To attach reliably on various inclined rough surfaces, many insects have evolved both claws and adhesive pads on their feet. However, the interaction between these organs still remains unclear. Here we designed an artificial attachment device, which mimics the structure and function of claws and adhesive pads, and tested it on stiff spheres of different dimensions. The results show that the attachment forces of claws decrease with an increase of the sphere radius. The forces may become very strong, when the sphere radius is smaller or comparable to the claw radius, because of the frictional self-lock. On the other hand, adhesive pads generate considerable adhesion on large sphere diameter due to large contact areas. The synergy effect between the claws and adhesive pads leads to much stronger attachment forces, if compared to the action of claw or adhesive pads independently (or even to the sum of both). The results carried out by our insect-inspired artificial attachment device clearly demonstrate why biological evolution employed two attachment organs working in concert. The results may greatly inspire the robot design, to obtain reliable attachment forces on various substrates.

SUMMARYMeasuring the interaction between each foot of an animal and the substrate is one of the most effective ways to understand the dynamics of legged locomotion. Here, a new facility -the force-measuring array (FMA) -was developed and applied to measure 3D reaction forces of geckos on different slope surfaces. The FMA consists of 16 3D sensors with resolution to the mN level. At the same time the locomotion behaviour of geckos freely moving on the FMA was recorded by high speed camera. The reaction forces acting on the gecko's individual feet measured by the FMA and correlated with locomotion behaviour provided enough information to reveal the mechanical and dynamic secrets of gecko locomotion. Moreover, dynamic forces were also measured by a force platform and correlated with locomotion behaviour. The difference between the forces measured by the two methods is discussed. From the results we conclude that FMA is the best way to obtain true reaction forces acting on the gecko's individual feet.

The excellent locomotion ability of geckos on various rough and/or inclined substrates has attracted scientists' attention for centuries. However, the moving ability of gecko-mimicking robots on various inclined surfaces still lags far behind that of geckos, mainly because our understanding of how geckos govern their locomotion is still very poor. To reveal the fundamental mechanism of gecko locomotion and also to facilitate the design of gecko-mimicking robots, we have measured the reaction forces (RFs) acting on each individual foot of moving geckos on inverted, vertical and horizontal substrates (i.e. ceiling, wall and floor), have associated the RFs with locomotion behaviors by using high-speed camera, and have presented the relationships of the force components with patterns of reaction forces (PRFs). Geckos generate different PRF on ceiling, wall and floor, that is, the PRF is determined by the angles between the direction of gravity and the substrate on which geckos move. On the ceiling, geckos produce reversed shear forces acting on the front and hind feet, which pull away from the body in both lateral and fore-aft directions. They use a very large supporting angle from 21° to 24° to reduce the forces acting on their legs and feet. On the floor, geckos lift their bodies using a supporting angle from 76° to 78°, which not only decreases the RFs but also improves their locomotion ability. On the wall, geckos generate a reliable self-locking attachment by using a supporting angle of 14.8°, which is only about half of the critical angle of detachment.

Remarkable progress has been made characterizing one of nature's most integrated, hierarchical structures––the fibrillar adhesive system of geckos. Nonetheless, we lack an understanding of how multiple toes coordinate to facilitate geckos' acrobatic locomotion. Here, we tested the control function of gecko toes by running them on vertical substrates varying in orientation, friction and roughness. Sideways wall-running geckos realigned the toes of their top feet upward to resist gravity. Toe contact area was not compromised, but redistributed. Geckos aligned all toes upward to resist slipping when encountering low-friction patches during sideways wall-running. Negotiation of intermittent slippery strips showed an increased contribution of particular toes to compensate for toes that lost adhesion. Increasing substrate roughness using discrete rods perpendicular to sideways locomotion resulted in geckos bending and/or rotating toes to conform to and even grasp the rods, with potential forces more than five times body weight. Geckos increase their effectiveness of manoeuvrability in demanding environments by taking advantage of the distributed control afforded by multiple toes. Our findings provide insight on biological attachment and offer inspiration to advance gecko-inspired robotics and other biomimetic applications.

Anurans are well known for their jumping abilities, making use of their strong hindlimbs. In contrast, the function of the forelimbs during take-off has rarely been studied. We measured the ground reaction forces exerted by forelimbs and hindlimbs during short jumps in the Dybowski's frog Rana dybowskii. Take-off occurred in two phases. Phase one (from the initial time until the forelimbs took off), which lasts a relatively long time (63.2 ± 4.1% of the total take-off phase, N = 20), provides sufficient time for the forelimbs to elevate the body to a suitable posture to deliver the best take-off angle. Phase two (from the forelimbs lift-off until hindlimbs lift-off) was dominated by the hindlimbs which provided a constant and fast elevation. The force angle (angle of the resultant vector from fore-aft and normal force components towards the plane of the substrate) of the hindlimbs and body trajectory was variable before the forelimbs lifted off of the substrate and then primarily followed the direction of the line from the foot-substrate point to the center of mass (COM). The preparation angle adopted when the forelimbs lifted off of the substrate was a good predictor of the take-off angle. The total normal force oscillated around body weight (BW) before the forelimb normal force peaked. The BW shifted from the hindlimbs to the forelimbs during the initial phase of take-off. A simple lever model suggests that the forelimbs are responsible for raising the COM, thus influencing the take-off angle in short jumps.

Locusts (Locusta migratoria manilensis) are characterized by their strong flying and grasping ability. Research on the grasping mechanism and behaviour of locusts on sloping substrates plays an important role in elucidating the mechanics of hexapod locomotion. Data on the maximum angles of slope at which locusts can grasp stably (critical angles of detachment) were obtained from high-speed video recordings at 215 fps. The grasping forces were collected by using two sensors, in situations where all left legs were standing on one and the right legs on the other sensor plate. These data were used to illustrate the grasping ability of locusts on slopes with varying levels of roughness. The grasping morphologies of locusts' bodies and tarsi were observed, and the surface roughness as well as diameters of their claw tips was measured under a microscope to account for the grasping mechanism of these insects on the sloping substrate. The results showed that the claw tips and part of the pads were in contact with the inverted substrate when the mean particle diameter was in the range of 15.3-40.5 µm. The interaction between pads and substrates may improve the stability of contact, and claw tips may play a key role in keeping the attachment reliable. A model was developed to explain the significant effects of the relative size of claw tips and mean particle diameter on grasping ability as well as the observed increase in lateral force (2.09-4.05 times greater than the normal force during detachment) with increasing slope angle, which indicates that the lateral force may be extremely important in keeping the contact reliable. This research lays the groundwork for the probable design and development of biomimetic robotics.

The study of the movement behavior of geckos on a vertical surface, including the measurement and recording of the reaction forces as they move in different directions, plays an important role in understanding the mechanics of the animals' locomotion. This study provides inspiration for the design of a control system for a bionics robot. The three-dimensional reaction forces of vertical surface-climbing geckos (Gekko gecko) were measured using a three-dimensional force-sensors-array. The behavior of gecko as it moved on a vertical surface was recorded with a high speed camera at 215 fps and the function of each foot of a gecko are discussed in this paper. The results showed that the gecko increased its velocity of movement mainly by increasing the stride frequency in the upward, downward and leftward direction and that the speed had no significant relationship to the attachment and detachment times. The feet above the center-of-mass play a key role in supporting the body, driving locomotion and balancing overturning etc. The movement behavior and foot function of geckos change correspondingly for different conditions, which results in safe and effective free vertical locomotion. This research will be helpful in designing gecko-like robots including the selection of gait planning and its control. gecko, three-dimensional reaction force, movement behavior, vertical surface, sensors-array Citation:

Locomotion is an essential character of animals, and excellent moving ability results from the delicate sensing of the substrate reaction forces (SRF) acting on body and modulating the behavior to adapt the motion requirement. The inclined substrates present in habitats pose a number of functional challenges to locomotion. In order to effectively overcome these challenges, climbing geckos execute complex and accurate movements that involve both the front and hind limbs. Few studies have examined gecko's SRF on steeper inclines of greater than 90°. To reveal how the SRFs acting on the front and hind limbs respond to angle incline changes, we obtained detailed measurements of the three-dimensional SRFs acting on the individual limbs of the tokay gecko while it climbed on an inclined angle of 0-180°. The fore-aft forces acting on the front and hind limbs show opposite trends on inverted inclines of greater than 120°, indicating propulsion mechanism changes in response to inclines. When the incline angles change, the forces exerted in the normal and fore-aft directions by gecko's front and hind limbs are reassigned to take full advantage of limbs' different roles in overcoming resistance and in propelling locomotion. This also ensures that weight acts in the angle range between the forces generated by the front and hind limbs. The change in the distribution of SRF with a change in the incline angle is directly linked to the favorable trade-off between locomotive maneuverability and stability.