During overground or treadmill walking, the stance phase and cycle durations are reduced as speed increases, whereas swing phase duration remains relatively invariant. When the speed of the left and right sides is unequal, as is the case during split-belt locomotion or when walking along a circular path, adjustments in stance and swing phases are observed, which could alter the phase/cycle duration relationships. Here, we tested this hypothesis in the left and right hindlimbs of four intact and two chronic spinal-transected adult cats during tied-belt (i.e., equal left and right speeds) and split-belt (i.e., unequal left and right speeds) walking. During split-belt walking, one side (i.e., constant limb) walked at a constant speed while the other side (varying limb) varied its speed from 0.3 to 1.0 m/s. We show that the phase/cycle duration relationships differed in both hindlimbs concurrently during split-belt walking. Specifically, the slope of the phase/cycle duration relationships for the stance/extension phase increased in the varying limb from tied-belt to split-belt walking, whereas that of the swing/flexion phase decreased. In contrast, in the constant limb, the slope of the phase/cycle duration relationships for the stance/extension phase decreased, whereas that of the swing/flexion phase increased. The results were qualitatively similar in intact and spinal-transected cats, indicating that the modulation was mediated within the spinal cord. In conclusion, we propose that neuronal networks within the spinal cord that control left and right hindlimb locomotion can differentially and simultaneously modulate phase variations when the two sides walk at different speeds.

Frigon A, D'Angelo G, Thibaudier Y, Hurteau MF, Telonio A, Kuczynski V, Dambreville C. Speed-dependent modulation of phase variations on a step-by-step basis and its impact on the consistency of interlimb coordination during quadrupedal locomotion in intact adult cats. J Neurophysiol 111: 1885-1902, 2014. First published February 12, 2014 doi:10.1152/jn.00524.2013.-It is well established that stance duration changes more than swing duration for a given change in cycle duration. Small variations in cycle duration are also observed at any given speed on a step-by-step basis. To evaluate the step-bystep effect of speed on phase variations, we measured the slopes of the linear regressions between the phases (i.e., stance, swing) and cycle duration during individual episodes at different treadmill speeds in five adult cats. We also determined the pattern of dominance, defined as the phase that varies most with cycle duration. We found a significant effect of speed on hindlimb phase variations, with significant differences observed between the slowest speed of 0.3 m/s compared with faster speeds. Moreover, although patterns of phase dominance were primarily stance/extensor dominated at the slowest speeds, as speed increased the patterns were increasingly categorized as covarying, whereby both stance/extensor and swing/flexor phases changed in approximately equal proportion with cycle duration. Speed significantly affected the relative duration of support periods as well as interlimb phasing between homolateral and diagonal pairs of limbs but not between homologous pairs of limbs. Speed also significantly affected the consistency of interlimb coordination on a step-by-step basis, being less consistent at the slowest speed of 0.3 m/s compared with faster speeds. We found a strong linear relationship between hindlimb phase variations and the consistency of interlimb coordination. Therefore, results show that phase variations on a step-by-step basis are modulated by speed, which appears to influence the consistency of interlimb coordination. locomotion; speed; phase variations; interlimb coordination DURING OVERGROUND OR TREADMILL locomotion, the cycle can be broadly divided into stance and swing phases. Numerous studies have shown that cycle duration is reduced as speed increases, which is accomplished primarily by a reduction in the duration of the stance phase, while the duration of the swing phase is much less affected (reviewed in Frigon 2012;Gossard et al. 2011). This can be demonstrated by plotting the durations of the stance and swing phases as a function of cycle duration and measuring the slopes of the linear regressions (Halbertsma 1983). In most terrestrial walking species, including insects, birds, rodents, reptiles, dogs, cats, macaques, and humans, the slope of the regression between the stance phase and cycle duration (r STA ) is steeper than the slope of the regression between the swing phase and cycle duration (r SW ) (Arshavskii et al.

Whole animal pressor responses are blunted during pregnancy; yet, uterine arteries, paradoxically, become significantly more sensitive to the constrictor effects of phenylephrine (PE). The objectives herein were to investigate 1) the regional variation (uterine vs. mesenteric arteries) in dose-lumen diameter relationship to alpha-adrenergic stimulation during pregnancy, and 2) the selectivity of these sensitivity shifts for this pathway (PE vs. KCl). Lumen diameter was measured in isolated, pressurized (50 mmHg) arterial segments from age-matched virgin (nonpregnant; NP) and late pregnant (LP; days 19-20) Sprague-Dawley rats. Uterine arcuate vs. mesenteric arteries from NP rats were equally sensitive to either vasoconstrictor. Arcuate arteries from LP rats, however, were 4.5-fold more sensitive to PE (P < 0.01) compared with those from NP controls. Furthermore, diameter curves became superimposed at [PE] > or = 0.1 microM, even though initial diameter of arcuate arteries from LP rats was significantly larger (P < 0.001). Conversely, mesenteric arteries from LP rats were three-fold less sensitive to PE (P < 0.02), and the diameter curve displayed a corresponding parallel rightward shift. Pregnancy did not affect the sensitivity to KCl depolarization in either arcuate or mesenteric arteries. The percent reduction in lumen diameter to the maximum [KCl] was significantly decreased only in arcuate arteries from LP rats (P < 0.001). Thus, during pregnancy, divergent constrictor responses specific to alpha-adrenergic stimulation occur in resistance arteries from the uterine vs. splanchnic circulations. Consequently, concentrations of PE that are subthreshold in NP uterine arteries can elicit large changes in lumen diameter and thereby have a pronounced effect on uterine vascular resistance in the pregnant state.

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