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      The flexion synergy, mother of all synergies and father of new models of gait

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          Abstract

          Recently there has been a growing interest in the modular organization of leg movements, in particular those related to locomotion. One of the basic modules involves the flexion of the leg during swing and it was shown that this module is already present in neonates (Dominici et al., 2011). In this paper, we question how these finding build upon the original work by Sherrington, who proposed that the flexor reflex is the basic building block of flexion during swing phase. Similarly, the relation between the flexor reflex and the withdrawal reflex modules of Schouenborg and Weng ( 1994) will be discussed. It will be argued that there is large overlap between these notions on modules and the older concepts of reflexes. In addition, it will be shown that there is a great flexibility in the expression of some of these modules during gait, thereby allowing for a phase-dependent modulation of the appropriate responses. In particular, the end of the stance phase is a period when the flexor synergy is facilitated. It is proposed that this is linked to the activation of circuitry that is responsible for the generation of locomotor patterns (CPG, “central pattern generator”). More specifically, it is suggested that the responses in that period relate to the activation of a flexor burst generator. The latter structure forms the core of a new asymmetric model of the CPG. This activation is controlled by afferent input (facilitation by a broad range of afferents, suppression by load afferent input). Meanwhile, many of these physiologic features have found their way in the control of very flexible walking bipedal robots.

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          Locomotor primitives in newborn babies and their development.

          How rudimentary movements evolve into sophisticated ones during development remains unclear. It is often assumed that the primitive patterns of neural control are suppressed during development, replaced by entirely new patterns. Here we identified the basic patterns of lumbosacral motoneuron activity from multimuscle recordings in stepping neonates, toddlers, preschoolers, and adults. Surprisingly, we found that the two basic patterns of stepping neonates are retained through development, augmented by two new patterns first revealed in toddlers. Markedly similar patterns were observed also in the rat, cat, macaque, and guineafowl, consistent with the hypothesis that, despite substantial phylogenetic distances and morphological differences, locomotion in several animal species is built starting from common primitives, perhaps related to a common ancestral neural network.
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            Circuits controlling vertebrate locomotion: moving in a new direction.

            Neurobiologists have long sought to understand how circuits in the nervous system are organized to generate the precise neural outputs that underlie particular behaviours. The motor circuits in the spinal cord that control locomotion, commonly referred to as central pattern generator networks, provide an experimentally tractable model system for investigating how moderately complex ensembles of neurons generate select motor behaviours. The advent of novel molecular and genetic techniques coupled with recent advances in our knowledge of spinal cord development means that a comprehensive understanding of how the motor circuitry is organized and operates may be within our grasp.
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              Organization of mammalian locomotor rhythm and pattern generation.

              Central pattern generators (CPGs) located in the spinal cord produce the coordinated activation of flexor and extensor motoneurons during locomotion. Previously proposed architectures for the spinal locomotor CPG have included the classical half-center oscillator and the unit burst generator (UBG) comprised of multiple coupled oscillators. We have recently proposed another organization in which a two-level CPG has a common rhythm generator (RG) that controls the operation of the pattern formation (PF) circuitry responsible for motoneuron activation. These architectures are discussed in relation to recent data obtained during fictive locomotion in the decerebrate cat. The data show that the CPG can maintain the period and phase of locomotor oscillations both during spontaneous deletions of motoneuron activity and during sensory stimulation affecting motoneuron activity throughout the limb. The proposed two-level CPG organization has been investigated with a computational model which incorporates interactions between the CPG, spinal circuits and afferent inputs. The model includes interacting populations of spinal interneurons and motoneurons modeled in the Hodgkin-Huxley style. Our simulations demonstrate that a relatively simple CPG with separate RG and PF networks can realistically reproduce many experimental phenomena including spontaneous deletions of motoneuron activity and a variety of effects of afferent stimulation. The model suggests plausible explanations for a number of features of real CPG operation that would be difficult to explain in the framework of the classical single-level CPG organization. Some modeling predictions and directions for further studies of locomotor CPG organization are discussed.
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                Author and article information

                Journal
                Front Comput Neurosci
                Front Comput Neurosci
                Front. Comput. Neurosci.
                Frontiers in Computational Neuroscience
                Frontiers Media S.A.
                1662-5188
                13 March 2013
                2013
                : 7
                : 14
                Affiliations
                [1] 1Department of Kinesiology, KU Leuven Heverlee, Belgium
                [2] 2Department of Research, Sint Maartenskliniek Nijmegen, Netherlands
                [3] 3Department of Mechanical Engineering, KU Leuven Heverlee, Belgium
                Author notes

                Edited by: Yuri P. Ivanenko, IRCCS Fondazione Santa Lucia, Italy

                Reviewed by: Yuri P. Ivanenko, IRCCS Fondazione Santa Lucia, Italy; Francesca Sylos Labini, IRCCS Santa Lucia Foundation, Italy

                *Correspondence: Jacques Duysens, Department of Kinesiology, Tervuursevest 101 - bus 01500, BE-3001 Heverlee, Belgium. e-mail: jacques.duysens@ 123456faber.kuleuven.be
                Article
                10.3389/fncom.2013.00014
                3595503
                23494365
                d948ffbf-2616-48f5-a9be-30c4aabf69f5
                Copyright © 2013 Duysens, De Groote and Jonkers.

                This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in other forums, provided the original authors and source are credited and subject to any copyright notices concerning any third-party graphics etc.

                History
                : 21 December 2012
                : 20 February 2013
                Page count
                Figures: 2, Tables: 0, Equations: 0, References: 106, Pages: 9, Words: 8555
                Categories
                Neuroscience
                Review Article

                Neurosciences
                flexion reflex,local sign,reflex modules,synergy,central pattern generator,gait,forward model

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