Anatomical study of the final common pathway for vocalization in the cat

Vocalization, the nonverbal production of sound, can be elicited in many vertebrates by stimulation in several regions of the limbic system but most easily in the caudal periaqueductal gray (PAG). This study shows that a specific cell group in the lateral part of the caudal PAG and in the tegmentum just lateral to it projects bilaterally to the nucleus retroambiguus (NRA) in the caudal medulla oblongata. Similar but much weaker projections are derived from the dorsal PAG. Neurons in the NRA in turn project via a contralateral pathway through the ventral funiculus of the spinal cord to motoneuronal cell groups, innervating intercostal and abdominal muscles. These projections are stronger on the contralateral side, although at lower thoracic and upper lumbar levels, many fibers recross to terminate in the ipsilateral motoneuronal cell groups. In the brainstem NRA neurons project to the motoneuronal cell groups innervating mouth-opening and perioral muscles as well as to motoneurons innervating the pharynx, soft palate, and tongue, and probably to the larynx. All these muscles are active in vocalization. The present anatomical results, combined with the physiological results of others, indicate that the projections from PAG via NRA to vocalization motoneurons form the final common pathway in vocalization. The role of this pathway in the total framework of emotional behavior is discussed.