pigeons used a vortex-ring gait at 6 and 8 m s-1, a transitional vortex-ring gait at 10 m s-1, and a continuous-vortex gait at faster speeds. Both species used a vortex-ring gait for acceleration and a continuous-vortex gait or a non-flapping phase for deceleration during flight at intermediate wind-tunnel speeds. Pigeons progressively flexed their wings during glides as flight speed increased but never performed bounds. Wingspan during glides in magpies did not vary with flight speed, but the percentage of bounds among non-flapping intervals increased with speed from 10 to 14 m s-1. The use of non-flapping wing postures seemed to be related to the gaits used during flapping and to the aspect ratio of the wings. We develop an 'adverse-scaling' hypothesis in which it is proposed that the ability to reduce metabolic and mechanical power output using flap-bounding flight at fast flight speeds is scaled negatively with body mass. This represents an alternative to the 'fixed-gear' hypothesis previously suggested by other authors to explain the use of intermittent flight in birds. Future comparative studies in the field would be worthwhile, especially if instantaneous flight speeds and within-wingbeat kinematics were documented; new studies in the laboratory should involve simultaneous recording of wing kinematics and aerodynamic forces on the wing.

Physiological modelling of oxygen consumption in birds during flight

This study combines data on changes in cardiovascular variables with body mass (Mb) and with exercise intensity to model the oxygen supply available to birds during flight. Its main purpose is to provide a framework for identifying the factors involved in limiting aerobic power input to birds during flight and to suggest which cardiovascular variables are the most likely to have been influenced by natural selection when considering both allometric and adaptive variation. It is argued that natural selection has acted on heart rate (fh) and cardiac stroke volume (Vs), so that the difference in the arteriovenous oxygen content (CaO2-Cv¯O2) in birds, both at rest and during flight, is independent of Mb. Therefore, the Mb exponent for oxygen consumption (V(dot)O2) during flight can be estimated from measurements of heart rate and stroke volume. Stroke volume is likely to be directly proportional to heart mass (Mh) and, using empirical data, values for the Mb coefficients and exponents of various cardiovascular variables are estimated. It is concluded that, as found for mammals, fh is the main adaptive variable when considering allometric variation, although Mh also shows a slight scaling effect. Relative Mh is likely to be the most important when considering adaptive specialisations. The Fick equation may be represented as: (V(dot)O2)Mbz = (fh)Mbw x (Vs)Mbx x (CaO2 - Cv¯O2)Mby , where w, x, y, z are the body mass exponents for each variable and the terms in parentheses represent the Mb coefficients. Utilising this formula and data from the literature, the scaling of minimum V(dot)O2 during flight for bird species with a 'high aerobic capacity' (excluding hummingbirds) is calculated to be: 166Mb0.77&plusmn;0.09 = 574Mb-0.19&plusmn;0.02 x 3.48Mb0.96&plusmn;0.02 x 0.083Mb0.00&plusmn;0.05 , and for hummingbirds (considered separately owing to their unique wing kinematics) it is: 314Mb0.90&plusmn;0.22 = 617Mb-0.10&plusmn;0.06 x 6.13Mb1.00&plusmn;0.11 x 0.083Mb0.00&plusmn;0.05 . These results are largely dependent on the cardiovascular values obtained from pigeons flying near to the minimum power speed of 10 m s-1, but would appear to provide realistic values. Both the measured and the estimated V(dot)O2 for hummingbirds appear to scale with a larger Mb exponent than that for all other birds, and it is suggested that this is as a result of the larger Mb exponent for flight muscle mass as the larger species of hummingbirds try to maintain hovering performance. It is proposed that estimated V(dot)O2 for birds during flight, which is based on Mh in combination with estimates of fh and CaO2-Cv¯O2, gives an indirect measure of relative aerobic power input and, when corrected for the estimated scaling influences of the mechano-chemical conversion efficiency and lift generation with respect to Mb, may be a useful indicator of the relative capacity of the muscle to sustain power output and lift production during flight.

How a Long Tail and Changes in Mass and Wing Shape Affect the Cost for Flight in Animals

1. Relationships between foraging strategy, flight performance and wing shape in animals can be demonstrated with the use of aerodynamic theory. The optimal morphology is dictated not only by foraging behaviour and habitat selection but also by size of prey and migratory habits, as well as flight display. Here I demonstrate how changes in body size and structure of wings and tail affect the optimal flight speeds and power required to fly. 2. Long-tailed birds flying with their tails hanging downwards-backwards (e. g. widowbirds) are predicted to fly more slowly than short-tailed birds to save energy or to have larger wings (broader and/or longer wings giving lower wing loading) to compensate for increased tail drag during flight. Assuming that the tail length is five times as long in a long-tailed bird than in a short-tailed (normal) one, the minimum power and maximum range power would be about 30% higher in the long-tailed bird and the corresponding speeds about 40% slower. Ignoring wing inertial loads as a cost (inertial power) the corresponding percentages would become 16-18% higher powers and slower speeds. 3. I predict that members of a bird family that display in hovering or vertical take-off flights should not have elongated wings if wing inertia is important. In contrast to most other widowbird species, males of Jackson's widowbird (Euplectes jacksoni) do not have elongated wings (Anderson 1992), although they have particularly costly tails (Thomas 1993). This species has abandoned forward flight display and instead uses a vertical take-off and hovering flight display. Inertial costs appear to explain this result. 4. An increase in weight causes an increase in wing loading, which in turn requires higher flight speed for production of enough lift. A 50% increase in body mass (as fat, an egg or a fetus inside the body), would increase P mr and P mp by about 45-50% and V mr and V mp by about 20-25% (symbols defined in Fig. 1), when inertial power is taken to be zero. 5. A bird or bat carrying prey in the beak/mouth or claws should use about the same speed for minimization of flight costs as when flying without prey. 6. For low flight costs, and assuming a perfect elastic storage (zero inertial power), fast-flying species should benefit from short, narrow, high-aspect-ratio wings, hovering species should have long wings and slow-flying species should have larger wings (lower wing loadings) but with no particular demands on the aspect ratio. Ignoring inertial power the best flight economy is attained by the combination of a low wing loading (enabling slow flight) and high aspect ratio (Norberg & Rayner 1987; Norberg 1990). If we assume that inertial power is an important cost in hovering, a hovering animal should have short wings to minimize mechanical power

Unsteady Aerodynamic Performance of Model Wings at Low Reynolds Numbers

ABSTRACT The synthesis of a comprehensive theory of force production in insect flight is hindered in part by the lack of precise knowledge of unsteady forces produced by wings. Data are especially sparse in the intermediate Reynolds number regime (10&amp;lt;Re&amp;lt;1000) appropriate for the flight of small insects. This paper attempts to fill this deficit by quantifying the time-dependence of aerodynamic forces for a simple yet important motion, rapid acceleration from rest to a constant velocity at a fixed angle of attack. The study couples the measurement of lift and drag on a two-dimensional model with simultaneous flow visualization. The results of these experiments are summarized below. At angles of attack below 13.5°, there was virtually no evidence of a delay in the generation of lift, in contrast to similar studies made at higher Reynolds numbers.At angles of attack above 13.5°, impulsive movement resulted in the production of a leading edge vortex that stayed attached to the wing for the first 2 chord lengths of travel, resulting in an 80% increase in lift compared to the performance measured 5 chord lengths later. It is argued that this increase is due to the process of detached vortex lift, analogous to the method of force production in delta-wing aircraft.As the initial leading edge vortex is shed from the wing, a second vortex of opposite vorticity develops from the trailing edge of the wing, correlating with a decrease in lift production. This pattern of alternating leading and trailing edge vortices generates a von Karman street, which is stable for at least 7.5 chord lengths of travel.Throughout the first 7.5 chords of travel the model wing exhibits a broad lift plateau at angles of attack up to 54°, which is not significantly altered by the addition of wing camber or surface projections.Taken together, these results indicate how the unsteady process of vortex generation at large angles of attack might contribute to the production of aerodynamic forces in insect flight. Because the fly wing typically moves only 2–4 chord lengths each half-stroke, the complex dynamic behavior of impulsively started wing profiles is more appropriate for models of insect flight than are steady-state approximations.

Limitations on Animal Flight Performance

ABSTRACT Flight performance seems to change systematically with body size: small animals can hover and fly over a wide range of speeds, but large birds taxi for takeoff and then fly over a narrow speed range. The traditional explanation for this is that the mass-specific power required for flight varies with speed according to a U-shaped curve, and it also scales between m0 and m1/6, where m is body mass. The mass-specific power available from the flight muscles is assumed to scale as m−1/3 As available power decreases with increasing body size, the range of attainable flight speeds becomes progressively reduced until the largest animals can only fly in the trough of the U-shaped curve. Above a particular size, the available power is insufficient and flapping flight is not possible. The underlying assumptions of this argument are examined in this review. Metabolic measurements are more consistent with a J-shaped curve, with little change in power from hovering to intermediate flight speeds, than with a U-shaped curve. Scaling of the mass-specific power required to fly agrees with predictions. The mass-specific power available from, the muscles, estimated from maximal loading studies, varies as m0.13. This scaling cannot be distinguished from that of the power required to fly, refuting the argument that power imposes an intrinsic scaling on flight performance. It is suggested instead that limitations on low-speed performance result from an adverse scaling of lift production with increasing body size

Kinematics of take‐off and climbing flight in butterflies

High speed flash photography (flash duration 0.1 ms) was used to analyse wing movements in over 30 species of butterfly. With few exceptions, the insects showed a clap and peel mechanism of lift production at the start of the downstroke. Early in the upstroke the wings showed pronounced ventral flexure which, combined with inertial lag in the posterior parts of both wing pairs and delayed supination in the hind wing, led to the formation of a funnel‐like space between the wings. These movements, and the resultant airflow patterns, appear to be an axi‐symmetric equivalent of the ‘near’ clap and peel (here referred to as the funnel). Hind wing movements throughout the stroke are hinged upon the claval furrow. The expanded anal lobes of the hind wing lying medially to the claval furrow help to provide an air‐tight seal around the abdomen between the upper and lower wing surfaces, which increases the efficiency of the peel and funnel mechanisms. The role of the intercalary flexion lines in controlling changes in wing surface corrugation during the cycle is also investigated.

An Overview of Waterflood Surveillance and Monitoring

Distinguished Author Series articles are general, descriptiverepresentations that summarize the state of the art in an area of technology bydescribing recent developments for readers who are not specialists in thetopics discussed. Written by individuals recognized as experts in the area, these articles provide key references to more definitive work and presentspecific details only to illustrate the technology. Purpose: to informthe general readership of recent advances in various areas of petroleumengineering. Introduction An essential key to a successful waterflooding project is a well-planned andwell-executed program of surveillance and monitoring. This program should betailored to address individual projects or fields because each waterflood willhave different characteristics. There are, however, some basic ingredients thatshould be common to all surveillance programs. In general, three majorcategories of field conditions must be included in any waterflood surveillanceprogram: reservoir conditions, injection/production-well conditions, andfacilities/operating conditions. The last and probably the most importantingredient is record keeping/performance control. There are, of course, economic conditions that must be taken into consideration. For the purposes ofthis paper, however, only the technical aspects of a waterflood surveillanceprogram will be addressed. The purpose of this paper is to provide an overviewof waterflood surveillance programs and to outline the various items to bemonitored. Selected tests for diagnosis of problems commonly associated withwaterfloods are discussed briefly, along with a look at developing surveillancetechnology. Surveillance Program Waterflood surveillance programs of the 1980's have been influenced to somedegree by the chemical waterflood projects of the 1970's. The desire tounderstand chemical-recovery-process applications better led to a significantincrease in project surveillance activities. Much time and effort was spent inevaluating both waterflood and tertiary recoveries. It was shown that byclosely monitoring field activities, improvement in waterflood recoveries couldbe achieved. Fig. 1 shows the key monitoring points in the traditionalwaterflood cycle. There was a time when most of the attention given awaterflood project focused on reservoir performance, and this usually waslimited to monitoring water cuts. Today we realize that it is equally importantto include well, facilities, and operating conditions in our surveillanceprograms. Thus, all the components of the waterflood cycle diagrammed in Fig. 1should be included in a well-planned surveillance program. Table 1 lists itemsthat should normally be included in the three major categories of surveillance. This is by no means a complete listing because waterfloods can have ratherunusual characteristics or conditions requiring additional items forobservation and evaluation-e.g., environmental and regulatory conditions. Reservoir Surveillance As listed in Table 1, reservoir pressures, injection and production rates, fluid volumes, WOR/GOR's, and fluid samples require constant surveillance. Adiscussion of these items and proposed schedules for obtaining the data waspresented by Barnes and Tinker. When these data are coupled with thereservoir-description information, waterflood-performance calculations can bemade. Reservoir-description information generally includes core, well log, andgeologic data. Holbert and Zeito list the reservoir-description data in moredetail. Numerous methods or techniques for estimating waterflood performancehave been reported in the literature. These methods range from the classictypes to sophisticated reservoir simulator models. There are four types ofwells requiring surveillance: production, injection, water-supply, andwater-disposal wells. Of these, production and injection wells require the mostattention. Monitoring well performance requires a program of selected welltests to be conducted regularly. The types of well tests selected will dependon surface/downhole equipment, well-completion characteristics, produced orinjected fluids, the stage of the waterflood project (early, middle, or late), and the reservoir description. Key items for surveillance are fluid entry intoor exit from target zones, cement/completion integrity, and mechanicalequipment, both downhole and surface. Well testing is discussed later. The wellsurveillance program should include plans for recording the information in asuitable manner or format such that it is both easily accessible and "userfriendly." Finally, the program should provide a systematic approach to dataanalysis, evaluation, recommendations, and corrective measures, as needed. Facilities/Operations Surveillance Waterflood operating procedures and conditions, along with the associatedproject facilities, are often taken for granted, yet they are key ingredientsto successful project management. Tinkers reported on the operating factorsthat affect waterflood performance, including well completions, injectionpatterns, high-volume lift, injection profiles, and bottomwater production. Operations and facilities vary considerably from project to project and undergochanges during the several stages of waterflood development. Injection-patternconfigurations, surface topography, reservoir characteristics, deviated wells, and field operating constraints are only a few of the conditions that can leadto problems associated with project management.

Forewing movements and motor activity during roll manoeuvers in flying desert locusts

Desert locusts, tethered on a roll torque meter and flying in a wind tunnel are surrounded by an artificial horizon (Fig. 1). Flight motor activity and movement of forewings are monitored continuously. Movements of the artificial horizon elicit roll manoeuvers of the animal with latencies of several seconds; concomitant changes in flight motor pattern and wing movement can be correlated with the animal's roll angle and roll torque. Flight sequences with constant torque and roll angle (steady state) have been analysed with the following results. Wing Kinematics. A phase difference between the movements of the forewings on either side is correlated with roll angle (Fig. 3). Pronation of a forewing is always greater on the side to which the animal rolls, i.e. on the side that produces less lift (Fig. 5). In some experiments the slope of the wing tip path is also decreased (Fig. 5). In both cases, the aerodynamic angle of attack is decreased and the forewing on this side produces less lift. In most experiments, changes in pronation are less pronounced in the contralateral wing (Fig. 11). All factors contribute to a net roll torque that sustains the animal's roll angle. Other kinematic parameters of forewing movement, e.g. wing stroke amplitude, were not found to be correlated with roll angle and torque (Fig. 4). Motor Pattern. Activity of several flight muscles (depressors M97, M98, M99, and M129; elevators M83, M84, and M90) was investigated for changes in burst length and temporal coordination in response to roll stimuli. Most flight muscles fired only once per wing beat cycle in our preparations. Thus, burst length was not found to be correlated with roll angle. Time intervals of firing between all muscle pairs investigated change in correlation with the torque and roll angle (Fig. 9). All mesothoracic muscles are active earlier-relative to the ipsilateral metathoracic subalar muscle M129-during roll to the ipsilateral side than during roll to the contralateral side. Correlations Between Motor and Movement Pattern. The phase of muscle firing within the wing beat cycle varies with roll angle (roll torque). The first basalar M97 and second tergosternal M84 muscles, when referenced e.g. to the upper (M97) or lower (M84) reversal point of the wing tip trajectory, are active earlier on the side the animal turns to (Fig. 10). The onset of the first basalar M97 relative to the beginning of downstroke is correlated with maximum pronation and roll angle (Fig. 11). Mechanisms of Lift Control. Wing pronation, which is very important for lift production is controlled by the central nervous system by altering the phase of muscle activity within the wing beat cycle.

The Forces Acting on Swimming Squid

ABSTRACT Analysis of ciné films and intramantle pressure records for squid Loligo opalescens Berry swimming in a tunnel respirometer provided estimates of all the forces acting in the horizontal and vertical planes for swimming speeds from 0·1 to 0·5 ms−1. Different speeds used different gaits; fin thrust was only important below 0·2 ms−1, ‘anaerobic’ circular muscles were recruited only at supracritical speeds, and hyperinflation caused by contraction of the radial muscle was not seen in steady swimming. The extent, rate and frequency of contraction of the obliquely striated circular muscles varied little with speed, and jet thrust was matched to speed primarily by active pressure control through adjustments in the size of the funnel orifice. Hydrodynamic lift production to compensate for negative buoyancy during enforced horizontal swimming in the tunnel required 30–90% of the total force over the speed range studied and appears less efficient than direct use of jet thrust. This suggests a new rationale for ‘climb-and-glide’ swimming which reduces previous estimates of the gross cost of transport for squid under natural conditions by at least 35%, with no loss of speed. The cost of accelerating water into the mantle of a squid moving at high speed appears to have been underestimated in previous studies. A simulation of a series of escape jets predicts a maximum speed of 8 body lengths s−1 (14ms−1), reached after only two jets, because of the high deceleration during refilling.

Maximum Lift Production During Takeoff in Flying Animals

ABSTRACT Maximum lift production during takeoff in still air was determined for a wide variety of insects and a small sample of birds and bats, and was compared with variation in morphology, taxonomy and wingbeat type. Maximum lift per unit flight muscle mass was remarkably similar between taxonomic groups (54–63 N kg−1), except for animals using clap-and-fling wingbeats, where muscle mass-specific lift increased by about 25 % (72–86 N kg−1). Muscle mass-specific lift was independent of body mass, wing loading, disk loading and aspect ratio. Birds and bats yielded results indistinguishable from insects using conventional wingbeats. Interspecific differences in short-duration powered flight and takeoff ability are shown to be caused primarily by differences in flight muscle ratio, which ranges from 0·115 to 0·560 among species studied to date. These results contradict theoretical predictions that maximum mass-specific power output and lift production should decrease with increasing body mass and wing disk loading.

Integration of Wing Proprioceptive and Descending Exteroceptive Sensory Inputs by Thoracic Interneurones of the Locust

ABSTRACT The campaniform sensilla on the wings of the locust are strain-sensitive mechanoreceptors that provide proprioceptive feedback about wing forces, particularly aerodynamic lift, experienced during flight. They can be excited by imposed deformations of the wing, including those caused by imposed wing twisting. The afferents of the single subcostal group of sensilla on the hindwing had the same directional selectivity for supinating twist and shared the properties of a dynamic sensitivity and adaptation. A group of strain-sensitive mechanoreceptors with similar properties, presumably campaniform sensilla, is also found in the forewings. Four types of thoracic interneurones influenced by these factors were recorded and stained intracellularly. The response of interneurone 5AA to imposed de-formations of the hindwing ipsilateral to its soma is determined by excitatory chemical synaptic input from the campaniform sensilla. Interneurone and sensilla have a common directional selectivity and optimal stimulus, and similar qualitative dynamics of response. Each spike of individual afferents is followed at short, constant latency by an excitatory postsynaptic potential (EPSP) in the interneurone, even at instantaneous frequencies of about 90 Hz. Physiological evidence is consistent with direct, chemically mediated synaptic inputs from campaniform sensilla afferents. Interneurone 5AA is also excited by a short-latency, chemical synaptic input from the ocelli when lights are turned off. EPSPs could be elicited by light-off stimuli to the median and contralateral, but not the ipsilateral, ocelli. In addition, the interneurone is excited when the head is moved relative to the thorax. The other three interneurones respond to strains in more than one wing. Inputs are derived from specific combinations of wings, with the sign of response depending on the neurone and the particular wing. Interneurones 3AA and 1AA are also phasically excited by light-off stimuli. In 1AA this response was shown to originate from the ocelli. Median and contralateral, but not ipsilateral, ocelli could evoke EPSPs. This neurone was also excited by imposed head movements. It is argued that the interneurones described here at suited to monitor lift production in particular wings and its pattern among several wings. Convergence of ocellar and head-motion inputs implies a function in the exteroceptive detection and correction of flight instability. It is inferred that these thoracic interneurones may act as the nexus for several different feedback pathways, proprioceptive and exteroceptive, which modulate flight motor output.