Convergent force fields organized in the frog's spinal cord

Abstract

Microstimulation of the gray matter of the frog's spinal cord was used to elicit motor responses. Force responses were recorded with the frog's ankle clamped while EMG activity was monitored. The collections of force patterns elicited at different leg configurations were summarized as force fields. These force fields showed convergence to an equilibrium point. The equilibrium paths were calculated from the force fields with the leg clamped. These paths predicted free limb motion in 75% of trials. The force fields were separated into active and prestimulation resting responses. The active force field responses had a fixed position equilibrium. These active force fields were modulated in amplitude over time, although the balance and orientations of forces in the pattern remained fixed. The active fields grouped into a few classes. These included both convergent and parallel fields. The convergent force fields (CFFS) could be observed in deafferented preparations. Motoneuron (MN) activity underlying the force fields was marked using sulforhodamine. The marked activity covered several segments. Several simulations and MN stimulations show that topography, limb geometry, and random activation could not account for the results. It is likely that propriospinal interneurons distribute the activity that underlies the responses observed here. Experiments showed that CFFs that resemble those elicited by microstimulation also underlie natural behaviors. The full variety of fields revealed by microstimulation was larger than the repertoire elicited by cutaneous stimulation. It was concluded that fixed-pattern force fields elicited in the spinal cord may be viewed as movement primitives. These force fields could form building blocks for more complex behaviors.