Biomechanical functions analysis of the Mallard webbed foot: A study of macroscopic and microscopic material assembly and tendon morphology.

The mallard webbed foot represents an exemplary model of biomechanical efficiency in avian locomotion. This study delves into the intricate material assembly and tendon morphology of the mallard webbed foot, employing both macroscopic and microscopic analyses. Through histological slices and scanning electron microscopy (SEM), we scrutinized the coupling assembly of rigid and flexible materials such as skin, tendon, and bone, while elucidating the biomechanical functions of tendons across various segments of the tarsometatarsophalangeal joint (TMTPJ). The histological examination unveiled a complex structural hierarchy extending from the external integument to the skeletal framework. Notably, the bone architecture, characterized by compact bone and honeycombed trabeculae, showcases a harmonious blend of strength and lightweight design. Tendons, traversing the phalangeal periphery, surrounded by elastic fibers, collagen fibers, and fat tissue. Fat chambers beneath the phalanx, filled with adipocytes, provide effective buffering, enabling the phalanx to withstand gravity, provide support, and facilitate locomotion. Furthermore, SEM analysis provided insights into the intricate morphology and arrangement of collagen fiber bundles within tendons. Flexor tendons in proximal and middle TMTPJ segments adopt a wavy-type, facilitating energy storage and release during weight-bearing activities. In contrast, distal TMTPJ flexor tendons assume a linear-type, emphasizing force transmission across phalangeal interfaces. Similarly, extensor tendons demonstrate segment-specific arrangements tailored to their respective biomechanical roles, with wavy-type in proximal and distal segments for energy modulation and linear-type in middle segments for enhanced force transmission and tear resistance. Overall, our findings offer a comprehensive understanding of the mallard webbed foot's biomechanical prowess, underscoring the symbiotic relationship between material composition, tendon morphology, and locomotor functionality. This study not only enriches our knowledge of avian biomechanics but also provides valuable insights for biomimetic design and tissue engineering endeavors.

Active Adaptive Strategies of Mallard Feet in Response to Changes in Wetness and Compactness of the Sand Terrain.

Mallards (Anas platyrhynchos) exhibit exceptional locomotive abilities in diverse terrains, such as beaches, swamps, and tidal flats. This capability is primarily attributed to their unique webbed toe structure and cooperative locomotion posture of their feet. Therefore, this study aims to further delve into the active adaptive strategies of mallard feet in response to diverse external environmental conditions. Six adult male mallards were selected for this research. Their locomotion on sandy surfaces with differing wetness levels and varying degrees of compaction were captured using a high-speed camera, and analysis of instantaneous and continuous changes in the primary joint angles of the mallards' feet, including the toe-webbed opening and closing angles, the tarsometatarsal-phalangeal joint (TMTPJ), and the intertarsal joint (ITJ). It was found that on loose sandy surfaces, increasing wetness expanded the ground contact area of the mallards' feet. This led to greater flexion at the TMTPJ joint during mid-stance, accompanied by decreased flexion of the ITJ during touch-down and mid-stance. Conversely, on compacted sand, increasing wetness resulted in a reduced foot effect area and lessened ITJ flexion at both touch-down and mid-stance. Furthermore, on looser sand, the ground contact area of the mallards' feet decreased, with an increase in ITJ buckling at touch-down. During the swing phase, sand wetness and compactness effected minimally on the feet of the mallards. On dry and loose sand ground, mallards will contract their second and fourth toes with webbing upon ground contact, covering and compacting the sand beneath while increasing ITJ flexion to mitigate sinking. This adaptation reduces the energy expended on sand and enhances body stability. In wet and compacted sand conditions, mallards expand their second and fourth toes upon ground contact and reduce ITJ flexion. Therefore, this coordinated foot and ITJ locomotion offers mallards a natural advantage when moving on various environmental media.

From the Analysis of Anatomy and Locomotor Function of Biological Foot Systems to the Design of Bionic Foot: An Example of the Webbed Foot of the Mallard.

This study utilized the mallard's foot as the subject, examining the bone distribution via computed tomography (CT) and analyzing pertinent parameters of the tarsometatarsal bones. Additionally, gross anatomy methods were employed to elucidate the characteristics of the toes and webbing bio-structures and their material composition. Biologically, the mallard's foot comprises tarsometatarsal bones and 10 phalanges, enveloped by fascia, tendons, and skin. Vernier calipers were used to measure the bones, followed by statistical analysis to acquire structural data. Tendons, originating in proximal muscles and terminating in distal bones beneath the fascia, facilitate force transmission and systematic movement of each segment's bones. Regarding material composition, the skin layer serves both encapsulation and wrapping functions. Fat pads, located on the metatarsal side of metatarsophalangeal joints and each phalanx, function as cushioning shock absorbers. The correlation between the force applied to the tarsometatarsal bones and the webbing opening angle was explored using a texture analyzer. A simplified model describing the driving force behind the webbing opening angle was introduced. Furthermore, we designed a bionic foot, contributing a foundational reference for anti-sinking bionic foot development.

Effects of particle size and thickness of quartz sand on the webbed foot kinematics of mallard (Anas platyrhynchos).

The webbed foot structure of mallards (Anas platyrhynchos) exhibits effective anti-subsidence properties when walking on soft ground. To investigate the effects of quartz sand particle size and thickness on joint angles and the movement patterns of webbed feet, we created a testing substrate with quartz sand and utilized high-speed cameras and kinematic analysis tools for data acquisition. Mallards mainly adjusted the tarsometatarso-phalangeal joint (TMTPJ) during touch-down and lift-off stages in response to increasing particle size or enhanced ground roughness. Conversely, adjustments to the intertarsal joint (ITJ) predominantly took place during mid-stance. Conversely, mallards predominantly adjusted the ITJ during touch-down and lift-off when coping with increased quartz sand thickness, with TMTPJ adjustments mainly occurring at touch-down. As quartz sand particle size increased, the TMTPJ angle increased, the ITJ angle decreased, toe closure advanced, and the duty factor decreased throughout the entire stride cycle. In contrast, increasing quartz sand thickness led to more delayed TMTPJ adjustments, slower webbed foot closure, and an increased duty factor throughout the stride cycle. Mallards modify their leg posture to notably decrease the touch-down foot angle upon encountering sandy terrain. This action subsequently forms a depression beneath their feet, contributing to sand consolidation and limiting flow. During the stance phase, the mallard's weight is distributed across the webbed foot, generating minimal pressure and preventing significant subsidence while walking on sandy ground.

Effects of the speed on the webbed foot kinematics of mallard (Anas platyrhynchos).

In this study, the effect of the speed on the webbed foot locomotion of the mallard was analyzed based on a considerable number of reliable indoor test data. Four adult male mallards were selected for analysis, and the locomotion speed of the mallard was controlled using the treadmill at an accurate and adjustable speed. The locomotion pattern of the webbed foot of the mallard at different speeds was recorded using a high-speed camera. The changes in the position and conformation of the webbed foot during locomotion on a treadmill were tracked and analyzed using Simi-Motion kinematics software. The results indicated that the stride length of the mallard increased, and the stance phase duration was shortened with the increase of the speed, whereas the swing phase duration did not vary significantly. The duty factor decreased with the increase of the mallard speed but not drop below to 0.5, because the mallards flew with their wings, or moved backward relative to the treadmill with the further increase of the speed. Using the energy method to further distinguish gait, and through the percentage of congruity analysis, it was found that between 0.73 and 0.93 m/s, the gait experienced a transition from walking to grounded running, with no significant changes in spatiotemporal parameters. At speeds between 0.93 and 1.6 m/s, mallards adopt a grounded running gait. The instantaneous changes of the tarsometatarso-phalangeal joint (TMTPJ) angle and the intertarsal joint (ITJ) angle at touch-down, mid-stance and lift-off concomitant with the change of the speed were examined with the TMTPJ and ITJ angle as the research objects. Moreover, the continuous changes of the joint angles were examined in a complete stride cycle. The result indicated that the increase of the speed will also make the TMTPJ and ITJ angle change ahead of time in a stride cycle, proving the shortened stance phase duration. The ITJ angle changed much more than the TMTPJ. Thus, the above result reveals that the mallard primarily responds with the increase of the speed by adjusting the ITJ, instead of the TMTPJ. The vertical displacement of the toe joint points and the toe joint angle was studied (α joint angle is between the second toe and the third toe; ß joint angle is between the third toe and the fourth toe) with a complete stride cycle as the research object. The distal phalanxes of the second, third and fourth toes first contacted the ground, and the proximal phalanx touched the ground in turn during the early stance phase duration of the mallard, as indicated by the result of this study. However, the toes got off the ground in turn from the proximal phalanxes when the mallard foot got off the ground. With the decrease of the interphalangeal α and ß joint angles, the foot web tended to be close and rapidly recovered before the next touch-down. The above result reveals that the webbed foot of the mallard is a coupling system that plays a role in the adjustment of speed.

Forelimb joints contribute to locomotor performance in reindeer (Rangifer tarandus) by maintaining stability and storing energy.

Reindeer (Rangifer tarandus) have lengthy seasonal migrations on land and their feet possess excellent locomotor characteristics that can adapt to complex terrains. In this study, the kinematics and vertical ground reaction force (GRF) of reindeer forelimb joints (interphalangeal joint b, metacarpophalangeal joint c, and wrist joint d) under walk, trot 1, and trot 2 were measured using a motion tracking system and Footscan pressure plates. Significant differences among different locomotor activities were observed in the joint angles, but not in changes of the joint angles (α b , α c , α d ) during the stance phase. Peak vertical GRF increased as locomotor speed increased. Net joint moment, power, and work at the forelimb joints were calculated via inverse dynamics. The peak joint moment and net joint power related to the vertical GRF increased as locomotor speed increased. The feet absorbed and generated more energy at the joints. During different locomotor activities, the contribution of work of the forelimbs changed with both gait and speed. In the stance phase, the metacarpophalangeal joint absorbed more energy than the other two joints while trotting and thus performed better in elastic energy storage. The joint angles changed very little (∼5°) from 0 to 75% of the stance phase, which reflected the stability of reindeer wrist joints. Compared to typical ungulates, reindeer toe joints are more stable and the stability and energy storage of forelimb joints contribute to locomotor performance in reindeer.

Study on bio-inspired feet based on the cushioning and shock absorption characteristics of the ostrich foot.

As the main actuator of high-speed running, the ostrich feet are highly capable of cushioning and shock absorption. In this study, based on the elastic modulus scales and assembly order of the 3rd toe soft tissues and the functions of the metatarsophalangeal (MTP) joint, we designed fourteen bio-inspired feet. The impact process on loose sand was simulated on the finite element software Abaqus. Also the stress distributions and deformations of each component of the bio-inspired feet were clarified. With the peak acceleration as the index, the cushioning performances of the bio-inspired feet were compared on both loose sand and solid ground through height-variable impact tests. The 15-15-15 HA (hardness unit) bio-inspired foot showed lower peak acceleration and thereby better cushioning performance, but larger deformation, less-uniform stress distribution and thereby lower stability than the 15-35-55 HA bio-inspired foot. In fact, the silicon rubbers with different hardness degrees (which simulate the elasticity modulus scales of the digital cushions, fascia and skin) and the spring mechanism (which simulates the functions of the MTP joint) work as an "integrated system" of cushioning and shock absorption.

Design and Analysis of the Bionic Mechanical Foot with High Trafficability on Sand.

The ostrich foot has excellent travelling performance on sand and plays a vital role in efficient locomotion. The tendon-bone assembly characteristics of an ostrich foot were studied by gross anatomy, and the 3D model of ostrich foot was reconstructed and analyzed using reverse engineering techniques. Further, the bionic mechanical foot, suitable for locomotion on loose sand, was designed based on the structural characteristics of ostrich foot and its rigid-flexible coupling mechanism of tendon-bone synergies. The travelling performance on sand of the bionic mechanical foot was tested on a test platform by using Simi Motion. After analyzing the angle changes of the ankle joint and the metatarsophalangeal (MTP) joint, the displacement changes of the knee joint, the ankle joint, the MTP joint, and each phalanx along the z-axis, the plantar pressure distribution, and the footprints, we drew the conclusion that the bionic mechanical foot is helpful to reduce the sinkage and improve the trafficability on sand ground. This study provides a new research method for the walking mechanism of a robot and deep space exploration platform walking on soft ground.

FEM analysis in excellent cushion characteristic of ostrich (Struthio camelus) toe pads.

African ostrich (Struthio camelus) is the largest and fastest extent bipedal animal. The ostrich mainly relies on the 3rd toe to support the entire body under high-speed motion. The short and severe impact concentrated on the limited area would produce tremendous momentary internal stress and strain, which may contribute to the phalanges disloaction, soft tissue damage and the like. The vibration and excessive negative acceleration caused by the ground reaction force also affect the stability of the touching process. Therefore, ostrich toe pads are required to have excellent cushion characteristics. However, current studies often explains the cushion properties by analyzing the macro-microscopic structure of the pad organism, and there is a paucity of research on its biomechanical behaviour. Consequently, from the perspective of multi-layer structure and biomaterial assembly, this study aims to explain the biomechanical characteristics of the ostrich toe pads by FEM (Finite Element Method) analysis. Based on results, we deem that the ostrich toe pad could absorb energy and reduce vibration effectively. Firstly, the multi-layer structure of the pads make the stress and strain decay from outside to inside. Secondly, the minimal response frequency of the pad is 164.22 Hz, making it effectively avoid resonance phenomenon. Finally, the composite material model has the best performance in decreasing the negative acceleration peak value, the impact force peak value and the maximal equivalent stress value at velocities of 0.669 m/s and 1.339 m/s. These results help to further understand the buffer mechanism of the ostrich toe pad, and also have important inter-species reference value for the pathogenesis of human foot soft tissue injury.

Phalangeal joints kinematics in ostrich (Struthio camelus) locomotion on sand.

In ostriches, the toes are the only body parts that contact loose sand surfaces. Thus, toe interphalangeal joint motions may play vital biomechanical roles. However, there is little research on ostrich phalangeal joint movements while walking or running on sand. The results from the three-dimensional motion track analysis system Simi Motion show that gait pattern has no significant effect on the key indicators (angles at touch-down, mid-stance, lift-off and range of motion) of the toe joint angles. The motion of the toe phalanges when walking and running on sand is basically the same. The ground medium is the key factor that changes the toe postures adopted by ostriches during the stance phase in slow to fast locomotion. The 3rd toe and the 4th toe are connected by the interphalangeal ligament, and the motions of the MTP3 and MTP4 joints are highly synchronized on a loose sand substrate. The 3rd toe and 4th toe are coupled to maintain static balance in slow locomotion and dynamic balance in fast locomotion. In addition, the gait pattern has a marked effect on the range of forward displacement of the toenail (YTN). The ostrich toenail plays an important role in preventing slip and provides traction at push-off in a sandy environment. The metatarsophalangeal joint plays an important role in energy saving during fast locomotion on loose sand substrates. Simulation reveals that the particle velocity field, particle force field and sand particle disturbance in the running gait are denser than those in the walking gait.

Macroscopic and microscopic analyses in flexor tendons of the tarsometatarso-phalangeal joint of ostrich (Struthio camelus) foot with energy storage and shock absorption.

Flexor tendons function as energy storage and shock absorption structures in the tarsometatarso-phalangeal joint (TMTPJ) of ostrich feet during high-speed and heavy-load locomotion. In this study, mechanisms underlying the energy storage and shock absorption of three flexor tendons of the third toe were studied using histology and scanning electron microscopy (SEM). Macroscopic and microscopic structures of the flexor tendons in different positions of TMTPJ were analyzed. Histological slices showed collagen fiber bundles of all flexor tendons in the middle TMTPJ were arranged in a linear-type, but in the proximal and distal TMTPJ, a wavy-type arrangement was found in the tendon of the M. flexor digitorum longus and tendon of the M. flexor perforans et perforatus digiti III, while no regular-type was found in the tendon of the M. flexor perforatus digiti III. SEM showed that the collagen fiber bundles of flexor tendons were arranged in a hierarchically staggered way (horizontally linear-type and vertically linear-type). Linear-type and wavy-type both existed in the proximal TMTPJ for the collagen fiber bundles of the tendon of the M. flexor perforatus digiti III, but only the linear-type was found in the distal TMTPJ. A number of fibrils were distributed among the collagen fiber bundles, which were likely effective in connection, force transmission and other functions. The morphology and arrangement of collagen fiber bundles were closely related to the tendon functions. We present interpretations of the biological functions in different positions and types of the tendons in the TMTPJ of the ostrich feet.

Plantar pressure distribution of ostrich during locomotion on loose sand and solid ground.

BACKGROUND: The ostrich is a cursorial bird with extraordinary speed and endurance, especially in the desert, and thus is an ideal large-scale animal model for mechanic study of locomotion on granular substrate. METHODS: The plantar pressure distributions of ostriches walking/running on loose sand/solid ground were recorded using a dynamic pressure plate. RESULTS: The center of pressure (COP) on loose sand mostly originated from the middle of the 3rd toe, which differed from the J-shaped COP trajectory on solid ground. At mid-stance, a high-pressure region was observed in the middle of the 3rd toe on loose sand, but three high-pressure regions were found on solid ground. The gait mode significantly affected the peak pressures of the 3rd and 4th toes (p = 1.5 × 10-6 and 2.39 × 10-8, respectively), but not that of the claw (p = 0.041). The effects of substrate were similar to those of the gait mode. DISCUSSION: Ground reaction force trials of each functional part showed the 3rd toe bore more body loads and the 4th toe undertook less loads. The pressure distributions suggest balance maintenance on loose sand was provided by the 3rd and 4th toes and the angle between their length axes. On loose sand, the middle of the 3rd toe was the first to touch the sand with a smaller attack angle to maximize the ground reaction force, but on solid ground, the lateral part was the first to touch the ground to minimize the transient loading. At push-off, the ostrich used solidification properties of granular sand under the compression of the 3rd toe to generate sufficient traction.