Dynamic shapes of the zygote and two-cell mouse and human.

Mouse zygote morphokinetics were measured during interphase, the mitotic period, cytokinesis, and two-cell stage. Sequences of rounder-distorted-rounder shapes were revealed, as were changing patterns of cross section area. A calcium chelator and an actin-disrupting agent inhibited the area changes that occurred between pronuclear envelope breakdown and cytokinesis. During cell division, two vortices developed in each nascent cell and they rotated in opposite directions at each end of the cell, a pattern that sometimes persisted for up to 10 h. Exchange with the environment may have been promoted by these shape and area cycles and persisting circulation in the cytoplasm may have a similar function between a cell's interior and periphery. Some of these movements were sporadically also seen in human zygotes with abnormal numbers of pronuclei and the two-cell stages that developed from these compromised human zygotes.

Terminal attack trajectories of peregrine falcons are described by the proportional navigation guidance law of missiles.

The ability to intercept uncooperative targets is key to many diverse flight behaviors, from courtship to predation. Previous research has looked for simple geometric rules describing the attack trajectories of animals, but the underlying feedback laws have remained obscure. Here, we use GPS loggers and onboard video cameras to study peregrine falcons, Falco peregrinus, attacking stationary targets, maneuvering targets, and live prey. We show that the terminal attack trajectories of peregrines are not described by any simple geometric rule as previously claimed, and instead use system identification techniques to fit a phenomenological model of the dynamical system generating the observed trajectories. We find that these trajectories are best-and exceedingly well-modeled by the proportional navigation (PN) guidance law used by most guided missiles. Under this guidance law, turning is commanded at a rate proportional to the angular rate of the line-of-sight between the attacker and its target, with a constant of proportionality (i.e., feedback gain) called the navigation constant (N). Whereas most guided missiles use navigation constants falling on the interval 3 ≤ N ≤ 5, peregrine attack trajectories are best fitted by lower navigation constants (median N < 3). This lower feedback gain is appropriate at the lower flight speed of a biological system, given its presumably higher error and longer delay. This same guidance law could find use in small visually guided drones designed to remove other drones from protected airspace.

Soaring energetics and glide performance in a moving atmosphere.

Here, we analyse the energetics, performance and optimization of flight in a moving atmosphere. We begin by deriving a succinct expression describing all of the mechanical energy flows associated with gliding, dynamic soaring and thermal soaring, which we use to explore the optimization of gliding in an arbitrary wind. We use this optimization to revisit the classical theory of the glide polar, which we expand upon in two significant ways. First, we compare the predictions of the glide polar for different species under the various published models. Second, we derive a glide optimization chart that maps every combination of headwind and updraft speed to the unique combination of airspeed and inertial sink rate at which the aerodynamic cost of transport is expected to be minimized. With these theoretical tools in hand, we test their predictions using empirical data collected from a captive steppe eagle (Aquila nipalensis) carrying an inertial measurement unit, global positioning system, barometer and pitot tube. We show that the bird adjusts airspeed in relation to headwind speed as expected if it were seeking to minimize its aerodynamic cost of transport, but find only weak evidence to suggest that it adjusts airspeed similarly in response to updrafts during straight and interthermal glides.This article is part of the themed issue 'Moving in a moving medium: new perspectives on flight'.

Wing tucks are a response to atmospheric turbulence in the soaring flight of the steppe eagle Aquila nipalensis.

Turbulent atmospheric conditions represent a challenge to stable flight in soaring birds, which are often seen to drop their wings in a transient motion that we call a tuck. Here, we investigate the mechanics, occurrence and causation of wing tucking in a captive steppe eagle Aquila nipalensis, using ground-based video and onboard inertial instrumentation. Statistical analysis of 2594 tucks, identified automatically from 45 flights, reveals that wing tucks occur more frequently under conditions of higher atmospheric turbulence. Furthermore, wing tucks are usually preceded by transient increases in airspeed, load factor and pitch rate, consistent with the bird encountering a headwind gust. The tuck itself immediately follows a rapid drop in angle of attack, caused by a downdraft or nose-down pitch motion, which produces a rapid drop in load factor. Positive aerodynamic loading acts to elevate the wings, and the resulting aerodynamic moment must therefore be balanced in soaring by an opposing musculoskeletal moment. Wing tucking presumably occurs when the reduction in the aerodynamic moment caused by a drop in load factor is not met by an equivalent reduction in the applied musculoskeletal moment. We conclude that wing tucks represent a gust response precipitated by a transient drop in aerodynamic loading.

Wake development behind paired wings with tip and root trailing vortices: consequences for animal flight force estimates.

Recent experiments on flapping flight in animals have shown that a variety of unrelated species shed a wake behind left and right wings consisting of both tip and root vortices. Here we present an investigation using Particle Image Velocimetry (PIV) of the behaviour and interaction of trailing vortices shed by paired, fixed wings that simplify and mimic the wake of a flying animal with a non-lifting body. We measured flow velocities at five positions downstream of two adjacent NACA 0012 aerofoils and systematically varied aspect ratio, the gap between the wings (corresponding to the width of a non-lifting body), angle of attack, and the Reynolds number. The range of aspect ratios and Reynolds number where chosen to be relevant to natural fliers and swimmers, and insect flight in particular. We show that the wake behind the paired wings deformed as a consequence of the induced flow distribution such that the wingtip vortices convected downwards while the root vortices twist around each other. Vortex interaction and wake deformation became more pronounced further downstream of the wing, so the positioning of PIV measurement planes in experiments on flying animals has an important effect on subsequent force estimates due to rotating induced flow vectors. Wake deformation was most severe behind wings with lower aspect ratios and when the distance between the wings was small, suggesting that animals that match this description constitute high-risk groups in terms of measurement error. Our results, therefore, have significant implications for experimental design where wake measurements are used to estimate forces generated in animal flight. In particular, the downstream distance of the measurement plane should be minimised, notwithstanding the animal welfare constraints when measuring the wake behind flying animals.

Operation of the alula as an indicator of gear change in hoverflies.

The alula is a hinged flap found at the base of the wings of most brachyceran Diptera. The alula accounts for up to 10 per cent of the total wing area in hoverflies (Syrphidae), and its hinged arrangement allows the wings to be swept back over the thorax and abdomen at rest. The alula is actuated via the third axillary sclerite, which is a component of the wing hinge that is involved in wing retraction and control. The third axillary sclerite has also been implicated in the gear change mechanism of flies. This mechanism allows rapid switching between different modes of wing kinematics, by imposing or removing contact with a mechanical stop limiting movement of the wing during the lower half of the downstroke. The alula operates in two distinct states during flight-flipped or flat-and we hypothesize that its state indicates switching between different flight modes. We used high-speed digital video of free-flying hoverflies (Eristalis tenax and Eristalis pertinax) to investigate whether flipping of the alula was associated with changes in wing and body kinematics. We found that alula state was associated with different distributions of multiple wing kinematic parameters, including stroke amplitude, stroke deviation angle, downstroke angle of incidence and timing of supination. Changes in all of these parameters have previously been linked to gear change in flies. Symmetric flipping of the alulae was associated with changes in the symmetric linear acceleration of the body, while asymmetric flipping of the alulae was associated with asymmetric angular acceleration of the body. We conclude that the wings produce less aerodynamic force when the alula is flipped, largely as a result of the accompanying changes in wing kinematics. The alula changes state at mid-downstroke, which is the point at which the gear change mechanism is known to come into effect. This transition is accompanied by changes in the other wing kinematic parameters. We therefore find that the state of the alula is linked to the same parameters as are affected by the gear change mechanism. We conclude that the state of the alula does indeed indicate the operation of different flight modes in Eristalis, and infer that a likely mechanism for these changes in flight mode is the gear change mechanism.

Deformable wing kinematics in free-flying hoverflies.

Here, we present a detailed analysis of the deforming wing kinematics of free-flying hoverflies (Eristalis tenax, Linnaeus) during hovering flight. We used four high-speed digital video cameras to reconstruct the motion of approximately 22 points on each wing using photogrammetric techniques. While the root-flapping motion of the wing is similar in both the downstroke and upstroke, and is well modelled as a simple harmonic motion, other wing kinematic parameters show substantial variation between the downstroke and upstroke. Whereas the magnitude of the angle of incidence varies considerably within and between different hoverflies, the twist distribution along the wing is highly stereotyped. The angle of incidence and camber both show a recoil effect as they change abruptly at stroke reversal. Pronation occurs consistently after stroke reversal, which is perhaps surprising, because this has been found to reduce lift production in modelling studies. We find that the alula, a hinged flap near the base of the wing, operates in two discrete states: either in plane with the wing, or flipped approximately normal to it. We hypothesize that the alula may be acting as a flow-control device.

Details of insect wing design and deformation enhance aerodynamic function and flight efficiency.

Insect wings are complex structures that deform dramatically in flight. We analyzed the aerodynamic consequences of wing deformation in locusts using a three-dimensional computational fluid dynamics simulation based on detailed wing kinematics. We validated the simulation against smoke visualizations and digital particle image velocimetry on real locusts. We then used the validated model to explore the effects of wing topography and deformation, first by removing camber while keeping the same time-varying twist distribution, and second by removing camber and spanwise twist. The full-fidelity model achieved greater power economy than the uncambered model, which performed better than the untwisted model, showing that the details of insect wing topography and deformation are important aerodynamically. Such details are likely to be important in engineering applications of flapping flight.

Deformable wing kinematics in the desert locust: how and why do camber, twist and topography vary through the stroke?

Here, we present a detailed analysis of the wing kinematics and wing deformations of desert locusts (Schistocerca gregaria, Forskål) flying tethered in a wind tunnel. We filmed them using four high-speed digital video cameras, and used photogrammetry to reconstruct the motion of more than 100 identified points. Whereas the hindwing motions were highly stereotyped, the forewing motions showed considerable variation, consistent with a role in flight control. Both wings were positively cambered on the downstroke. The hindwing was cambered through an 'umbrella effect' whereby the trailing edge tension compressed the radial veins during the downstroke. Hindwing camber was reversed on the upstroke as the wing fan corrugated, reducing the projected area by 30 per cent, and releasing the tension in the trailing edge. Both the wings were strongly twisted from the root to the tip. The linear decrease in incidence along the hindwing on the downstroke precisely counteracts the linear increase in the angle of attack that would otherwise occur in root flapping for an untwisted wing. The consequent near-constant angle of attack is reminiscent of the optimum for a propeller of constant aerofoil section, wherein a linear twist distribution allows each section to operate at the unique angle of attack maximizing the lift to drag ratio. This implies tuning of the structural, morphological and kinematic parameters of the hindwing for efficient aerodynamic force production.

Photogrammetric reconstruction of high-resolution surface topographies and deformable wing kinematics of tethered locusts and free-flying hoverflies.

Here, we present a suite of photogrammetric methods for reconstructing insect wing kinematics, to provide instantaneous topographic maps of the wing surface. We filmed tethered locusts (Schistocerca gregaria) and free-flying hoverflies (Eristalis tenax) using four high-speed digital video cameras. We digitized multiple natural features and marked points on the wings using manual and automated tracking. Epipolar geometry was used to identify additional points on the hoverfly wing outline which were anatomically indistinguishable. The cameras were calibrated using a bundle adjustment technique that provides an estimate of the error associated with each individual data point. The mean absolute three-dimensional measurement error was 0.11 mm for the locust and 0.03 mm for the hoverfly. The error in the angle of incidence was at worst 0.51 degrees (s.d.) for the locust and 0.88 degrees (s.d.) for the hoverfly. The results we present are of unprecedented spatio-temporal resolution, and represent the most detailed measurements of insect wing kinematics to date. Variable spanwise twist and camber are prominent in the wingbeats of both the species, and are of such complexity that they would not be adequately captured by lower resolution techniques. The role of spanwise twist and camber in insect flight has yet to be fully understood, and accurate insect wing kinematics such as we present here are required to be sure of making valid predictions about their aerodynamic effects.

New experimental approaches to the biology of flight control systems.

Here we consider how new experimental approaches in biomechanics can be used to attain a systems-level understanding of the dynamics of animal flight control. Our aim in this paper is not to provide detailed results and analysis, but rather to tackle several conceptual and methodological issues that have stood in the way of experimentalists in achieving this goal, and to offer tools for overcoming these. We begin by discussing the interplay between analytical and empirical methods, emphasizing that the structure of the models we use to analyse flight control dictates the empirical measurements we must make in order to parameterize them. We then provide a conceptual overview of tethered-flight paradigms, comparing classical ;open-loop' and ;closed-loop' setups, and describe a flight simulator that we have recently developed for making flight dynamics measurements on tethered insects. Next, we provide a conceptual overview of free-flight paradigms, focusing on the need to use system identification techniques in order to analyse the data they provide, and describe two new techniques that we have developed for making flight dynamics measurements on freely flying birds. First, we describe a technique for obtaining inertial measurements of the orientation, angular velocity and acceleration of a steppe eagle Aquila nipalensis in wide-ranging free flight, together with synchronized measurements of wing and tail kinematics using onboard instrumentation and video cameras. Second, we describe a photogrammetric method to measure the 3D wing kinematics of the eagle during take-off and landing. In each case, we provide demonstration data to illustrate the kinds of information available from each method. We conclude by discussing the prospects for systems-level analyses of flight control using these techniques and others like them.

Automatic aeroelastic devices in the wings of a steppe eagle Aquila nipalensis.

Here we analyse aeroelastic devices in the wings of a steppe eagle Aquila nipalensis during manoeuvres. Chaotic deflections of the upperwing coverts observed using video cameras carried by the bird (50 frames s(-1)) indicate trailing-edge separation but attached flow near the leading edge during flapping and gust response, and completely stalled flows upon landing. The underwing coverts deflect automatically along the leading edge at high angle of attack. We use high-speed digital video (500 frames s(-1)) to analyse these deflections in greater detail during perching sequences indoors and outdoors. Outdoor perching sequences usually follow a stereotyped three-phase sequence comprising a glide, pitch-up manoeuvre and deep stall. During deep stall, the spread-eagled bird has aerodynamics reminiscent of a cross-parachute. Deployment of the underwing coverts is closely phased with wing sweeping during the pitch-up manoeuvre, and is accompanied by alula protraction. Surprisingly, active alula protraction is preceded by passive peeling from its tip. Indoor flights follow a stereotyped flapping perching sequence, with deployment of the underwing coverts closely phased with alula protraction and the end of the downstroke. We propose that the underwing coverts operate as an automatic high-lift device, analogous to a Kruger flap. We suggest that the alula operates as a strake, promoting formation of a leading-edge vortex on the swept hand-wing when the arm-wing is completely stalled, and hypothesise that its active protraction is stimulated by its initial passive deflection. These aeroelastic devices appear to be used for flow control to enhance unsteady manoeuvres, and may also provide sensory feedback.

Digital particle image velocimetry measurements of the downwash distribution of a desert locust Schistocerca gregaria.

Actuator disc models of insect flight are concerned solely with the rate of momentum transfer to the air that passes through the disc. These simple models assume that an even pressure is applied across the disc, resulting in a uniform downwash distribution. However, a correction factor, k, is often included to correct for the difference in efficiency between the assumed even downwash distribution, and the real downwash distribution. In the absence of any empirical measurements of the downwash distribution behind a real insect, the values of k used in the literature have been necessarily speculative. Direct measurement of this efficiency factor is now possible, and could be used to compare the relative efficiencies of insect flight across the Class. Here, we use Digital Particle Image Velocimetry to measure the instantaneous downwash distribution, mid-downstroke, of a tethered desert locust (Schistocerca gregaria). By integrating the downwash distribution, we are thereby able to provide the first direct empirical measurement of k for an insect. The measured value of k = 1.12 corresponds reasonably well with that predicted by previous theoretical studies.

The aerodynamics of Manduca sexta: digital particle image velocimetry analysis of the leading-edge vortex.

Here we present the first digital particle image velocimetry (DPIV) analysis of the flow field around the wings of an insect (the tobacco hawkmoth Manduca sexta, tethered to a 6-component force-moment balance in a wind tunnel). A leading-edge vortex (LEV) is present above the wings towards the end of the downstroke, as the net upward force peaks. Our DPIV analyses and smoke visualisations match the results of previous flow visualisation experiments at midwing, and we extend the experiments to provide the first analysis of the flow field above the thorax. Detailed DPIV measurements show that towards the end of the downstroke, the LEV structure is consistent with that recently reported in free-flying butterflies and dragonflies: the LEV is continuous across the thorax and runs along each wing to the wingtip, where it inflects to form the wingtip trailing vortices. The LEV core is 2-3 mm in diameter (approximately 10% of local wing chord) both at the midwing position and over the centreline at 1.2 m s(-1) and at 3.5 m s(-1) flight speeds. At 1.2 m s(-1) the measured LEV circulation is 0.012+/-0.001 m(2) s(-1) (mean +/-S.D.) at the centreline and 0.011+/-0.001 m(2) s(-1) halfway along the wing. At 3.5 m s(-1) LEV circulation is 0.011+/-0.001 m(2) s(-1) at the centreline and 0.020+/-0.004 m(2) s(-1) at midwing. The DPIV measurements suggest that if there is any spanwise flow in the LEV towards the end of the downstroke its velocity is less than 1 m s(-1). Estimates of force production show that the LEV contributes significantly to supporting body weight during bouts of flight at both speeds (more than 10% of body weight at 1.2 m s(-1) and 35-65% of body weight at 3.5 m s(-1)).

Dragonfly flight: free-flight and tethered flow visualizations reveal a diverse array of unsteady lift-generating mechanisms, controlled primarily via angle of attack.

Here we show, by qualitative free- and tethered-flight flow visualization, that dragonflies fly by using unsteady aerodynamic mechanisms to generate high-lift, leading-edge vortices. In normal free flight, dragonflies use counterstroking kinematics, with a leading-edge vortex (LEV) on the forewing downstroke, attached flow on the forewing upstroke, and attached flow on the hindwing throughout. Accelerating dragonflies switch to in-phase wing-beats with highly separated downstroke flows, with a single LEV attached across both the fore- and hindwings. We use smoke visualizations to distinguish between the three simplest local analytical solutions of the Navier-Stokes equations yielding flow separation resulting in a LEV. The LEV is an open U-shaped separation, continuous across the thorax, running parallel to the wing leading edge and inflecting at the tips to form wingtip vortices. Air spirals in to a free-slip critical point over the centreline as the LEV grows. Spanwise flow is not a dominant feature of the flow field--spanwise flows sometimes run from wingtip to centreline, or vice versa--depending on the degree of sideslip. LEV formation always coincides with rapid increases in angle of attack, and the smoke visualizations clearly show the formation of LEVs whenever a rapid increase in angle of attack occurs. There is no discrete starting vortex. Instead, a shear layer forms behind the trailing edge whenever the wing is at a non-zero angle of attack, and rolls up, under Kelvin-Helmholtz instability, into a series of transverse vortices with circulation of opposite sign to the circulation around the wing and LEV. The flow fields produced by dragonflies differ qualitatively from those published for mechanical models of dragonflies, fruitflies and hawkmoths, which preclude natural wing interactions. However, controlled parametric experiments show that, provided the Strouhal number is appropriate and the natural interaction between left and right wings can occur, even a simple plunging plate can reproduce the detailed features of the flow seen in dragonflies. In our models, and in dragonflies, it appears that stability of the LEV is achieved by a general mechanism whereby flapping kinematics are configured so that a LEV would be expected to form naturally over the wing and remain attached for the duration of the stroke. However, the actual formation and shedding of the LEV is controlled by wing angle of attack, which dragonflies can vary through both extremes, from zero up to a range that leads to immediate flow separation at any time during a wing stroke.

Tuning of Strouhal number for high propulsive efficiency accurately predicts how wingbeat frequency and stroke amplitude relate and scale with size and flight speed in birds.

The wing kinematics of birds vary systematically with body size, but we still, after several decades of research, lack a clear mechanistic understanding of the aerodynamic selection pressures that shape them. Swimming and flying animals have recently been shown to cruise at Strouhal numbers (St) corresponding to a regime of vortex growth and shedding in which the propulsive efficiency of flapping foils peaks (St approximately fA/U, where f is wingbeat frequency, U is cruising speed and A approximately bsin(theta/2) is stroke amplitude, in which b is wingspan and theta is stroke angle). We show that St is a simple and accurate predictor of wingbeat frequency in birds. The Strouhal numbers of cruising birds have converged on the lower end of the range 0.2 < St < 0.4 associated with high propulsive efficiency. Stroke angle scales as theta approximately 67b-0.24, so wingbeat frequency can be predicted as f approximately St.U/bsin(33.5b-0.24), with St0.21 and St0.25 for direct and intermittent fliers, respectively. This simple aerodynamic model predicts wingbeat frequency better than any other relationship proposed to date, explaining 90% of the observed variance in a sample of 60 bird species. Avian wing kinematics therefore appear to have been tuned by natural selection for high aerodynamic efficiency: physical and physiological constraints upon wing kinematics must be reconsidered in this light.

Flying and swimming animals cruise at a Strouhal number tuned for high power efficiency.

Dimensionless numbers are important in biomechanics because their constancy can imply dynamic similarity between systems, despite possible differences in medium or scale. A dimensionless parameter that describes the tail or wing kinematics of swimming and flying animals is the Strouhal number, St = fA/U, which divides stroke frequency (f) and amplitude (A) by forward speed (U). St is known to govern a well-defined series of vortex growth and shedding regimes for airfoils undergoing pitching and heaving motions. Propulsive efficiency is high over a narrow range of St and usually peaks within the interval 0.2 < St < 0.4 (refs 3-8). Because natural selection is likely to tune animals for high propulsive efficiency, we expect it to constrain the range of St that animals use. This seems to be true for dolphins, sharks and bony fish, which swim at 0.2 < St < 0.4. Here we show that birds, bats and insects also converge on the same narrow range of St, but only when cruising. Tuning cruise kinematics to optimize St therefore seems to be a general principle of oscillatory lift-based propulsion.

Dynamic flight stability in the desert locust Schistocerca gregaria.

Here we provide the first formal quantitative analysis of dynamic stability in a flying animal. By measuring the longitudinal static stability derivatives and mass distribution of desert locusts Schistocerca gregaria, we find that their static stability and static control responses are insufficient to provide asymptotic longitudinal dynamic stability unless they are sensitive to pitch attitude (measured with respect to an inertial or earth-fixed frame) as well as aerodynamic incidence (measured relative to the direction of flight). We find no evidence for a 'constant-lift reaction', previously supposed to keep lift production constant over a range of body angles, and show that such a reaction would be inconsequential because locusts can potentially correct for pitch disturbances within a single wingbeat. The static stability derivatives identify three natural longitudinal modes of motion: one stable subsidence mode, one unstable divergence mode, and one stable oscillatory mode (which is present with or without pitch attitude control). The latter is identified with the short period mode of aircraft, and shown to consist of rapid pitch oscillations with negligible changes in forward speed. The frequency of the short period mode (approx. 10 Hz) is only half the wingbeat frequency (approx. 22 Hz), so the mode would become coupled with the flapping cycle without adequate damping. Pitch rate damping is shown to be highly effective for this purpose - especially at the small scales associated with insect flight - and may be essential in stabilising locust flight. Although having a short period mode frequency close to the wingbeat frequency risks coupling, it is essential for control inputs made at the level of a single wingbeat to be effective. This is identified as a general constraint on flight control in flying animals.

HOW TO COMPENSATE FOR COSTLY SEXUALLY SELECTED TAILS: THE ORIGIN OF SEXUALLY DIMORPHIC WINGS IN LONG-TAILED BIRDS.

Recent work on birds suggests that certain morphological differences between the sexes may have evolved as an indirect consequence of sexual selection because they offset the cost of bearing extravagant ornaments used for fighting or mate attraction. For example, long-tailed male sunbirds and widowbirds also have longer wings than females, perhaps to compensate for the aerodynamic costs of tail elaboration. We used comparative data from 57 species to investigate whether this link between sexual dimorphism in wing and tail length is widespread among long-tailed birds. We found that within long-tailed families, variation in the extent of tail dimorphism was associated with corresponding variation in wing dimorphism. One nonfunctional explanation of this result is simply that the growth of wings and tails is controlled by a common developmental mechanism, such that long-tailed individuals inevitably grow long wings as well. However, this hypothesis cannot account for a second pattern in our data set: as predicted by aerodynamic theory, we found that, comparing across long-tailed families, sexual dimorphism in wing length varied with tail shape as well as with sex differences in tail length. Thus, wing dimorphism was generally greater in species with aerodynamically costly graduated tails than in birds with cheaper, streamer-shaped tails. This result was not caused by confounding phylogenetic effects, because it persisted when phylogeny was controlled for, using an independent comparisons method. Our findings therefore confirm that certain aspects of sexual dimorphism may sometimes have evolved through selection for traits that reduce the costs of elaborate sexually selected characters. We suggest that future work aimed at understanding sexual selection by investigating patterns of sexual dimorphism should attempt to differentiate between the direct and indirect consequences of sexual selection.