In decerebrate, acutely labyrinthectomized cats we used neck rotation to study the role of direct upper cervical afferents to the cervical enlargement and of cervical and lumbar propriospinal neurons in the tonic neck reflex. Interruption of the dorsal columns between C4 and C5 had no qualitative effect on the dynamics of the reflex although gain usually increased. Direct upper cervical afferents to the cervical enlargement therefore have no unique role in producing the reflex. Many medially located propriospinal neurons in C4 were modulated by neck rotation. About 40% had axons, mostly crossed, that terminated in the cervical enlargement. The others projected more caudally, some as far as L3-L4 or even the lumbar enlargement. For a population of C4 neurons, including propriospinal neurons, we measured the response vector with combinations of roll and pitch stimuli. These vectors ranged from pitch to roll. Many propriospinal neurons in L3-L4, projecting to the lumbosacral enlargement, were also modulated by neck rotation with a variety of response vectors. Some of these neurons had an ascending projection. As in previous experiments, C4 neurons were modulated by neck rotation after spinal transection rostral to the C1 dorsal root entry zone; a wide variety of response vectors was observed. In contrast, almost no modulated L3-L4 neurons were found in the same experiments. The results suggest a role for propriospinal neurons in the tonic neck reflex. They also demonstrate that responses of lumbar neurons to neck rotation are much more dependent on supraspinal pathways than are those of cervical neurons.