Most primates use diagonal sequence (DS), diagonal couplets (DC) gaits when they walk or run quadrupedally, and it has been suggested that DSDC gaits contribute to stability in their natural arboreal habitats compared to other symmetrical gaits. However, this postulate is based solely on studies of primate gaits using continuous terrestrial and arboreal substrates. A particular species may select suitable gaits according to the substrate properties. Here, we analyzed the gaits of Japanese macaques moving on a horizontal ladder with rung intervals ranging from 0.40 to 0.80 m to elucidate the relative advantages of each observed form of gait. The rung arrangement forced our macaques to choose either diagonal coupling or DS gaits. One macaque consistently used diagonal coupling (i.e., DSDC and LSDC gaits) across narrow and intermediate rung intervals, whereas the other macaque used DS gaits (i.e., DSDC and DSLC gaits). At wider rung intervals, both macaques shifted to a two-one sequence (TOS), which is characterized by two nearly simultaneous touchdowns of both forelimbs and one touchdown of each hind limb in a stride. The transition to the TOS sequence increased the duration of support on multiple limbs, but always included periods of a whole-body aerial phase. These results suggest that Japanese macaques prefer DSDC gaits, because the diagonal coupling and DS contribute separately to stability on complex supports compared to the lateral coupling and lateral sequence. We also postulate that stability triggers the transition from symmetrical gaits to the TOS sequence.

The primate hand adopts a variety of postures during locomotion. Habitually terrestrial cercopithecine primates are known to use a palmigrade posture at faster speeds to possibly mitigate stresses on the hand skeleton; however, it is unclear whether arboreal or semi-terrestrial species use a similar strategy for adjusting hand posture. Here, we explored intra-individual variation in hand contact patterns during terrestrial locomotion in the Japanese macaque (Macaca fuscata), a semi-terrestrial cercopithecine primate. Two monkeys were required to walk on the ground at their own preferred speeds or were encouraged to move faster for food rewards. The contact area under the hand and ground reaction forces (GRFs) were measured simultaneously using a tactile pressure sensor, and then hand pressures were calculated offline. We found that hand contact patterns could vary within individuals. The monkeys used predominantly a palmigrade posture within the range of speeds covered in this study (0.72-2.56 m s). There were two subtypes of palmigrade posture. In one subtype, the hypothenar pad did not contact the substrate, whereas the entire hand contacted the substrate in the other. The palm of the palmigrade hand with total-hand contact experienced similar or lower peak pressure and pressure-time integral than those of the palmigrade hand without hypothenar pad contact even though it experienced higher peak GRFs. The moderate peak pressure experienced by the palmigrade hand with total-hand contact was due to increased contact area under the palm. The total contact area of the fingers and peak GRF to the fingers were relatively unchanged with different patterns of hand contact. These findings provide evidence that when walking on the ground, semi-terrestrial Japanese macaques use a palmigrade posture with total-hand contact to attenuate stresses on hand bones, as do habitually terrestrial species.

Primate hands and feet are versatile and offer an opportunity to examine how morphology reflects compromises among different functions in daily activities. In this study, we investigated the diaphyseal robusticity of metacarpals (MCs) and metatarsals (MTs) and discussed their correlation with locomotor and other behaviors in the semiterrestrial Japanese macaque (Macaca fuscata). The objectives of this study were to determine (1) whether more robust MCs and MTs experience higher forces during terrestrial locomotion than less robust bones; and (2) whether MT3, which is suggested to be the functional axis of the foot in Japanese macaques, is more robust than MT2, MT4, and MT5. Computed tomography of MCs1-5 and MTs1-5 was performed in 10 monkeys. As a measure of bone robusticity, the buckling strength of each MT and MC was calculated as J/L 2 where J is the polar second moment of area and L is the bone length. Hand and foot pressure were recorded using plantar pressure measurement systems while two monkeys moved on a flat floor over a range of speeds (0.72-2.56 m/s). The relationship between the bone robusticity and the load applied to the bones during terrestrial locomotion was analyzed. Our results did not support the two predictions. There was no positive correlation between diaphyseal robusticity and the peak force in both male and female Japanese macaques. There was no clear difference in bone robusticity among MTs2-5 in both males and females. These results suggest that the relation between MC and MT robusticity and mechanical loading during locomotion is not as straightforward as might be expected, possibly due to the complex multifunctionality of primate hands and feet. Additional integrative studies that similarly incorporate morphological and experimental approaches are expected to provide useful insights into macaque hand and foot morphology.

To better place the form and motion of the human foot in an evolutionary context, understanding how foot motions change when quadrupeds walk bipedally can be informative. For this purpose, we compared the pressures beneath the foot during bipedal and quadrupedal walking in Japanese macaques (Macaca fuscata). The pressure at nine plantar regions was recorded using a pressure mat (120 Hz), while the animals walked on a level walkway at their preferred speeds. The results revealed substantial differences in foot use between the two modes of locomotion, and some features observed during bipedal walking resembled human gait, such as the medial transfer of the center of pressure (COP), abrupt declines in forefoot pressures, and the increased pressure beneath the hallux, all occurring during the late-stance phase. In particular, the medial transfer of the COP, which is also observed in bonobos (Vereecke et al.: Am J Phys Anthropol 120 (2003) 373-383), was due to a biomechanical requirement for a hind limb dominant gait, such as bipedal walking. Features shared by bipedal and quadrupedal locomotion that were quite different from human locomotion were also observed: the heel never contacted the ground, a foot longitudinal arch was absent, the hallux was widely abducted, and the functional axis was on the third digit, not the second.