Brainstem control of head movements during orienting; organization of the premotor circuits

When an object appears in the visual field, animals orient their head, eyes, and body toward it in a well-coordinated manner (orienting movement). The head movement is a major portion of the orienting movement. Interest in the neural control of head movements in the monkey and human have increased in the 1990's, however, fundamental knowledge about the neural circuits controlling the orienting head movement continues to be based on a large number of experimental studies performed in the cat. Thus, it is crucial now to summarize information that has been clarified in the cat for further advancement in understanding the neural control of head movements in different animal species. The superior colliculus (SC) has been identified as the primary brainstem center controlling the orienting. Its output signal is transmitted to neck motoneurons via two major separate pathways: one through the reticulospinal neurons (RSNs) in the pons and medulla and the other through neurons in Forel's field H (FFH) in the mesodiencephalic junction. The tecto-reticulo-spinal pathway controls orienting chiefly in the horizontal direction, while the tecto-FFH-spinal pathway controls orienting in the vertical direction. In each pathway, a subgroup of neurons functions as premotor neurons for both extraocular and neck motoneurons, while others are specified for each, which allows both coordinated and separate control of eye and head movements. Head movements almost always produce shifts in the center of gravity that might cause postural disturbances. The postural equilibrium may be maintained by transmitting the orienting command to the limb segments via descending axons of the reticulospinal and long propriospinal neurons. The SC and brainstem relay neurons receive descending inputs from higher order structures such as the cerebral cortex, cerebellum, and basal ganglia. These inputs may serve context-dependent control of orienting by modulating the activities of the primary brainstem pathways.