In this study, we describe functional characteristics of neurons forming networks generating oral ingestive motor behaviours. Neurons in medial reticular nuclei on the right side of the brainstem between the trigeminal and hypoglossal motor nuclei were recorded in anaesthetized and paralysed rabbits during two types of masticatory-like motor patterns induced by electrical stimulation of the left (contralateral) or right (ipsilateral) cortical masticatory areas. Sixty-seven neurons in nucleus reticularis pontis caudalis (nPontc), nucleus reticularis parvocellularis (nParv), and nucleus reticularis gigantocellularis (Rgc) were studied. These were classified as phasic or tonic depending on their firing pattern during the fictive jaw movement cycle. Phasic neurons located in the dorsal part of nPontc were active during the jaw opening phase, whilst those in dorsal nParv tended to fire during the closing phase. In most neurons, burst duration and firing frequency changed between the two motor patterns, but there was little change in phase of firing. Tonic units were mainly recorded in the ventral half of nPontc, and at the junction between Rgc and caudal nParv. Cortical inputs with short latency from the contralateral masticatory area were more frequent in phasic (82%) than tonic (44%) neurons, whilst inputs from the ipsilateral cortex were equal in the two subgroups (57% and 56%). Phasic neurons had significantly shorter mean contralateral than ipsilateral cortical latencies, whilst there was no difference among tonic neurons. Intra- and perioral primary afferent inputs activated both types of neurons at oligo-synaptic latencies. Our results show that subpopulations of neurons in medial reticular nuclei extending from the caudal part of the trigeminal motor nucleus to the rostral third of the hypoglossal motor nucleus are active during the fictive masticatory motor behaviour. Unlike masticatory neurons in the lateral tegmentum, the medial subpopulations are spatially organized according to discharge pattern.
Neurons of the dorsal nucleus reticularis pontis caudalis (nPontc) fire rhythmically during fictive mastication, while neurons of the ventral half tend to fire tonically (Westberg et al., 2001). This paper describes the changes in the pattern of rhythmical mastication elicited by stimulation of the sensorimotor cortex during inhibition or excitation of neurons in this nucleus and adjacent parts of nucleus reticularis gigantocellularis (Rgc) in the anaesthetized rabbit. Masticatory movements and electromyographic (EMG) activity of the masseter and digastric muscles produced by cortical stimulation were recorded before, during and after injections of a local anaesthetic (lidocaine) or excitatory amino acid N-methyl-d-aspartate (NMDA) into nPontc and Rgc through a microsyringe with attached microelectrode to record neuronal activity. Lidocaine inhibited local neurons and modified the motor program, and the effects varied with the site of injection. Most injections into the ventral half of nPontc increased cycle duration, digastric burst duration and burst area. The action of lidocaine in dorsal nPontc was more variable, although burst duration and area were often decreased. The effects on the muscle activity were always bilateral. Lidocaine block of the rostromedial part of Rgc had no effect on movements or on EMGs. Injections of NMDA excited local neurons and when injected into ventral nPontc, it completely blocked mastication. Dorsal injections either had no effect or increased cycle frequency, while decreasing burst duration and area. No increases in EMG burst duration or area were observed with NMDA. Our findings suggest that neurons of ventral nPontc tonically inhibit other parts of the central pattern generator during mastication, while dorsal neurons have mixed effects. We incorporated these findings into a new model of the masticatory central pattern generator.
The rabbit digastric muscle has a single belly that opens and retracts the mandible. It does not contain connective tissue partitions, and all fibers arise from the same tendon and insert into a single broad site. Historically, it was assumed that the muscle functioned as a single unit. Since we had preliminary evidence that this might not be the case, we carried out five small studies in rabbits. First, we showed that electromyographic (EMG) activity varies between recording sites within the muscle during the masticatory cycle induced by repetitive stimulation of the sensorimotor cortex. We found that EMG activity in the caudal region sometimes began before the anterior EMG during mastication when the jaw swung to the side of the muscle, but the two regions became active at the same time during other patterns. We next showed that separate branches of the mylohyoid nerve enter the anterior, intermediate and caudal regions of the digastric. However, a separate study showed that the motor endplates were distributed across a continuous sheet, consistent with a single anatomical partition. We then stimulated single nerve branches to deplete glycogen. By comparing the optical density of fibers labeled by the periodic acid-Schiff method for glycogen, we were able to show that the three branches innervate separate regions of the muscle. Finally, we applied either FluoroGold or Fast Blue dyes to the central cut ends of the branches to label the cell bodies of the three pools of motoneurons. These were found within the middle and caudal thirds of the trigeminal motor nucleus, but there appeared to be no spatial separation of the three pools or double labeling of cells. We conclude that the digastric muscle contains two and possibly three functional subregions. The fact that the motoneurons are intermingled suggests that the distribution of motor commands to the three pools is not based on their location.
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