Acquisition of operant-trained bipedal locomotion in juvenile Japanese monkeys (Macaca fuscata): a longitudinal study

This study investigated developmental aspects of the acquisition of operant-trained bipedal (Bp) standing and Bp walking in the normally quadrupedal (Qp) juvenile Japanese monkey (M. fuscata). Four male monkeys (age: 1.6 to 2.4 years, body weight: 3.3 to 4.6 kg) were initially operantly trained to stand upright on a smooth floor and a stationary treadmill belt (width = 60 cm, walking length = 150 cm). They were then trained to walk bipedally on the moving treadmill belt (speed: 0.4-0.7 m/s). A regular training program (5 days/week; 30-60 min/day) was given to each monkey for the first 40 to 60 days, followed by less intensive training. After the beginning of locomotor training, upright postural stability and Bp walking capability were assessed kinematically for 592, 534, 526, and 537 days on monkeys A, B, C, and D, respectively. Left side- and back-views of the walking monkey were photographed (10 frames/s) and videotaped (250 frames/s). Stick figures of the head, body, and hindlimbs were drawn with reference to ink-marks positioned in front of the ear and over the pivot points of hindlimb joints. All kinematic data were digitized and analyzed using image-analyzing software. After sufficient physical growth and locomotor training, all the monkeys gradually acquired: (a) a more upright and a more stable posture with a constant body axis orientation during Bp locomotion; (b) a more stable and a stronger functional coupling between the body and hindlimb movements with a less anterior (A)-posterior (P) fluctuation of a body axis; (c) a smaller leftward (Lt)-rightward (Rt) displacement of the midline pelvic position, allowing the monkey to walk along a straight course; (d) a more coordinated relationship among hip-knee, knee-ankle, and ankle-metatarsophalangeal (MTP) joints; and finally (e) the acquisition of well-coordinated Bp walking even at high treadmill belt speeds up to 1.5 m/s. All of these results demonstrated the capability of the physically developing monkey to integrate the neural and musculoskeletal mechanisms required for sufficient coordination of upper (head, neck, trunk) and lower (hindlimbs) motor segments so that Bp standing and Bp walking could be elaborated.