Organization within the cranial IX–X complex in ranid frogs: A horseradish peroxidase transport study

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Abstract
Cranial nerves IX and X in frogs have been described as originating from a nuclear group referred to as the IX–X complex. We studied the central nervous system components of this complex in Rana pipiens and R. catesbiana by labeling peripheral branches of cranial nerves IX and X and identifying the central nervous system contributions of these branches. Various peripheral nerves (IX and the cardiac, gastric, pulmonary, and laryngeal branches of X) were identified and soaked in horseradish peroxidase (HRP). One to 2 weeks later, the frogs were killed and processed for HRP by the tetramethylbenzidine method. Glossopharyngeal efferents originated from a small ventrolateral cell group found at the level of the IX root exit. Vagal efferents formed a single column of cells in a ventrolateral position from the level of the brainstem exit of the vagus nerve (approximately 2,000 μm above the obex) to 200 μm below the obex (values given are for an 80-g frog). This cell group was separate from and just caudal to efferent cells of the glossopharyngeal nerve. Within the vagal portion of the column, cells projecting through the gastric branch were found throughout the rostral-caudal extent of the nucleus. “Cardiac” cells tended to be more rostral than “pulmonary” cells, and both groups of cells were located in the middle of the nucleus. “Laryngeal” cells were located more caudally in the nucleus. This peripheral representation within the vagal nucleus corresponds more closely to the organization found in the mammalian nucleus ambiguus, rather than to the apparent lack of organization found in the mammalian dorsal motor nucleus. Afferents of IX and X entered slightly rostral to the ventral roots of their respective nerves and descended in two tracts. The majority entered the tractus solitarius and descended in a medial position to cervical spinal cord. A portion of the afferents from the vagus nerve crossed the midline in the lower myelencephalon just dorsal to the central canal and ascended a short distance on the contralateral side. Within the solitary tract, vagal afferents were located in a ventrolateral position as they descended to below the obex. Glossopharyngeal afferents filled the remainder of the tract. A smaller portion of afferents from both IX and X did not enter the solitary tract but descended in the spinal tract of V and the dorsolateral funiculus of the spinal cord (Lissauer's tract) to thoracic levels. Afferents of IX also formed a rostral bundle which extended in the solitary tract to the caudal metencephalon.