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THE ipsilateral and contralateral corticocortical connections to the fifth somatosensory area (SV) in the feline cortex were determined from the location of retrogradely labelled cells following a single injection of HRP into SV. The injection was made into physiologically defined components of the body representation in SV. After injection of HRP into the face regions of SV, HRP-labelled cells were located ipsilaterally in areas 6 beta, 3b and 1-2 of the primary somatosensory (SI), in the second somatosensory (SII), third somatosensory (SIII), and fourth somatosensory (SIV) areas, along the ansate sulcus, and in areas 5a and 6a beta of the ipsilateral cortex, as well as in area 1-2 of SI and in SV of the contralateral cortex. On the other hand, after HRP had been injected into the trunk/hindlimb area, HRP-labelled cells were located in areas 3a, 1-2 of SI, in area 5, in SII, in SIII and in SIV of the ipsilateral cortex, as well as in area 1-2 of SI, and in SV of the contralateral cortex. The extent of these interconnections suggests that SV receives multiple sensory inputs and may function to integrate this information.

Forelimb crossed extension reflexes were examined in 22 thalamic cats. These reflexes were elicited either by backward passive movement or by repetitive electrical stimulation of cutaneous and joint afferent nerves in the contralateral forelimb. Single stimulation of the superficial radial nerve evoked two types of reflex responses—early (ER) and late (LR)—from the triceps brachii muscle on the contralateral side. The latencies were about 7 and 16–25 ms, corresponding to the propriospinal (PSR) and spino-bulbo-spinal (SBS) reflexes of the ipsilateral flexor, respectively. Repetitive stimulation of the superficial radial nerve evoked the LR but not the ER. The crossed extension reflex and LR were abolished by lesions of the dorsolateral funiculus of the cervical cord on the side opposite to the recording. The tonic EMG activity, crossed extension reflex and LR in the extensor on the side of lesions were abolished by lesions of the ventrolateral funiculus of the cervical cord. During forelimb stepping, the amplitudes of both ER and LR fluctuated depending on the phase of the step cycle. The ER appeared during a narrow period in the early phase of the stance, whereas the LR was observed during a wide period from the middle of the swing to the middle of the stance. Both responses were absent from the middle of the stance to the middle of the swing. These observations suggest that forelimb crossed extension reflexes involve both spinal and supraspinal (SBS) loop mechanisms, and that these are utilized during stepping, with the latter mechanism in particular playing an important part in the extension phase of the forelimb forward movement.

To analyze the descending pathways of the paralemniscal pontine reticular formation (PLRF), a technique was used for the selective activation of cell bodies by localized injection of putative neurotransmitters in the PLRF. When a small amount (less than 0.1 microliter) of 0.1 M glutamate was injected into the PLRF unilaterally in thalamic cats, the forelimb contralateral (c-forelimb) to the injection was extended, and occasionally the ipsilateral forelimb was flexed. These responses were similar to those obtained by electrical stimulation of the PLRF, but were relatively weaker. Unit spikes of PLRF neurons were increased in frequency following administration of glutamate. The latent periods and durations of increases in spike frequency varied depending on the concentration and quantity of the glutamate solution, and were roughly similar to those of the extensor EMG in the c-forelimb. Since the firing of PLRF neurons preceded the EMG with 11 ms latency, the unit spike of PLRF neurons could be used as a triggering signal to observe a spike triggered averaged EMG response in the extensor muscle of the c-forelimb. Results similar to those with glutamate were observed upon administration of quisqualate, kainate and aspartate. The most effective compound was quisqualate. Application to the PLRF of 1-naphthylacetyl spermine (1-NA-Spm), an analogue of the natural spider toxin JSTX-3 and an antagonist of glutamate, suppressed both the PLRF neuron activity and the extensor EMG of the c-forelimb. These observations suggest that extensor muscles of the forelimb are excited by the contralateral PLRF, perhaps via the crossed reticulospinal tract from the PLRF. PLRF neurons may be activated by glutamate (quisqualate) receptors.

To analyze changes in the excitability of both the spinal cord and brainstem in thalamic cats stepping on a moving treadmill, we examined the cutaneous propriospinal (PSR) and spino-bulbo-spinal (SBS) reflex responses in 20 adult cats. Tracheal cannulation, spinal transection at the T10 segment, and decerebration at the stereotaxic A12 level were performed under ether anesthesia. Immediately after decerebration, the ether was withdrawn. The head was fixed in a stereotaxic device, the T2 spinal process clamped to a metal frame, and the lumbar region suspended by a hammock, with bilateral forelimb contact on the floor of a treadmill. Electrical stimulation was applied to the superficial radial nerve with a cuff electrode, and two reflex responses (PSR and SBS) were recorded from the biceps brachii muscle in the same forelimb. Shortly before the appearance of forelimb stepping, both PSR and SBS reflex responses were elevated in amplitude. During forelimb stepping, the amplitudes of PSR and SBS reflex responses fluctuated depending on the phase of the step cycle. The PSR response was enhanced in the early phase of the swing, whereas the SBS response was elevated during a wider period from the beginning of the stance to the middle of the swing. The SBS response was completely absent in the late phase of the swing. This period corresponded to the transfer from flexion to extension and the appearance of the EMG of the triceps brachii muscle of the same forelimb. The fluctuation of the SBS response during stepping may be produced at the brainstem level, and not the spinal cord level, because the PSR response was enhanced only during narrow periods. The generation of locomotion thus seems to result in an enhancement of excitability of reflex pathways in the spinal cord and particularly in the brainstem.

Summary We recorded the neuronal activities from, stimulated, and produced lesions in, various regions in the brainstem of thalamic cats. Neuronal activities were recorded during stepping on a motor-driven treadmill, and were averaged by using the flexor EMG as the trigger. In wide regions of the brainstem, an averaged response was obtained with similar latency as the ipsilateral flexor EMG; however, in the paralemniscal pontine reticular formation (PLRF) the averaged response preceded the flexor EMG. When repetitive electrical stimulation was applied to the PLRF on one side, coordinated movements of bilateral forelimbs were elicited. The ipsilateral forelimb was flexed and the contralateral forelimb extended. EMG recordings revealed that on the side ipsilateral to the stimulation, flexor EMGs appeared and increased gradually in amplitude, and the extensor EMG was suppressed, while the extensor EMG was increased and no flexor EMG was obtained on the side contralateral to the stimulation. When pulse train stimulation was alternatingly applied to the PLRF on bilateral sides, alternating forelimb movement, as in stepping, was evoked, depending on each stimulation. A small, unilateral lesion in the PLRF reduced forelimb stepping on a motor-driven treadmill, particularly the flexor muscle contraction in the forelimb ipsilateral to the lesion. Bilateral PLRF lesions abolished forelimb stepping on both sides. Unilateral transection of the ventral quadrant at the cervical cord abolished PLRF-elicited flexor EMGs and associated suppression of extensor EMG ipsilateral to the transection; however, the side contralateral to the transection remained intact. These results suggest that the PLRF exerts a patterned control of the bilateral forelimb muscles and plays a role in initiating and maintaining forelimb stepping.

The paralemniscal pontine reticular neurons (PLRF) exhibited burst discharges when thalamic cats stepped on the moving treadmill, during stepping induced by mesencephalic locomotor region (MLR) stimulation, fictive locomotion by MLR stimulation after immobilization and during MLR stimulation after transection at the C2 level. These results imply that MLR stimulation can evoke burst discharge within the brainstem which may obtain alternating limb movements via descending tract to the cervical cord.

To analyze the roles of joint afferents in relation to initiation of forelimb stepping in thalamic cats, we recorded the unit spikes of the cervical dorsal roots, stimulated the joint afferents, and applied local anesthesia to the joint capsule. Almost all of the joint afferents of the shoulder, elbow, wrist and finger adapted slowly and exhibited alternating firing during forelimb stepping. About 45% of the afferents of each joint showed firings as the limb moved from forward to backward. About 44% of the afferents exhibited discharges as the limb moved from backward to forward. The remaining afferents showed firings as the limb moved in both directions. The application of local anesthesia to joints of the shoulder, elbow or wrist resulted in a marked reduction of forelimb stepping. Forelimb stepping was evoked by electric stimulation of the joint capsule, when excitabilities of flexor motoneurons were increased due to muscle stretching. Impulses originating in the joint afferents of the forelimb entered the spinal cord and ascended to the dorsolateral funiculus of the cervical cord, since forelimb stepping was abolished after bilateral transection of this part. Our results indicate that joint afferents may play an important role in the initiation of forelimb stepping in thalamic cats walking on a motor-driven treadmill.