Do the enlarged hind legs of male thick-legged flower beetles contribute to take-off or mating?

The volume of the hind femora in the adult male flower beetle Oedemera nobilis is 38 times greater than in adult females. To determine what advantage limbs with swollen femora might provide, the behaviour of these insects was analysed with high-speed videography. First, because large hind legs are often associated with jumping and take-off, the performance of this behaviour by the two sexes was determined. Take-off was generated by a series of small-amplitude wing beats followed by larger ones, with the hind legs contributing little or no propulsion. The mean acceleration time to take-off was not significantly different in males (46.2 ms) and females (45.5 ms), but the mean take-off velocity of males was 10% higher than in females. Second, to determine if enlarged hind legs were critical in specifically male behaviour, interactions between males and females, and between males were videoed. The male mounted a female and then encircled her abdomen between the enlarged femora and tibiae of both his hind legs. The joint between these leg parts acted like a mole wrench (vice grip) so that when the tibia was fully flexed, a triangular space of 0.3 mm2 remained, in which a female abdomen (cross-sectional area 0.9 mm2) could be compressed and restrained firmly without inflicting damage. The flexor tibiae muscle in a male hind femur was 5.9 times larger than the extensor. In interactions between males, attempts to achieve a similar entrapment were frequently thwarted by the pursued male extending his hind legs vertically.

Effectiveness and efficiency of two distinct mechanisms for take-off in a derbid planthopper insect.

Analysis of the kinematics of take-off in the planthopper Proutista moesta (Hemiptera, Fulgoroidea, family Derbidae) from high-speed videos showed that these insects used two distinct mechanisms involving different appendages. The first was a fast take-off (55.7% of 106 take-offs by 11 insects) propelled by a synchronised movement of the two hind legs and without participation of the wings. The body was accelerated in 1 ms or less to a mean take-off velocity of 1.7 m s-1 while experiencing average forces of more than 150 times gravity. The power required from the leg muscles implicated a power-amplification mechanism. Such take-offs propelled the insect along its trajectory a mean distance of 7.9 mm in the first 5 ms after take-off. The second and slower take-off mechanism (44.3% of take-offs) was powered by beating movements of the wings alone, with no discernible contribution from the hind legs. The resulting mean acceleration time was 16 times slower at 17.3 ms, the mean final velocity was six times lower at 0.27 m s-1, the g forces experienced were 80 times lower and the distance moved in 5 ms after take-off was 7 times shorter. The power requirements could be readily met by direct muscle contraction. The results suggest a testable hypothesis that the two mechanisms serve distinct behavioural actions: the fast take-offs could enable escape from predators and the slow take-offs that exert much lower ground reaction forces could enable take-off from more flexible substrates while also displacing the insect in a slower and more controllable trajectory.

Take-off mechanisms in parasitoid wasps.

High-speed video analyses of the natural behaviour of parasitoid wasps revealed three strategies used to launch the insects into the air. Which strategy is the most energy efficient? In Pteromalus puparum, 92% of take-offs by were propelled entirely by movements of the middle and hind legs, which were depressed at their coxo-trochanteral and extended at their femoro-tibial joints. The front legs left the ground first, followed by the hind legs, so that the middle legs provided the final propulsion. Second, in other species of a similar mass, Cotesia glomerata and Leptopilina boulardi, all take-offs were propelled by a mean of 2.8 and 3.8 wingbeats, respectively, with little or no contribution from the legs. The first strategy resulted in take-off times that were four times shorter (5 versus 22.8 ms) and take-off velocities that were four times faster (0.8 versus 0.2 m s-1). Calculations from the kinematics indicate that propulsion by the legs was the most energy-efficient strategy, because more energy is put into propulsion of the body, whereas in take-off propelled by repetitive wing movements energy is lost to generating these movements and moving the air. In heavier species such as Netelia testacea and Amblyteles armatorius, take-off was propelled by the combined movements of the middle and hind legs and wingbeats. In A. armatorius, this resulted in the longest mean take-off time of 33.8 ms but an intermediate take-off velocity of 0.4 m s-1 In all three strategies the performance could be explained without invoking energy storage and power amplification mechanisms.

Development and deposition of resilin in energy stores for locust jumping.

Locusts jump by using a catapult mechanism in which energy produced by slow contractions of the extensor tibiae muscles of the hind legs is stored in distortions of the exoskeleton, most notably (1) the two semi-lunar processes at each knee joint and (2) the tendons of the extensor muscles themselves. The energy is then suddenly released from these stores to power the rapid, propulsive movements of the hind legs. The reliance on the mechanical storage of energy is likely to impact on jumping because growth occurs by a series of five moults, at each of which the exoskeleton is replaced by a new one. All developmental stages (instars) nevertheless jump as a means of forward locomotion, or as an escape movement. Here, I show that in each instar, resilin is added to the semi-lunar processes and to the core of the extensor tendons so that their thickness increases. As the next moult approaches, a new exoskeleton forms within the old one, with resilin already present in the new semi-lunar processes. The old exoskeleton, the tendons and their resilin are discarded at moulting. The resilin of the semi-lunar processes and tendons of the new instar is initially thin, but a similar pattern of deposition results in an increase of their thickness. In adults, resilin continues to be deposited so that at 4 weeks old the thickness in the semi-lunar processes has increased fourfold. These changes in the energy stores accompany changes in jumping ability and performance during each moulting cycle.

Increased muscular volume and cuticular specialisations enhance jump velocity in solitarious compared with gregarious desert locusts, Schistocerca gregaria.

The desert locust, Schistocerca gregaria, shows a strong phenotypic plasticity. It can develop, depending upon population density, into either a solitarious or gregarious phase that differs in many aspects of behaviour, physiology and morphology. Prominent amongst these differences is that solitarious locusts have proportionately longer hind femora than gregarious locusts. The hind femora contain the muscles and energy-storing cuticular structures that propel powerful jumps using a catapult-like mechanism. We show that solitarious locusts jump on average 23% faster and 27% further than gregarious locusts, and attribute this improved performance to three sources: first, a 17.5% increase in the relative volume of their hind femur, and hence muscle volume; second, a 24.3% decrease in the stiffness of the energy-storing semi-lunar processes of the distal femur; and third, a 4.5% decrease in the stiffness of the tendon of the extensor tibiae muscle. These differences mean that solitarious locusts can generate more power and store more energy in preparation for a jump than can gregarious locusts. This improved performance comes at a cost: solitarious locusts expend nearly twice the energy of gregarious locusts during a single jump and the muscular co-contraction that energises the cuticular springs takes twice as long. There is thus a trade-off between achieving maximum jump velocity in the solitarious phase against the ability to engage jumping rapidly and repeatedly in the gregarious phase.

Jumping mechanisms in adult caddis flies (Insecta, Trichoptera).

To understand the jumping mechanisms and strategies of adult caddis flies, leg morphology and movements were analysed in three species with mean masses of 3.9 to 38 mg. Two distinct jumping strategies were found. First (67% of 90 jumps), take-off was propelled solely by the middle and hind legs while the wings remained closed. Second (33% of jumps), the same leg movements were combined with wing movements before take-off. The hind legs were 70% and the middle legs were 50% longer than the front legs and represented 105% and 88%, respectively, of body length. Both hind and middle trochantera were depressed together, approximately 15 ms before take-off. The front legs apparently did not contribute to thrust in either strategy and were the first to be lifted from the ground. The hind legs were the next to lose contact, so that the middle legs alone provided the final thrust before take-off. Jumping performance did not differ significantly in the two jumping strategies or between species, in acceleration times (range of means for the three species 14.5-15.4 ms), take-off velocities (range 0.7-1 m s(-1)) and trajectory angles. A significant difference in jumps propelled only by the legs was the lower angle (9.3 ± 1.9 deg) of the body relative to the horizontal at take-off compared with jumps involving wing movements (35.3 ± 2.5 deg). Calculations from the kinematics indicated that jumps were produced by direct muscle contractions and did not require power amplification or energy storage.

Jumping mechanisms and strategies in moths (Lepidoptera).

To test whether jumping launches moths into the air, take-off by 58 species, ranging in mass from 0.1 to 220 mg, was captured in videos at 1000 frames s(-1). Three strategies for jumping were identified. First, rapid movements of both middle and hind legs provided propulsion while the wings remained closed. Second, middle and hind legs again provided propulsion but the wings now opened and flapped after take-off. Third, wing and leg movements both began before take-off and led to an earlier transition to powered flight. The middle and hind legs were of similar lengths and were between 10 and 130% longer than the front legs. The rapid depression of the trochantera and extension of the middle tibiae began some 3 ms before similar movements of the hind legs, but their tarsi lost contact with the ground before take-off. Acceleration times ranged from 10 ms in the lightest moths to 25 ms in the heaviest ones. Peak take-off velocities varied from 0.6 to 0.9 m s(-1) in all moths, with the fastest jump achieving a velocity of 1.2 m s(-1). The energy required to generate the fastest jumps was 1.1 µJ in lighter moths but rose to 62.1 µJ in heavier ones. Mean accelerations ranged from 26 to 90 m s(-2) and a maximum force of 9 G: was experienced. The highest power output was within the capability of normal muscle so that jumps were powered by direct contractions of muscles without catapult mechanisms or energy storage.

Jumping mechanisms in flatid planthoppers (Hemiptera, Flatidae).

The jumping performance of three species of hemipterans from Australia and Europe belonging to the family Flatidae was analysed from images captured at a rate of 5000 s(-1). The shape of a flatid was dominated by large triangular or wedge-shaped front wings, which, when folded, covered and extended above and behind the body to give a laterally compressed and possibly streamlined appearance. The body lengths of the three species of adults ranged from 7 to 9 mm and their mass from 8 to 19 mg. The propulsive hind legs were 30% longer than the front legs but only 36-54% of the body length. Jumps with the fastest take-off velocities of 2.8-3.2 m s(-1) had acceleration times of 1.4-1.8 ms. During such jumps, adults experienced an acceleration of 174-200 G: . These jumps required an energy expenditure of 76-225 µJ, a power output of 13-60 mW and exerted a force of 9-37 mN. The required power output per mass of jumping muscle in adults ranged from 24,000 to 27,000 W kg(-1) muscle, 100 times greater than the maximum active contractile limit of normal muscle. The free-living nymphs were also proficient jumpers, reaching take-off velocities of 2.2 m s(-1). To achieve such a jumping performance requires a power amplification mechanism. The energy store for such a mechanism was identified as the internal skeleton linking a hind coxa to the articulation of a hind wing. These pleural arches fluoresced bright blue when illuminated with UV light, indicating the presence of the elastic protein resilin. The energy generated by the prolonged contractions of the trochanteral depressor muscles was stored in distortions of these structures, and the rapid elastic recoil of these muscles powered the synchronous propulsive movements of the hind legs.

Jumping mechanisms in dictyopharid planthoppers (Hemiptera, Dicytyopharidae).

The jumping performance of four species of hemipterans belonging to the family Dictyopharidae, from Europe, South Africa and Australia, were analysed from high-speed images. The body shape in all was characterised by an elongated and tapering head that gave a streamlined appearance. The body size ranged from 6 to 9 mm in length and from 6 to 23 mg in mass. The hind legs were 80-90% of body length and 30-50% longer than the front legs, except in one species in which the front legs were particularly large so that all legs were of similar length. Jumping was propelled by rapid and simultaneous depression of the trochantera of both hind legs, powered by large muscles in the thorax, and was accompanied by extension of the tibiae. In the best jumps, defined as those with the fastest take-off velocity, Engela minuta accelerated in 1.2 ms to a take-off velocity of 5.8 m s(-1), which is the fastest achieved by any insect described to date. During such a jump, E. minuta experienced an acceleration of 4830 m s(-2) or 490 g, while other species in the same family experienced 225-375 g. The best jumps in all species required an energy expenditure of 76-225 µJ, a power output of 12-80 mW and exerted a force of 12-29 mN. The required power output per mass of jumping muscle ranged from 28,000 to 140,200 W kg(-1) muscle and thus greatly exceeded the maximum active contractile limit of normal muscle. To achieve such a jumping performance, these insects must be using a power amplification mechanism in a catapult-like action. It is suggested that their streamlined body shape improves jumping performance by reducing drag, which, for a small insect, can substantially affect forward momentum.

Jumping mechanisms in lacewings (Neuroptera, Chrysopidae and Hemerobiidae).

Lacewings launch themselves into the air by simultaneous propulsive movements of the middle and hind legs as revealed in video images captured at a rate of 1000 s(-1). These movements were powered largely by thoracic trochanteral depressor muscles but did not start from a particular preset position of these legs. Ridges on the lateral sides of the meso- and metathorax fluoresced bright blue when illuminated with ultraviolet light, suggesting the presence of the elastic protein resilin. The middle and hind legs were longer than the front legs but their femora and tibiae were narrow tubes of similar diameter. Jumps were of two types. First, those in which the body was oriented almost parallel to the ground (-7±8 deg in green lacewings, 13.7±7 deg in brown lacewings) at take-off and remained stable once animals were airborne. The wings did not move until 5 ms after take-off when flapping flight ensued. Second, were jumps in which the head pointed downwards at take-off (green lacewings, -37±3 deg; brown lacewings, -35±4 deg) and the body rotated in the pitch plane once airborne without the wings opening. The larger green lacewings (mass 9 mg, body length 10.3 mm) took 15 ms and the smaller brown lacewings (3.6 mg and 5.3 mm) 9 ms to accelerate the body to mean take-off velocities of 0.6 and 0.5 m s(-1). During their fastest jumps green and brown lacewings experienced accelerations of 5.5 or 6.3 G: , respectively. They required an energy expenditure of 5.6 or 0.7 µJ, a power output of 0.3 or 0.1 mW and exerted a force of 0.6 or 0.2 mN. The required power was well within the maximum active contractile limit of normal muscle, so that jumping could be produced by direct muscle contractions without a power amplification mechanism or an energy store.

Jumping from the surface of water by the long-legged fly Hydrophorus (Diptera, Dolichopodidae).

The fly Hydrophorus alboflorens (4 mm long, 4.7 mg mass) moves around upon and jumps from water without its tarsi penetrating the surface. All six tarsi have a surface area of 1.3 mm(-2) in contact with the water, but they did not dimple its surface when standing. Jumping was propelled by depression of the trochantera of both hind and middle legs, which are 40% longer than the front legs and 170% longer than the body. As these four legs progressively propelled the insect to take-off, they each created dimples on the water surface that expanded in depth and area. No dimples were associated with the front legs, which were not moved in a consistent sequence. The wings opened while the legs were moving and then flapped at a frequency of 148 Hz. The body was accelerated in a mean time of 21 ms to a mean take-off velocity of 0.7 m s(-1). The best jumps reached velocities of 1.6 m s(-1), and required an energy output of 7 µJ and a power output of 0.6 mW, with the fly experiencing a force of 140 g. The required power output indicates that direct muscle contractions could propel the jump without the need for elaborate mechanisms for energy storage. Take-off trajectories were steep, with a mean of 87 deg to the horizontal. Take-off velocity fell if a propulsive tarsus penetrated the surface of the water. If more tarsi became submerged, take-off was not successful. A second strategy for take-off was powered only by the wings and was associated with slower (1 deg ms(-1) compared with 10 deg ms(-1) when jumping) and less extensive movements of the propulsive joints of the middle and hind legs. No dimples were then created on the surface of the water. When jumping was combined with wing flapping, the acceleration time to take-off was reduced by 84% and the take-off velocity was increased by 168%. Jumping can potentially therefore enhance survival when threatened by a potential predator.

Jumping mechanisms in gum treehopper insects (Hemiptera, Eurymelinae).

Jumping in a species of Australian gum treehopper was analysed from high-speed images. Pauroeurymela amplicincta adults and nymphs lived together in groups that were tended by ants, but only adults jumped. The winged adults with a body mass of 23 mg and a body length of 7 mm had some morphological characteristics intermediate between those of their close relatives the leafhoppers (Cicadellidae) and the treehoppers (Membracidae). They, like leafhoppers, lacked the prominent prothoracic helmets of membracid treehoppers, and their large hind coxae were linked by press studs (poppers), that are present in leafhoppers but not treehoppers. The hindlegs were only 30-40% longer than the other legs and 67% of body length. They are thus of similar proportion to the hindlegs of treehoppers but much shorter than those of most leafhoppers. Jumping was propelled by the hindlegs, which moved in the same plane as each other beneath and almost parallel to the longitudinal axis of the body. A jump was preceded by full levation of the coxo-trochanteral joints of the hindlegs. In its best jumps, the rapid depression of these joints then accelerated the insect in 1.4 ms to a take-off velocity of 3.8 m s(-1) so that it experienced a force of almost 280 g. In 22% of jumps, the wings opened before take-off but did not flap until the gum treehopper was airborne, when the body rotated little in any plane. The energy expended was 170 µJ, the power output was 122 mW and the force exerted was 64 mN. Such jumps are predicted to propel the insect forwards 1450 mm (200 times body length) and to a height of 430 mm if there is no effect of wind resistance. The power output per mass of jumping muscle far exceeded the maximum active contractile limit of muscle and indicates that a catapult-like action must be used. This eurymelid therefore out-performs both leafhoppers and treehoppers in i ts faster acceleration and in its higher take-off velocity.

Locusts use a composite of resilin and hard cuticle as an energy store for jumping and kicking.

Locusts jump and kick by using a catapult mechanism in which energy is first stored and then rapidly released to extend the large hind legs. The power is produced by a slow contraction of large muscles in the hind femora that bend paired semi-lunar processes in the distal part of each femur and store half the energy needed for a kick. We now show that these energy storage devices are composites of hard cuticle and the rubber-like protein resilin. The inside surface of a semi-lunar process consists of a layer of resilin, particularly thick along an inwardly pointing ridge and tightly bonded to the external, black cuticle. From the outside, resilin is visible only as a distal and ventral triangular area that tapers proximally. High-speed imaging showed that the semi-lunar processes were bent in all three dimensions during the prolonged muscular contractions that precede a kick. To reproduce these bending movements, the extensor tibiae muscle was stimulated electrically in a pattern that mimicked the normal sequence of its fast motor spikes recorded in natural kicking. Externally visible resilin was compressed and wrinkled as a semi-lunar process was bent. It then sprung back to restore the semi-lunar process rapidly to its original natural shape. Each of the five nymphal stages jumped and kicked and had a similar distribution of resilin in their semi-lunar processes as adults; the resilin was shed with the cuticle at each moult. It is suggested that composite storage devices that combine the elastic properties of resilin with the stiffness of hard cuticle allow energy to be stored by bending hard cuticle over only a small distance and without fracturing. In this way all the stored energy is returned and the natural shape of the femur is restored rapidly so that a jump or kick can be repeated.

A cockroach that jumps.

We report on a newly discovered cockroach (Saltoblattella montistabularis) from South Africa, which jumps and therefore differs from all other extant cockroaches that have a scuttling locomotion. In its natural shrubland habitat, jumping and hopping accounted for 71 per cent of locomotory activity. Jumps are powered by rapid and synchronous extension of the hind legs that are twice the length of the other legs and make up 10 per cent of the body weight. In high-speed images of the best jumps the body was accelerated in 10 ms to a take-off velocity of 2.1 m s(-1) so that the cockroach experienced the equivalent of 23 times gravity while leaping a forward distance of 48 times its body length. Such jumps required 38 µJ of energy, a power output of 3.4 mW and exerted a ground reaction force through both hind legs of 4 mN. The large hind legs have grooved femora into which the tibiae engage fully in advance of a jump, and have resilin, an elastic protein, at the femoro-tibial joint. The extensor tibiae muscles contracted for 224 ms before the hind legs moved, indicating that energy must be stored and then released suddenly in a catapult action to propel a jump. Overall, the jumping mechanisms and anatomical features show remarkable convergence with those of grasshoppers with whom they share their habitat and which they rival in jumping performance.

Jumping mechanisms and performance of snow fleas (Mecoptera, Boreidae).

Flightless snow fleas (snow scorpion flies, Mecoptera, Boreidae) live as adults during northern hemisphere winters, often jumping and walking on the surface of snow. Their jumping mechanisms and performance were analysed with high speed imaging. Jumps were propelled by simultaneous movements of both the middle and hind pairs of legs, as judged by the 0.2 ms resolution afforded by image rates of 5000 frames s(-1). The middle legs of males represent 140% and the hindlegs 187% of the body length (3.4 mm), and the ratio of leg lengths is 1:1.3:1.7 (front:middle:hind). In preparation for a jump the middle legs and hindlegs were rotated forwards at their coxal joints with the fused mesothorax and metathorax. The first propulsive movement of a jump was the rotation of the trochantera about the coxae, powered by large depressor muscles within the thorax. The acceleration time was 6.6 ms. The fastest jump by a male had a take-off velocity of 1 m s(-1), which required 1.1 µJ of energy and a power output of 0.18 mW, and exerted a force about 16 times its body weight. Jump distances of about 100 mm were unaffected by temperature. This, and the power per mass of muscle requirement of 740 W kg(-1), suggests that a catapult mechanism is used. The elastic protein resilin was revealed in four pads at the articulation of the wing hinge with the dorsal head of the pleural ridge of each middle leg and hindleg. By contrast, fleas, which use just their hindlegs for jumping, have only two pads of resilin. This, therefore, provides a functional reference point for considerations about the phylogenetic relationships between snow fleas and true fleas.

Biomechanics of jumping in the flea.

It has long been established that fleas jump by storing and releasing energy in a cuticular spring, but it is not known how forces from that spring are transmitted to the ground. One hypothesis is that the recoil of the spring pushes the trochanter onto the ground, thereby generating the jump. A second hypothesis is that the recoil of the spring acts through a lever system to push the tibia and tarsus onto the ground. To decide which of these two hypotheses is correct, we built a kinetic model to simulate the different possible velocities and accelerations produced by each proposed process and compared those simulations with the kinematics measured from high-speed images of natural jumping. The in vivo velocity and acceleration kinematics are consistent with the model that directs ground forces through the tibia and tarsus. Moreover, in some natural jumps there was no contact between the trochanter and the ground. There were also no observable differences between the kinematics of jumps that began with the trochanter on the ground and jumps that did not. Scanning electron microscopy showed that the tibia and tarsus have spines appropriate for applying forces to the ground, whereas no such structures were seen on the trochanter. Based on these observations, we discount the hypothesis that fleas use their trochantera to apply forces to the ground and conclude that fleas jump by applying forces to the ground through the end of the tibiae.

Elucidating the complex organization of neural micro-domains in the locust Schistocerca gregaria using dMRI.

To understand brain function it is necessary to characterize both the underlying structural connectivity between neurons and the physiological integrity of these connections. Previous research exploring insect brain connectivity has typically used electron microscopy techniques, but this methodology cannot be applied to living animals and so cannot be used to understand dynamic physiological processes. The relatively large brain of the desert locust, Schistercera gregaria (Forksȧl) is ideal for exploring a novel methodology; micro diffusion magnetic resonance imaging (micro-dMRI) for the characterization of neuronal connectivity in an insect brain. The diffusion-weighted imaging (DWI) data were acquired on a preclinical system using a customised multi-shell diffusion MRI scheme optimized to image the locust brain. Endogenous imaging contrasts from the averaged DWIs and Diffusion Kurtosis Imaging (DKI) scheme were applied to classify various anatomical features and diffusion patterns in neuropils, respectively. The application of micro-dMRI modelling to the locust brain provides a novel means of identifying anatomical regions and inferring connectivity of large tracts in an insect brain. Furthermore, quantitative imaging indices derived from the kurtosis model that include fractional anisotropy (FA), mean diffusivity (MD) and kurtosis anisotropy (KA) can be extracted. These metrics could, in future, be used to quantify longitudinal structural changes in the nervous system of the locust brain that occur due to environmental stressors or ageing.

Spatiotemporal receptive field properties of a looming-sensitive neuron in solitarious and gregarious phases of the desert locust.

Desert locusts (Schistocerca gregaria) can transform reversibly between the swarming gregarious phase and a solitarious phase, which avoids other locusts. This transformation entails dramatic changes in morphology, physiology, and behavior. We have used the lobula giant movement detector (LGMD) and its postsynaptic target, the descending contralateral movement detector (DCMD), which are visual interneurons that detect looming objects, to analyze how differences in the visual ecology of the two phases are served by altered neuronal function. Solitarious locusts had larger eyes and a greater degree of binocular overlap than those of gregarious locusts. The receptive field to looming stimuli had a large central region of nearly equal response spanning 120 degrees x 60 degrees in both phases. The DCMDs of gregarious locusts responded more strongly than solitarious locusts and had a small caudolateral focus of even further sensitivity. More peripherally, the response was reduced in both phases, particularly ventrally, with gregarious locusts showing greater proportional decrease. Gregarious locusts showed less habituation to repeated looming stimuli along the eye equator than did solitarious locusts. By contrast, in other parts of the receptive field the degree of habituation was similar in both phases. The receptive field organization to looming stimuli contrasts strongly with the receptive field organization of the same neurons to nonlooming local-motion stimuli, which show much more pronounced regional variation. The DCMDs of both gregarious and solitarious locusts are able to detect approaching objects from across a wide expanse of visual space, but phase-specific changes in the spatiotemporal receptive field are linked to lifestyle changes.

A single muscle moves a crustacean limb joint rhythmically by acting against a spring containing resilin.

BACKGROUND: The beating or fanning movements of three pairs of maxilliped flagella in crabs and crayfish modify exhalent gill currents while drawing water over chemoreceptors on the head. They play an integral part both in signalling by distributing urine odours, and in active chemosensation. RESULTS: The rhythmical maxilliped movements start with maxilliped 3 followed after a delay of 15 to 20 ms in shore crabs by maxilliped 2 and then maxilliped 1, at a frequency of 18 to 20 Hz in crabs and 10 to 13 Hz in signal crayfish. The contraction of a single abductor muscle controls the power stroke (abduction) of each flagellum, which is accompanied by flaring of feather-like setae which increase its surface area. No muscle can bring about the return stroke (adduction). Release of an isolated flagellum from an imposed abduction is followed by a rapid recoil to its resting adducted position. The relationship between the extent of abduction and the angular velocity of the return stroke indicates the operation of a spring. Blue fluorescence under UV light, and its dependence on the pH of the bathing medium, indicates that resilin is present at the joint between an exopodite and flagellum, at the annuli of a flagellum and at the base of the setae. CONCLUSION: Resilin is progressively bent as a flagellum is abducted and resumes its natural shape when the joint recoils. Other distortions of the exopodites may also contribute to this spring-like action. The joint is therefore controlled by a single abductor muscle operating against a spring in which the elastic properties of resilin play a key role.

Jumping performance of planthoppers (Hemiptera, Issidae).

The structure of the hind limbs and the kinematics of their movements that propel jumping in planthopper insects (Hemiptera, Auchenorrhyncha, Fulgoroidea, Issidae) were analysed. The propulsion for a jump was delivered by rapid movements of the hind legs that both move in the same plane beneath the body and parallel to its longitudinal axis, as revealed in high-speed sequences of images captured at rates up to 7500 images s(-1). The first and key movement was the depression of both trochantera about their coxae, powered by large depressor muscles in the thorax, accompanied by rapid extension of the tibiae about their femora. The initial movements of the two trochantera of the hind legs were synchronised to within 0.03 ms. The hind legs are only 20% longer than the front and middle legs, represent 65% of the body length, and have a ratio of 1.8 relative to the cube root of the body mass. The two hind coxae have a different structure to those in frog- and leafhoppers. They are fused at the mid-line, covered ventrally by transparent cuticle, and each is fixed laterally to a part of the internal skeleton called the pleural arch that extends to the articulation of a hind wing. A small and pointed, ventral coxal protrusion covered in microtrichia engages with a raised, smooth, white patch on a dorsal femur when a hind leg is levated (cocked) in preparation for a jump. In the best jumps by a male Issus, the body was accelerated in 0.8 ms to a take-off velocity of 5.5 m s(-1), was subjected to a force of 719 g and was displaced a horizontal distance of 1.1 m. This performance required an energy output of 303 microJ, a power output of 388 mW and exerted a force of 141 mN, or more than 700 times its body mass. This performance implies that a catapult mechanism must be used, and that Issus ranks alongside the froghopper Philaenus as one of the best insect jumpers.