Dextran-amines were used as retrograde tracers to investigate the organization of cortical projections to different cytoarchitectonic regions of the pontomedullary reticular formation of the cat. Injections into the nucleus reticularis pontis oralis resulted in labelling of neurones in the proreus cortex and area 6a beta of the premotor cortex, with little labelling in the motor cortex (area 4). This labelling was predominantly ipsilateral to the injection site. In contrast, injections into the nucleus reticularis pontis caudalis (NRPc), nucleus reticularis gigantocellularis (NRGc), and nucleus reticularis magnocellularis (NRMc) resulted in bilateral labelling--primarily in areas 6a beta, 6a gamma, and in the rostromedial region of area 4--with little labelling in the proreus cortex. In general, the cortical projections to the caudal NRGc and the NRMc were larger than those to the NRPc. More than 25% of the total projections to each of the latter three reticular regions arose from the medial part of area 4. Labelling in the hindlimb regions of area 4 was largest following the NRMc injections and smallest after injections in the NRPc. The projections to the NRPc originated from more medial parts of areas 4 and 6 than did the projections to the caudal region of the NRGc. These results suggest that areas 4 and 6 may be able to differentially activate different regions of the pontomedullary reticular formation depending on the movement that is made and perhaps also on the context of that movement.
Effects of red nucleus microstimulation on the locomotor pattern and timing in the intact cat: a comparison with the motor cortex. To determine the extent to which the rubrospinal tract is capable of modifying locomotion in the intact cat, we applied microstimulation (cathodal current, 330 Hz; pulse duration 0.2 ms; maximal current, 25 microA) to the red nucleus during locomotion. The stimuli were applied either as short trains (33 ms) of impulses to determine the capacity of the rubrospinal tract to modify the level of electromyographic (EMG) activity in different flexors and extensors at different phases of the step cycle or as long trains (200 ms) of pulses to determine the effect of the red nucleus on cycle timing. Stimuli were also applied with the cat at rest (33-ms train). This latter stimulation evoked short-latency (average = 11.8-19.0 ms) facilitatory responses in all of the physiological flexor muscles of the forelimb that were recorded; facilitatory responses were also common in the elbow extensor, lateral head of triceps but were rare in the physiological wrist and digit extensor, palmaris longus. Responses were still evoked in most muscles when the current was decreased to near threshold (3-10 microA). Stimulation during locomotion with the short trains of stimuli evoked shorter-latency (average = 6.0-12.5 ms) facilitatory responses in flexor muscles during the swing phase of locomotion and, except in the case of the extensor digitorum communis, evoked substantially smaller responses in stance. The same stimuli also evoked facilitatory responses in the extensor muscles during swing and produced more complex effects involving both facilitation and suppression in stance. Increasing the duration of the train to 200 ms modified the amplitude and duration of the EMG activity of both flexors and extensors but had little significant effect on the cycle duration. In contrast, whereas stimulation of the motor cortex with short trains of stimuli during locomotion had very similar effects to that of the red nucleus, increasing the train duration to 200 ms frequently produced a marked reset of the step cycle by curtailing stance and initiating a new period of swing. The results suggest that whereas both the motor cortex and the red nucleus have access to the interneuronal circuits responsible for controlling the structure of the EMG activity in the step cycle, only the motor cortex has access to the circuits responsible for controlling cycle timing.
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