Infant stepping: a window to the behaviour of the human pattern generator for walking

2004 ◽  
Vol 82 (8-9) ◽  
pp. 662-674 ◽  
Author(s):  
Jaynie F Yang ◽  
Tania Lam ◽  
Marco Y.C Pang ◽  
Erin Lamont ◽  
Kristin Musselman ◽  
...  
Keyword(s):  
Pattern Generator ◽  
Human Infant ◽  
Cycle Duration ◽  
Step Cycle ◽  
Human Infants ◽  
Sensory Inputs ◽  
Weight Support ◽  
Pattern Generators ◽  
Proprioceptive Inputs ◽  
Insight Into

The aim of this paper is to provide evidence, both published and new, to support the notion that human infants are particularly good subjects for the study of the pattern generator for walking. We and others have shown that stepping can be initiated by sensory input from the legs or by general heightened excitability of the infant. New results are presented here to suggest that weight support through the feet and rapid extension of the legs are important proprioceptive inputs to initiate stepping. Our previous work has shown that infants can step at many different speeds when supported on a treadmill. The step cycle duration shortens as the speed increases, with the changes coming largely from the stance phase, just as in most other terrestrial animals. Moreover, we have shown that infants will step in all directions. Regardless of the direction of stepping, the step cycle changes in the same way with walking speed, suggesting the circuitry that controls different directions of walking share common elements. We have also shown that infant stepping is highly organized. Sensory inputs, whether proprioceptive or touch, are gated in a functional way so that only important sensory inputs generate a response. For example, touch to the lateral surface of the foot elicits a response only in sideways walking, and only in the leading limb. New data is presented here to show that the pattern generators from each limb can operate somewhat independently. On a split-belt treadmill with the 2 belts running at different speeds or in different directions, the legs showed considerable independence in behaviour. Yet, the pattern generators on each side interact to ensure that swing phase does not occur at the same time. These studies have provided insight into the organization of the pattern generator for walking in humans. It will be interesting in the future to study how maturation of the descending tracts changes walking behaviour to allow independent bipedal walking.Key words: locomotion, central pattern generator, human, infant.

Could Different Directions of Infant Stepping Be Controlled by the Same Locomotor Central Pattern Generator?

2000 ◽  
Vol 83 (5) ◽  
pp. 2814-2824 ◽  
Author(s):  
Tania Lamb ◽  
Jaynie F. Yang
Keyword(s):  
Central Pattern Generator ◽  
Muscle Activation ◽  
Pattern Generator ◽  
Swing Phase ◽  
Smooth Transition ◽  
Cycle Duration ◽  
Human Infants ◽  
Activation Patterns ◽  
Muscle Activation Patterns ◽  
The Relationship

This study examined the idea of whether the same central pattern generator (CPG) for locomotion can control different directions of walking in humans. Fifty-two infants, aged 2–11 mo, were tested. Infants were supported to walk on a treadmill at a variety of speeds. If forward stepping was elicited, stepping in the other directions (primarily sideways and backward) was attempted. The orientation of the infant on the treadmill belt determined the direction of stepping. In some infants, we also attempted to obtain a smooth transition from one direction to another by gradually changing the orientation of the infant during a stepping sequence. Limb segment motion and surface electromyography from the muscles of the lower limb were recorded. Most infants who showed sustained forward walking also could walk in all other directions. Thirty-three of 34 infants tested could step sideways. The success of eliciting backward stepping was 69%. Most of the infants who did not meet our backward stepping criteria did, however, make stepping movements. The different directions of stepping had similar responses to changes in treadmill speed. The relationship between stance and swing phase durations and cycle duration were the same regardless of the direction of stepping across a range of speeds. Some differences were noted in the muscle activation patterns during different directions of walking. For example, the hamstrings were much more active during the swing phase of backward walking compared with forward walking. The quadriceps was more active in the trailing leg during sideways walking. In some infants, we were able to elicit stepping along a continuum of directions. We found no discrete differences in either the electromyographic patterns or the temporal parameters of stepping as the direction of stepping was gradually changed. The results support the idea that the same locomotor CPG controls different directions of stepping in human infants. The fact that most infants were able to step in all directions, the similarity in the response to speed changes, and the absence of any discrete changes as the direction of stepping was changed gradually are all consistent with this hypothesis.

Stimulation of the Parapyramidal Region of the Neonatal Rat Brain Stem Produces Locomotor-Like Activity Involving Spinal 5-HT7 and 5-HT2A Receptors

2005 ◽  
Vol 94 (2) ◽  
pp. 1392-1404 ◽  
Author(s):  
Jun Liu ◽  
Larry M. Jordan
Keyword(s):  
Brain Stem ◽  
Pattern Generator ◽  
Neonatal Rat ◽  
Cycle Duration ◽  
Receptor Antagonists ◽  
Step Cycle ◽  
Output Stage ◽  
Discrete Population ◽  
First Time ◽  
Neonatal Rat Brain

Locomotion can be induced in rodents by direct application 5-hydroxytryptamine (5-HT) onto the spinal cord. Previous studies suggest important roles for 5-HT7 and 5-HT2A receptors in the locomotor effects of 5-HT. Here we show for the first time that activation of a discrete population of 5-HT neurons in the rodent brain stem produces locomotion and that the evoked locomotion requires 5-HT7 and 5-HT2A receptors. Cells localized in the parapyramidal region (PPR) of the mid-medulla produced locomotor-like activity as a result of either electrical or chemical stimulation, and PPR-evoked locomotor-like activity was blocked by antagonists to 5-HT2A and 5-HT7 receptors located on separate populations of neurons concentrated in different rostro-caudal regions. 5-HT7 receptor antagonists blocked locomotor-like activity when applied above the L3 segment; 5-HT2A receptor antagonists blocked locomotor-like activity only when applied below the L2 segment. 5-HT7 receptor antagonists decreased step cycle duration, consistent with an action on neurons involved in the rhythm-generating function of the central pattern generator (CPG) for locomotion. 5-HT2A antagonists reduced the amplitude of ventral root activity with only small effects on step cycle duration, suggesting an action directly on cells involved in the output stage of the pattern generator for locomotion, including motoneurons and premotor cells. Experiments with selective antagonists show that dopaminergic (D1, D2) and noradrenergic (α1, α2) receptors are not critical for PPR-evoked locomotor-like activity.

Effects of Intrathecal α1- and α2-Noradrenergic Agonists and Norepinephrine on Locomotion in Chronic Spinal Cats

1998 ◽  
Vol 79 (6) ◽  
pp. 2941-2963 ◽  
Author(s):  
Connie Chau ◽  
Hugues Barbeau ◽  
Serge Rossignol
Keyword(s):  
Time Course ◽  
Spinal Injury ◽  
Emg Activity ◽  
Cycle Duration ◽  
Flexor Muscle ◽  
Step Cycle ◽  
Differential Effects ◽  
Cutaneous Reflex ◽  
Weight Support ◽  
Locomotor Deficits

Chau, Connie, Hugues Barbeau, and Serge Rossignol. Effects of intrathecal α1- and α2-noradrenergic agonists and norepinephrine on locomotion in chronic spinal cats. J. Neurophysiol. 79: 2941–2963, 1998. Noradrenergic drugs, acting on α adrenoceptors, have been found to play an important role in the initiation and modulation of locomotor pattern in adult cats after spinal cord transection. There are at least two subtypes of α adrenoceptors, α1 and α2 adrenoceptors. The aim of this study was to investigate the effects of selective α1 and α2 agonists in the initiation and modulation of locomotion in adult chronic cats in the early and late stages after complete transection at T13. Five cats, chronically implanted with an intrathecal cannula and electromyographic (EMG) electrodes were used in this study. Noradrenergic drugs including α2 agonists (clonidine, tizanidine, and oxymetazoline) and an antagonist, yohimbine, one α1 agonist (methoxamine), and a blocker, prazosin, as well as norepinephrine were injected intrathecally. EMG activity synchronized to video images of the hindlimbs were recorded before and after each drug injection. The results show differential effects of α1 and α2 agonists in the initiation of locomotion in early spinal cats (i.e., in the first week or so when there is no spontaneous locomotion) and in the modulation of locomotion and cutaneous reflexes in the late-spinal cats (i.e., when cats have recovered spontaneous locomotion). In early spinal cats, all three α2 agonists were found to initiate locomotion, although their action had a different time course. The α1 agonist methoxamine induced bouts of nice locomotor activity in three spinal cats some hours after injection but only induced sustained locomotion in one cat in which the effects were blocked by the α1 antagonist prazosin. In late spinal cats, although α2 agonists markedly increased the cycle duration and flexor muscle burst duration and decreased the weight support or extensor activity (effects blocked by an α2 antagonist, yohimbine), α1 agonist increased the weight support and primarily the extensor activity of the hindlimbs without markedly changing the timing of the step cycle. Although α2 agonists, especially clonidine, markedly reduced the cutaneous excitability and augmented the foot drag, the α1 agonist was found to increase the cutaneous reflex excitability. This is in line with previously reported differential effects of activation of the two receptors on motoneuron excitability and reflex transmission. Noradrenaline, the neurotransmitter itself, increased the cycle duration and at the same time retained the cutaneous excitability, thus exerting both α1 and α2 effects. This work therefore suggests that different subclasses of noradrenergic drugs could be used to more specifically target aspects of locomotor deficits in patients after spinal injury or diseases.

Sensory regulation of quadrupedal locomotion: a top-down or bottom-up control system?

2012 ◽  
Vol 108 (3) ◽  
pp. 709-711 ◽  
Author(s):  
Yann Thibaudier ◽  
Marie-France Hurteau
Keyword(s):  
Control System ◽  
Central Pattern Generators ◽  
Top Down ◽  
Bottom Up ◽  
Sensory Inputs ◽  
Sensory Regulation ◽  
Before And After ◽  
Pattern Generators

Propriospinal pathways are thought to be critical for quadrupedal coordination by coupling cervical and lumbar central pattern generators (CPGs). However, the mechanisms involved in relaying information between girdles remain largely unexplored. Using an in vitro spinal cord preparation in neonatal rats, Juvin and colleagues ( Juvin et al. 2012 ) have recently shown sensory inputs from the hindlimbs have greater influence on forelimb CPGs than forelimb sensory inputs on hindlimb CPGs, in other words, a bottom-up control system. However, results from decerebrate cats suggest a top-down control system. It may be that both bottom-up and top-down control systems exist and that the dominance of one over the other is task or context dependent. As such, the role of sensory inputs in controlling quadrupedal coordination before and after injury requires further investigation.

Behavior-Dependent Activities of a Central Pattern Generator in Freely Behaving Lymnaea stagnalis

1997 ◽  
Vol 78 (6) ◽  
pp. 3415-3427 ◽  
Author(s):  
Rene F. Jansen ◽  
Anton W. Pieneman ◽  
Andries ter Maat
Keyword(s):  
Electrical Activity ◽  
Central Pattern Generator ◽  
Lymnaea Stagnalis ◽  
Central Pattern Generators ◽  
Pattern Generator ◽  
Egg Laying ◽  
Buccal Ganglion ◽  
Level Of Activity ◽  
Freely Behaving ◽  
Pattern Generators

Jansen, Rene F., Anton W. Pieneman, and Andries ter Maat. Behavior-dependent activities of a central pattern generator in freely behaving Lymnaea stagnalis. J. Neurophysiol. 78: 3415–3427, 1997. Cyclic or repeated movements are thought to be driven by networks of neurons (central pattern generators) that are dynamic in their connectivity. During two unrelated behaviors (feeding and egg laying), we investigated the behavioral output of the buccal pattern generator as well as the electrical activity of a pair of identified interneurons that have been shown to be involved in setting the level of activity of this pattern generator (PG). Analysis of the quantile plots of the parameters that describe the behavior (movements of the buccal mass) reveals that during egg laying, the behavioral output of the PG is different compared with that during feeding. Comparison of the average durations of the different parts of the buccal movements showed that during egg laying, the duration of one specific part of buccal movement is increased. Correlated with these changes in the behavioral output of the PG were changes in the firing rate of the cerebral giant neurons (CGC), a pair of interneurons that have been shown to modulate the activity of the PG by means of multiple synaptic contacts with neurons in the buccal ganglion. Interval- and autocorrelation histograms of the behavioral output and CGC spiking show that both the PG output and the spiking properties of the CGCs are different when comparing egg-laying animals with feeding animals. Analysis of the timing relations between the CGCs and the behavioral output of the PG showed that both during feeding and egg laying, the electrical activity of the CGCs is largely in phase with the PG output, although small changes occur. We discuss how these results lead to specific predictions about the kinds of changes that are likely to occur when the animal switches the PG from feeding to egg laying and how the hormones that cause egg laying are likely to be involved.

scholarly journals Statistical learning in domestic chicks is modulated by strain and sex

2020 ◽  
Vol 10 (1) ◽  
Author(s):  
Chiara Santolin ◽  
Orsola Rosa-Salva ◽  
Bastien S. Lemaire ◽  
Lucia Regolin ◽  
Giorgio Vallortigara
Keyword(s):  
Sex Differences ◽  
Statistical Learning ◽  
Comparative Research ◽  
Learning Task ◽  
Human Infants ◽  
Sensory Inputs ◽  
Automated Tracking ◽  
Domestic Chicks ◽  
Continuous Stream

Abstract Statistical learning is a key mechanism for detecting regularities from a variety of sensory inputs. Precocial newborn domestic chicks provide an excellent model for (1) exploring unsupervised forms of statistical learning in a comparative perspective, and (2) elucidating the ecological function of statistical learning using imprinting procedures. Here we investigated the role of the sex of the chicks in modulating the direction of preference (for familiarity or novelty) in a visual statistical learning task already employed with chicks and human infants. Using both automated tracking and direct human coding, we confirmed chicks’ capacity to recognize the presence of a statistically defined structure underlying a continuous stream of shapes. Using a different chicken strain than previous studies, we were also able to highlight sex differences in chicks’ propensity to approach the familiar or novel sequence. This could also explain a previous failure to reveal statistical learning in chicks which sex was however not determined. Our study confirms chicks’ ability to track visual statistics. The pivotal role of sex in determining familiarity or novelty preferences in this species and the interaction with the animals’ strain highlight the importance to contextualize comparative research within the ecology of each species.

Comparison of the Effect of Intrathecal Administration of Clonidine and Yohimbine on the Locomotion of Intact and Spinal Cats

2001 ◽  
Vol 85 (6) ◽  
pp. 2516-2536 ◽  
Author(s):  
Nathalie Giroux ◽  
Tomás A. Reader ◽  
Serge Rossignol
Keyword(s):  
Acid Methyl Ester ◽  
Intrathecal Administration ◽  
Cycle Duration ◽  
Descending Pathways ◽  
Step Cycle ◽  
Locomotor Pattern ◽  
Before And After ◽  
Spontaneous Expression ◽  
High Doses ◽  
Treadmill Locomotion

Several studies have shown that noradrenergic mechanisms are important for locomotion. For instance, L-dihydroxyphenylalanine (L-DOPA) can initiate “fictive” locomotion in immobilized acutely spinalized cats and α2-noradrenergic agonists, such as 2,6,-dichloro- N-2-imidazolidinylid-enebenzenamine (clonidine), can induce treadmill locomotion soon after spinalization. However, the activation of noradrenergic receptors may be not essential for the basic locomotor rhythmicity because chronic spinal cats can walk with the hindlimbs on a treadmill in the absence of noradrenergic stimulation because the descending pathways are completely severed. This suggests that locomotion, in intact and spinal conditions, is probably expressed and controlled through different neurotransmitter mechanisms. To test this hypothesis, we compared the effect of the α2 agonist, clonidine, and the antagonist (16α, 17α)-17-hydroxy yohimbine-16-carboxylic acid methyl ester hydrochloride (yohimbine), injected intrathecally at L3–L4before and after spinalization in the same cats chronically implanted with electrodes to record electromyograms (EMGs). In intact cats, clonidine (50–150 μg/100 μl) modulated the locomotor pattern slightly causing a decrease in duration of the step cycle accompanied with some variation of EMG burst amplitude and duration. In the spinal state, clonidine could trigger robust and sustained hind limb locomotion in the first week after the spinalization at a time when the cats were paraplegic. Later, after the spontaneous recovery of a stable locomotor pattern, clonidine prolonged the cycle duration, increased the amplitude and duration of flexor and extensor bursts, and augmented the foot drag at the onset of swing. In intact cats, yohimbine at high doses (800–1600 μg/100 μl) caused major walking difficulties characterized by asymmetric stepping, stumbling with poor lateral stability, and, at smaller doses (400 μg/100 μl), only had slight effects such as abduction of one of the hindlimbs and the turning of the hindquarters to one side. After spinalization, yohimbine had no effect even at the largest doses. These results indicate that, in the intact state, noradrenergic mechanisms probably play an important role in the control of locomotion since blocking the receptors results in a marked disruption of walking. In the spinal state, although the receptors are still present and functional since they can be activated by clonidine, they are seemingly not critical for the spontaneous expression of spinal locomotion since their blockade by yohimbine does not impair spinal locomotion. It is postulated therefore that the expression of spinal locomotion must depend on the activation of other types of receptors, probably related to excitatory amino acids.

Intrinsic and Extrinsic Modulation of a Single Central Pattern Generating Circuit

2000 ◽  
Vol 84 (3) ◽  
pp. 1186-1193 ◽  
Author(s):  
Peter T. Morgan ◽  
Ray Perrins ◽  
Philip E. Lloyd ◽  
Klaudiusz R. Weiss
Keyword(s):  
Neural Circuits ◽  
Central Pattern Generators ◽  
Motor Program ◽  
Pattern Generator ◽  
Motor Programs ◽  
Appetitive Behavior ◽  
Protraction Phase ◽  
Pattern Generators ◽  
Rhythmic Behaviors ◽  
Appetitive Behaviors

Intrinsic and extrinsic neuromodulation are both thought to be responsible for the flexibility of the neural circuits (central pattern generators) that control rhythmic behaviors. Because the two forms of modulation have been studied in different circuits, it has been difficult to compare them directly. We find that the central pattern generator for biting in Aplysia is modulated both extrinsically and intrinsically. Both forms of modulation increase the frequency of motor programs and shorten the duration of the protraction phase. Extrinsic modulation is mediated by the serotonergic metacerebral cell (MCC) neurons and is mimicked by application of serotonin. Intrinsic modulation is mediated by the cerebral peptide-2 (CP-2) containing CBI-2 interneurons and is mimicked by application of CP-2. Since the effects of CBI-2 and CP-2 occlude each other, the modulatory actions of CBI-2 may be mediated by CP-2 release. Although the effects of intrinsic and extrinsic modulation are similar, the neurons that mediate them are active predominantly at different times, suggesting a specialized role for each system. Metacerebral cell (MCC) activity predominates in the preparatory (appetitive) phase and thus precedes the activation of CBI-2 and biting motor programs. Once the CBI-2s are activated and the biting motor program is initiated, MCC activity declines precipitously. Hence extrinsic modulation prefacilitates biting, whereas intrinsic modulation occurs during biting. Since biting inhibits appetitive behavior, intrinsic modulation cannot be used to prefacilitate biting in the appetitive phase. Thus the sequential use of extrinsic and intrinsic modulation may provide a means for premodulation of biting without the concomitant disruption of appetitive behaviors.

Effects of Intrathecal Glutamatergic Drugs on Locomotion. II. NMDA and AP-5 in Intact and Late Spinal Cats

2003 ◽  
Vol 90 (2) ◽  
pp. 1027-1045 ◽  
Author(s):  
Nathalie Giroux ◽  
Connie Chau ◽  
Hugues Barbeau ◽  
Tomás A. Reader ◽  
Serge Rossignol
Keyword(s):  
Excitatory Amino Acids ◽  
Early Stage ◽  
Pattern Generator ◽  
Major Effect ◽  
Low Doses ◽  
Receptor Stimulation ◽  
Locomotor Pattern ◽  
Weight Support ◽  
Spontaneous Expression ◽  
Nmda Receptor Blocker

In a previous article, we have shown that, in cats, intrathecal injections of N-methyl-d-aspartate (NMDA) in the first few days after spinalization at T13 do not induce locomotion as in many other spinal preparations. This is in contrast to alpha-2 noradrenergic receptor stimulation, which can trigger locomotion at this early stage. However, it is known that spinal cats do recover spontaneous locomotion in the absence of descending noradrenergic pathways and that the spinal pattern generator must then depend on other neurotransmitters still present in the cord such as excitatory amino acids. In the present paper, therefore we look at the effects of intrathecal NMDA, a glutamatergic agonist, and 2-amino-5-phosphonovaleric acid (AP-5), an NMDA receptor blocker, in both intact and late spinal cats. Low doses of NMDA had no major effect on the locomotor pattern in both intact and late spinal cats. Larger doses of NMDA in the chronic spinal cat initially produced an increase in the general excitability followed by more regular locomotion. AP-5 in intact cats caused a decrease in the amplitude of the flexion reflex and induced a bilateral foot drag as well as some decrease in weight support but it did not prevent locomotion. However, in late spinal cats, the same dose of AP-5 blocked locomotion completely. These results indicate that NMDA receptors may be critical for the spontaneous expression of spinal locomotion. It is proposed that the basic locomotor rhythmicity in cats is NMDA-dependent and that normally this glutamatergic mechanism is modulated by other neurotransmitters, such as 5-HT and NA.

Contribution of the Motor Cortex to the Structure and the Timing of Hindlimb Locomotion in the Cat: A Microstimulation Study

2005 ◽  
Vol 94 (1) ◽  
pp. 657-672 ◽  
Author(s):  
Frédéric Bretzner ◽  
Trevor Drew
Keyword(s):  
Motor Cortex ◽  
Muscle Activity ◽  
Pyramidal Tract ◽  
Cycle Duration ◽  
Step Cycle ◽  
Standard Glass ◽  
Level Of Activity ◽  
Flexor Muscles ◽  
Multijoint Movements ◽  
Stimulation Of

We used microstimulation to examine the contribution of the motor cortex to the structure and timing of the hindlimb step cycle during locomotion in the intact cat. Stimulation was applied to the hindlimb representation of the motor cortex in 34 sites in three cats using either standard glass-insulated microelectrodes (16 sites in 1 cat) or chronically implanted microwire electrodes (18 sites in 2 cats). Stimulation at just suprathreshold intensities with the cat at rest produced multijoint movements at a majority of sites (21/34, 62%) but evoked responses restricted to a single joint, normally the ankle, at the other 13/34 (38%) sites. Stimulation during locomotion generally evoked larger responses than the same stimulation at rest and frequently activated additional muscles. Stimulation at all 34 sites evoked phase-dependent responses in which stimulation in swing produced transient increases in activity in flexor muscles while stimulation during stance produced transient decreases in activity in extensors. Stimulation with long (200 ms) trains of stimuli in swing produced an increased level of activity and duration of flexor muscles without producing changes in cycle duration. In contrast, stimulation during stance decreased the duration of the extensor muscle activity and initiated a new and premature period of swing, resetting the step cycle. Stimulation of the pyramidal tract in two of these three cats as well as in two additional ones produced similar effects. The results show that the motor cortex is capable of influencing hindlimb activity during locomotion in a similar manner to that seen for the forelimb.

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