1. The effect of stimulating the contralateral pyramid has been investigated with intracellular recording from 128 long propriospinal neurones (long PNs) in the C3-Th1 segments of the cat. Long PNs were identified by the antidromic activation from the Th13 segment. They were located in laminae VII-VIII of Rexed. Single pyramidal stimulation evoked monosynaptic EPSPs in 15/40 of the long PNs in cats with intact pyramid. In 15 other long PNs, a train of three to four pyramidal stimuli evoked EPSPs with latencies indicating a minimal disynaptic linkage. The remaining 25% of the long PNs lacked mono- or disynaptic pyramidal EPSPs. In a few cases longer latency excitation was observed. 2. The location of the intercalated neurones which mediate the disynaptic pyramidal EPSPs was investigated by making four different lesions of the corticofugal fibres: 1) at the border of the C5 and C6 segments, 2) at the border of the C2 and C3 segments, 3) at the caudal part of the pyramid; three mm rostral to the decussation and 4) at the level of the trapezoid body. Stimulation of the corticofugal fibres was made either rostral to lesion 3 (rPyr) in order to activate neurones in a cortico-bulbospinal pathway or caudal to lesion 3 (cPyr) to activate neurones in a corticospinal pathway. In the former case, in one experiment, stimulation was made in the pyramid between lesions 3 and 4 (double pyramidal lesion). In case of cPyr stimulation, lesions 1 and 2 were added sequentially in order to investigate if the corticospinal excitation was mediated via C3-C4 PNs. All lesions were made mechanically, except lesion 2 which in some of the experiments was performed by reversible cooling. 3. Stimulation in the pyramid rostral to lesion 3 and in between lesions 3 and 4 evoked disynaptic EPSPs in the long PNs, which shows that they were mediated via reticulospinal neurones. Stimulation in cPyr after lesion 3 elicited disynaptic EPSPs, which remained after lesion 1 but were abolished after adding lesion 2. It is concluded that the disynaptic cPyr EPSPs were mediated via intercalated neurones in the C3-C4 segments. 4. When the disynaptic cPyr EPSP was conditioned with a single volley in nucleus ruber and/or in tectum, it was markedly facilitated, especially when the conditioned volley was applied simultaneously with the effective cPyr volley. The results show that the intercalated neurones in the C3-C4 segments receive monosynaptic convergence from cortico-, rubro- and tectospinal fibres. Stimulation in the lateral reticular nucleus (LRN) evoked monosynaptic EPSPs.(ABSTRACT TRUNCATED AT 400 WORDS)