Excitability level-setting mechanisms in the pons: their behavioral support in decerebrate, reflex standing and freely moving, intact cats.
In the acute precollicular-postmammillary decerebrate cat, stimulation of the mesencephalic locomotor region (MLR) induces "controlled locomotion" on a moving treadmill. Stimulation of the dorsal area and of the ventral area of the pons at its midline elicited a long-lasting decrease and an increase in the tone of the hindlimb extensor muscles, respectively. By selecting the stimulus strength according to the stimulus site, it was possible to set the extensor muscle tone level, that is the "background excitability" of the brain stem and the spinal cord. Locomotor effects induced by MLR stimulation were greatly modified by the set level of background excitability. When the background excitability was high, MLR stimulation evoked "spastic" locomotor movement, while "atonic" locomotor movement was evoked when it was low. Furthermore, stimulation of the ventral area alone also evoked "spastic" locomotor movement. During locomotion in intact cats, stimulation of the dorsal area evoked a series of postural changes. Within a few seconds from the beginning of stimulation, the cat ceased to walk, but maintained a standing posture with or without a locomotor figure. With continuation of this stimulation, it squatted and then lay down on the floor in a sequential manner. Stimulation of the ventral area of the pons evoked an almost opposite series of postural changes. Within a few seconds from the beginning of stimulation, the cat changed from a lying to a squatting posture, and then stood, started to walk and continued to walk during the period of stimulation. All these results demonstrate that an increase in extensor muscle tone and activation of the spinal stepping generator are not separate phenomena, and suggest that integration of neuronal mechanisms involved in the setting of the background excitability and in locomotor movement is a prerequisite for successful expression of locomotor behavior both in decerebrate and intact cats.