Mechanical performance of spider orb webs is tuned for high-speed prey.

Spiders in the Orbiculariae spin orb webs that dissipate the mechanical energy of their flying prey, bringing the insects to rest and retaining them long enough for the spider to attack and subdue their meals. Small prey are easily stopped by webs but provide little energetic gain. While larger prey offer substantial nourishment, they are also challenging to capture and can damage the web if they escape. We therefore hypothesized that spider orb webs exhibit properties that improve their probability of stopping larger insects while minimizing damage when the mechanical energy of those prey exceeds the web's capacity. Large insects are typically both heavier and faster flying than smaller prey, but speed plays a disproportionate role in determining total kinetic energy, so we predicted that orb webs may dissipate energy more effectively under faster impacts, independent of kinetic energy per se. We used high-speed video to visualize the impact of wooden pellets fired into orb webs to simulate prey strikes and tested how capture probability varied as a function of pellet size and speed. Capture probability was virtually nil above speeds of ~3 m s(-1). However, successful captures do not directly measure the maximum possible energy dissipation by orb webs because these events include lower-energy impacts that may not significantly challenge orb web performance. Therefore, we also compared the total kinetic energy removed from projectiles that escaped orb webs by breaking through the silk, asking whether more energy was removed at faster speeds. Over a range of speeds relevant to insect flight, the amount of energy absorbed by orb webs increases with the speed of prey (i.e. the rates at which webs are stretched). Orb webs therefore respond to faster - and hence higher kinetic energy - prey with better performance, suggesting adaptation to capture larger and faster flying insect prey. This speed-dependent toughness of a complex structure suggests the utility of the intrinsic toughness of spider silk and/or features of the macro-design of webs for high-velocity industrial or military applications, such as ballistic energy absorption.

Hydrodynamic pumping by serial gill arrays in the mayfly nymph Centroptilum triangulifer.

Aquatic nymphs of the mayfly Centroptilum triangulifer produce ventilatory flow using a serial array of seven abdominal gill pairs that operates across a Reynolds numbers (Re) range from 2 to 22 during ontogeny. Net flow in small animals is directed ventrally and essentially parallel to the stroke plane (i.e. rowing), but net flow in large animals is directed dorsally and essentially transverse to the stroke plane (i.e. flapping). Detailed flow measurements based on Particle Image Velocimetry (PIV) ensemble-correlation analysis revealed that the phasing of the gills produces a time-dependent array of vortices associated with a net ventilatory current, a fluid kinematic pattern, here termed a 'phased vortex pump'. Absolute size of vortices does not change with increasing animal size or Re, and thus the vortex radius (R(v)) decreases relative to inter-gill distance (L(is)) during mayfly growth. Given that effective flapping in appendage-array animals requires organized flow between adjacent appendages, we hypothesize that rowing should be favored when L(is)/R(v)<1 and flapping should be favored when L(is)/R(v)>1. Significantly, the rowing-to-flapping transition in Centroptilum occurs at Re∼5, when the mean dynamic inter-gill distance equals the vortex radius. This result suggests that the Re-based rowing-flapping demarcation observed in appendage-array aquatic organisms may be determined by the relative size of the propulsive mechanism and its self-generated vortices.

Spider orb webs rely on radial threads to absorb prey kinetic energy.

The kinetic energy of flying insect prey is a formidable challenge for orb-weaving spiders. These spiders construct two-dimensional, round webs from a combination of stiff, strong radial silk and highly elastic, glue-coated capture spirals. Orb webs must first stop the flight of insect prey and then retain those insects long enough to be subdued by the spiders. Consequently, spider silks rank among the toughest known biomaterials. The large number of silk threads composing a web suggests that aerodynamic dissipation may also play an important role in stopping prey. Here, we quantify energy dissipation in orb webs spun by diverse species of spiders using data derived from high-speed videos of web deformation under prey impact. By integrating video data with material testing of silks, we compare the relative contributions of radial silk, the capture spiral and aerodynamic dissipation. Radial silk dominated energy absorption in all webs, with the potential to account for approximately 100 per cent of the work of stopping prey in larger webs. The most generous estimates for the roles of capture spirals and aerodynamic dissipation show that they rarely contribute more than 30 per cent and 10 per cent of the total work of stopping prey, respectively, and then only for smaller orb webs. The reliance of spider orb webs upon internal energy absorption by radial threads for prey capture suggests that the material properties of the capture spirals are largely unconstrained by the selective pressures of stopping prey and can instead evolve freely in response to alternative functional constraints such as adhering to prey.

Supercontraction forces in spider dragline silk depend on hydration rate.

Spider dragline silk is a model biological polymer for biomimetic research due to its many desirable and unusual properties. 'Supercontraction' describes the dramatic shrinking of dragline silk fibers when wetted. In restrained silk fibers, supercontraction generates substantial stresses of 40-50 MPa above a critical humidity of approximately 70% relative humidity (RH). This stress may maintain tension in webs under the weight of rain or dew and could be used in industry for robotics, sensor technology, and other applications. Our own findings indicate that supercontraction can generate stress over a much broader range than previously reported, from 10 to 140 MPa. Here we show that this variation in supercontraction stress depends upon the rate at which the environment reaches the critical level of humidity causing supercontraction. Slow humidity increase, over several minutes, leads to relatively low supercontraction stress, while fast humidity increase, over a few seconds, typically results in higher supercontraction stress. Slowly supercontracted fibers take up less water and differ in thermostability from rapidly supercontracted fibers, as shown by thermogravimetric analysis. This suggests that spider silk achieves different molecular configurations depending upon the speed at which supercontraction occurs. Ultimately, rate-dependent supercontraction may provide a mechanism to tailor the properties of silk or biomimetic fibers for various applications.

Mechanics of cuticular elastic energy storage in leg joints lacking extensor muscles in arachnids.

Certain leg joints in arachnids lack extensor muscles and have elastically deformable transarticular sclerites spanning their arthrodial membranes, an arrangement consistent with a model in which flexor muscles load transarticular sclerites during flexion and energy from elastic recoil is used for extension. This study quantifies the potential contribution of elastic recoil to extension torque at joints of the fourth leg of representative arachnids. Extension torques of isolated joints with and without transarticular sclerites were measured as the joint was rotated through angles and at angular velocities comparable with those used by walking animals. The procedure was repeated with the joint subjected to different internal fluid pressures in order to assess the potential role of hydraulically induced extension. The efficiency of elastic energy storage (resilience) in the absence of internal fluid pressure was 70-90% for joints with well-developed transarticular sclerites, and the magnitude of torque was similar to those produced by different joint extension mechanisms in other arthropods. Increased internal fluid pressure acted synergistically with transarticular sclerites in some joints but had little or no effect in others. Joints that lacked both extensor muscles and transarticular sclerites appeared to be specialized for hydraulic extension, and joints operated by antagonistic muscles lacked apparent specializations for either elastic or hydraulic extension. It is well known that elastic energy storage is a significant contributor to propulsion in running vertebrates and certain arthropods, where elastic elements are loaded as the center of mass falls during one phase of the locomotor cycle. However, transarticular sclerites are apparently loaded by contraction of flexor muscles when the leg is not in contact with the substratum. Hence the mechanism of a transarticular sclerite is more similar to the flight and jumping mechanisms of other arthropods than to running vertebrates. The evolutionary significance and potential mechanical advantages of the transarticular elastic mechanism are discussed.