Functional differentiation of trailing and leading forelimbs during locomotion on the ground and on a horizontal branch in the European red squirrel (Sciurus vulgaris, Rodentia)

Mammalian locomotion is characterized by the frequent use of in-phase gaits in which the footfalls of the left and right fore- or hindlimbs are unevenly spaced in time. Although previous studies have identified a functional differentiation between the first limb (trailing limb) and the second limb (leading limb) to touch the ground during terrestrial locomotion, the influence of a horizontal branch on limb function has never been explored. To determine the functional differences between trailing and leading forelimbs during locomotion on the ground and on a horizontal branch, X-ray motion analysis and force measurements were carried out in two European red squirrels (Sciurus vulgaris, Rodentia). The differences observed between trailing and leading forelimbs were minimal during terrestrial locomotion, where both limbs fulfill two functions and go through a shock-absorbing phase followed by a generating phase. During locomotion on a horizontal branch, European red squirrels reduce speed and all substrate reaction forces transmitted may be due to the reduction of vertical oscillation of the center of mass. Further adjustments during locomotion on a horizontal branch differ significantly between trailing and leading forelimbs and include limb flexion, lead intervals, limb protraction and vertical displacement of the scapular pivot. Consequently, trailing and leading forelimbs perform different functions. Trailing forelimbs function primarily as shock-absorbing elements, whereas leading forelimbs are characterized by a high level of stiffness. This functional differentiation indicates that European red squirrels ‘test’ the substrate for stability with the trailing forelimb, while the leading forelimb responds to or counteracts swinging or snapping branches.