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Our laboratory has reported with near-infrared spectroscopy (NIRS) that prefrontal oxygenated-hemoglobin concentration (Oxy-Hb), measured as index of regional cerebral blood flow, increased before and at the onset of arbitrary (i.e., noncued) ergometer exercise in a laboratory environment. In the current study, we hypothesized that naturally occurring over-ground locomotion, despite "very light" motor effort, as indicated by a Borg scale of 8.0 ± 0.3, likewise causes preexercise activation of the prefrontal cortex. Using wireless NIRS, we examined in this study how early and to what extent prefrontal activity changed before the onset of arbitrary walking in 13 subjects. Prefrontal Oxy-Hb increased 2 s before the onset of arbitrary walking, and the increased Oxy-Hb reached a peak at 5 s from walking onset. The preexercise and initial increase in prefrontal Oxy-Hb was absent when over-ground walking was forced to start by cue. The difference in the Oxy-Hb response between arbitrary and cued start, which was considered to be related to central command, became significant 2 s before walking onset, preceding the difference in the heart rate (HR) response by 8 s. This demonstrated a positive relationship with the HR difference in 69% of subjects. Imagery of arbitrary walking was, likewise, able to increase prefrontal oxygenation to the same extent as actual walking. Thus, it is likely that prefrontal oxygenation increases before the onset of naturally occurring walking in daily life, despite "very light" effort. The increased prefrontal oxygenation may contribute at least partly to cardiac adjustment, synchronized with the beginning of motor performance. NEW & NOTEWORTHY We found using wireless near-infrared spectroscopy that prefrontal oxygenation increased before the onset of arbitrary over-ground walking, whereas the preexercise increase was absent when walking was suddenly started by cue. The difference in prefrontal oxygenation between start modes (considered related to central command) preceded heart rate response variances and demonstrated a positive relationship with the difference in heart rate. The central command-related prefrontal activity may contribute to cardiac adjustment, synchronized with the beginning of over-ground walking.

We examined whether the cerebrum is essential for producing the rapid autonomic adjustment at the onset of spontaneous overground locomotion. Renal sympathetic nerve activity (RSNA), mean arterial pressure (MAP), heart rate (HR), and electromyogram of the forelimb triceps brachialis were measured when freely moving, decerebrate cats spontaneously produced overground locomotion, supporting body weight. Decerebration was performed at the level of the precollicular-premammillary body. RSNA increased 95 +/- 14 impulses/s (68 +/- 10% of baseline value) at the onset of spontaneous locomotion, which was followed by rises in MAP and HR (7 +/- 1 mmHg and 18 +/- 2 beats/min, respectively). Concomitantly with the MAP rise, RSNA declined toward control values and then increased again during the subsequent period of locomotion. The same rapid increase in RSNA at the onset of locomotion was observed after sinoaortic denervation and vagotomy. It is concluded that some central site(s), other than the cerebrum and the rostral part of the diencephalon, can generate the centrally induced autonomic adjustment at the onset of spontaneous overground locomotion, which is independent of arterial baroreceptor and vagal afferents.

To examine whether the cerebrum is essential for producing the rapid cardiovascular adjustment at the beginning of overground locomotion, we examined heart rate (HR), mean arterial blood pressure (MAP), and integrated electromyogram (iEMG) of the forelimb triceps brachialis muscle in freely moving decerebrate cats during locomotion. Two to four days after decerebration surgery performed at the level of the precollicular-premammillary body, the animals spontaneously produced coordinated overground locomotion, supporting body weight. HR began to increase immediately before the onset of iEMG, and MAP began to rise almost simultaneously with the iEMG onset. Their increases in HR and MAP (24 +/- 3 beats/min and 22 +/- 4 mmHg) were sustained during locomotion. Sinoaortic denervation (SAD) did not affect the abrupt changes in HR and MAP at the beginning of locomotion (0-4 s from the onset of iEMG), whereas SAD had a contrasting effect during the subsequent period, a decrease in the HR response (9 +/- 1 beats/min) and an increase in the MAP response (30 +/- 3 mmHg). These results suggest that the cerebrum and the rostral part of the diencephalon are not essential for producing the rapid cardiovascular adjustment at the beginning of spontaneous overground locomotion. The arterial baroreflex does not contribute to this rapid adjustment but plays an important role in regulating the cardiovascular responses during the later period of spontaneous locomotion.

In walking cats decerebrated at the premammillary level, single neurone activity of Purkinje cells (P-cells) with long corticofugal axons was recorded in the cerebellar vermis. The P-cells (N = 145) were identified as they showed spontaneous simple and complex spikes and also antidromic activation from Deiters' nucleus. These P-cells were classified into 6 groups according to the receptive fields of the climbing fibre responses (CFRs) which were evoked by electrical stimulation in each limb at the radial and sciatic nerve bundles. One group designated as forelimb units received the CFRs from both forelimbs and from neither hindlimb. According to previous studies, this group of P-cells is thought to make inhibitory connections with Deiters neurones projecting to the ipsilateral cervicothoracic spinal cord. For the forelimb units, two types of discharge patterns for simple spikes were found in relation to limb movements during locomotion. Type I cells showed one peak in their firing rate in the late swing (E1) or early stance (E2) phase of the ipsilateral forelimb. Type II cells showed two peaks and two valleys during one step cycle: one peak was in the E1 phase, the other in the late stance (E3) or early swing (F) phase; each of the two valleys followed the peak. Complex spikes of the forelimb units occurred more frequently in the E1 phase than during the other phases. The increased activity of simple and complex spikes of the forelimb units in the E2 phase is suggested to have a functional significance in preparing the appropriate floor reaction forces that appear upon touchdown on the ipsilateral forelimb.

During locomotion of decerebrate and awake walking cats, perturbation (mechanical tap) was applied to the paw dorsum of the left forelimb (LF), and the responses of both forelimbs were recorded cinematographically and electromyographically (EMG). When the tap was applied during the LF stance phase, the duration of the ongoing LF stance was shortened by 10%; in the right forelimb (RF), the duration of the concomitant swing was shortened by 32%. A tap during the LF swing phase prolonged the duration of the ongoing LF swing phase and the concomitant RF stance phase by 55 and 15%, respectively. Analysis of RF joint angle excursions showed that the shortening of the RF swing phase was related mainly to acceleration of extension movement in the late swing phase; the prolongation of the RF stance phase was related to prolonged extension movement in the late stance phase. While EMG activities were relevant to these limb movements, a notable observation was that, by tapping the LF during the LF stance phase, EMG activity in the RF extensor started well before onset of the elbow extension movement to place down the limb; without the tap, the extensor activity started shortly after onset of the extension. Closely related to changes in phase durations of each forelimb, the period of bisupport phase where both forelimbs were in stance, was retained for more than 40% of that of unperturbed steps, even when the RF or LF made the first touchdown after the tap. The rostrocaudal level at RF touchdown after the tap was comparable to unperturbed steps. These findings on interlimb relation suggest that neural control ensures coordinated movements between symmetric limbs during locomotion.

The effects of mechanical stimulation (tap) on single unit activity of Deiter's neurons were analysed in walking cats decerebrated at the premammillary level. Deiters' neurons projecting to the ipsilateral cervical, but not to the lumbosacral, spinal cord (C-Deiters' neurons) were identified by antidromic activation, cerebellar stimulation, and localization of the neurons. During each unperturbed cycle of quadrupedal locomotion, most C-Deiters' neurons showed two frequency modulation peaks in their impulse discharges: one (A peak) in the late swing (E1) or the early stance (E2) phase, the other (B peak) in the late stance (E3) or the early swing (F) phase, of the ipsilateral forelimb. The A peak started to rise shortly before the ipsilateral forelimb was placed. When mechanical perturbation was applied during locomotion to the paw dorsum of the left forelimb (LF) in its stance phase, the ongoing LF stance phase shortened and the simultaneous swing phase of the right forelimb (RF) shortened. Accordingly, in the RF, extensor activity in the swing phase to place down the limb occurred earlier than in unperturbed step cycles. The same LF tap induced a marked enhancement of impulse discharges in C-Deiters' neurons on the right side (with a magnitude of 20-100 imp/s, and the shortest latency of 25 ms). This enhancement was more pronounced than that induced when the perturbation was applied to the LF during its swing phase. The latency manifested a close time relation to the RF extensor activity supporting the postulate that the increased C-Deiters' activity in the RF swing phase contributes to the earlier onset of RF extensor activity which plays an important role in maintaining alternating footfalls after perturbation.

During the locomotion of cats which had been decerebrated at the precollicular and premammillary level, mechanical perturbations (taps of 50-550 g wt.) were applied to the paw dorsum of the left forelimb. Purkinje cells were recorded from the vermis of the cerebellar anterior lobe, and those connected to Deiters' neurons controlling the right forelimb were identified by antidromic and orthodromic stimuli. Taps on the left forelimb induced in these Purkinje cells two types of responses; I-type is a depression of simple spike discharge, often preceded by a brief phase of facilitation, and E-type is entirely a facilitation of simple spike discharge. Complex spikes, representing activation through climbing fiber afferents, were frequently evoked by the taps in both I- and E-types. The I-type depression in Purkinje cells closely corresponds to the previously reported facilitation in Deiters' neurons and forelimb extensor muscles, suggesting that the interlimb coordination during cat's locomotion is effected by linked activity of vermal Purkinje cells and Deiters' neurons.