The human hand has so many degrees of freedom that it may seem impossible to control. A potential solution to this problem is 'synergy control' which combines dimensionality reduction with great flexibility. With applicability to a wide range of tasks, this has become a very popular concept. In this review, we describe the evolution of the modern concept using studies of kinematic and force synergies in human hand control, cortical and spinal neurons, and electromyographic activity of hand muscles. We go beyond the often purely descriptive usage of the concept of synergy by reviewing the organization of the underlying neuronal circuitry in order to provide mechanistic explanations for various observed synergy phenomena. Finally, we propose a theoretical framework to reconcile important and still debated concepts such as the definitions of ‘fixed’ versus ‘flexible’ synergies and mechanisms underlying the combination of synergies for hand control.

It has been shown that when subjects can predict object properties [e.g., weight or center of mass (CM)], fingertip forces are appropriately scaled before the object is lifted, i.e., before somatosensory feedback can be processed. However, it is not known whether subjects, in addition to these anticipatory force mechanisms, exploit the ability to choose where digits can be placed to facilitate object manipulation. We addressed this question by asking subjects to reach and grasp an object whose CM was changed to the left, center, or right of the object in either a predictable or unpredictable manner. The only task requirement was to minimize object roll during lift. We hypothesized that subjects would modulate contact points but only when object CM location could be predicted. As expected, object roll was significantly smaller in the predictable condition. This experimental condition was also associated with statistically distinct spatial distributions of contact points as a function of object CM location but primarily when large torques had to be counteracted, i.e., for right and left CM locations. In contrast, when subjects could not anticipate CM location, a “default” distribution of contact points was used, this being statistically indistinguishable from that adopted for the center CM location in the predictable condition. We conclude that choice of contact points is integrated with anticipatory force control mechanisms to facilitate object manipulation. These results demonstrate that planning of digit placement is an important component of grasp control.

Dexterous object manipulation requires anticipatory control of digit positions and forces. Despite extensive studies on sensorimotor learning of digit forces, how humans learn to coordinate digit positions and forces has never been addressed. Furthermore, the functional role of anticipatory modulation of digit placement to object properties remains to be investigated. We addressed these questions by asking human subjects (12 females, 12 males) to grasp and lift an inverted T-shaped object using precision grip at constrained or self-chosen locations. The task requirement was to minimize object roll during lift. When digit position was not constrained, subjects could have implemented many equally valid digit position-force coordination patterns. However, choice of digit placement might also have resulted in large trial-to-trial variability of digit position, hence challenging the extent to which the CNS could have relied on sensorimotor memories for anticipatory control of digit forces. We hypothesized that subjects would modulate digit placement for optimal force distribution and digit forces as a function of variable digit positions. All subjects learned to minimize object roll within the first three trials, and the unconstrained device was associated with significantly smaller grip forces but larger variability of digit positions. Importantly, however, digit load force modulation compensated for position variability, thus ensuring consistent object roll minimization on each trial. This indicates that subjects learned object manipulation by integrating sensorimotor memories with sensory feedback about digit positions. These results are discussed in the context of motor equivalence and sensorimotor integration of grasp kinematics and kinetics.

1 The role of vision in controlling leg muscle activation in landing from a drop was investigated. Subjects (n= 8) performed 10 drops from four heights (0.2, 0.4, 0.6 and 0.8 m) with and without vision. Drop height was maintained constant throughout each block of trials to allow adaptation. The aim of the study was to assess the extent to which proprioceptive and vestibular information could substitute for the lack of vision in adapting landing movements to different heights. 2 At the final stages of the movement, subjects experienced similar peak centre of body mass (CM) displacements and joint rotations, regardless of the availability of vision. This implies that subjects were able to adapt the control of landing to different heights. The amplitude and timing of electromyographic signals from the leg muscles scaled to drop height in a similar fashion with and without vision. 3 However, variables measured throughout the execution of the movement indicated important differences. Without vision, landings were characterised by 10 % larger ground reaction forces, 10 % smaller knee joint rotations, different time lags between peak joint rotations, and more variable ground reaction forces and times to peak CM displacement. 4 We conclude that non-visual sensory information (a) could not fully compensate for the lack of continuous visual feedback and (b) this non-visual information was used to reorganise the motor output. These results suggest that vision is important for the very accurate timing of muscle activity onset and the kinematics of landing.