Computer simulation study on central pattern generator: from biology to engineering

Central pattern generator (CPG) is a neuronal circuit in the nervous system that can generate oscillatory patterns for the rhythmic movements. Its simplified format, neural oscillator, is wildly adopted in engineering application. This paper explores the CPG from an integral view that combines biology and engineering together. Biological CPG and simplified CPG are both studied. Computer simulation reveals the mechanism of CPG. Some properties, such as effect of tonic input and sensory feedback, stable oscillation, robustness, entrainment etc., are further studied. The promising results provide foundation for the potential engineering application in future.     

Velocity of head movements and sensory-motor adaptation during and after short spaceflight

To investigate to time course of sensory-motor adaptation to microgravity, we tested spatially-directed voluntary head movements before, during and after short spaceflight. We also tested the re-adaptation of postural responses to sensory stimulation after space flight. The cosmonaut performed in microgravity six cycles of voluntary head rotation in pitch, roll and yaw directions. During the first days of weightlessness the angular velocity of head movements increased. Over the next days of microgravity the velocity of head movements gradually decreased. On landing day a significant decrease of head rotation velocity was observed compared to the head movement velocity before spaceflight. Re-adaptation to Earth condition measured by body sway on soft support showed similar time course, but re-adaptation measured by postural responses to vestibular galvanic stimulation was prolonged. These results showed that the angular velocity of aimed head movements of cosmonauts is a good indicator of sensory-motor adaptation in altered gravity conditions.     

Pattern generating and reflex-like processes controlling aiming movements in the presence of inertia,damping and gravity

A model is proposed, in which goal-directed movements of the forearm are controlled by a central pattern generator (CPG) initiated for exactly one period, and by reflex-analogous processes. Movement width is proportional to the amplitude factor of the CPG's output, and to the square of the CPG's period length. The period duration can be freely selected, thus enabling the CPG to accommodate its time scale to the period of others CPG's. Parameters which influence movement accuracy can be adjusted by means of closed control loop, which are discrete with respect to time: The time unit corresponds to the period of the CPG. For instance, momentum adjustment balances the CPG in such a manner that the velocity of the arm becomes zero on termination of the period, while gain adjustment serves to attain a correct movement length in the presence of an inertial load. Friction, stiffness and gravitational force are neutralized by additional reflex-type processes, interpretable as positive feedback loops with adjustable gain factors, using position and velocity signals.     

CPG Control for Biped Hopping Robot in Unpredictable Environment

A CPG control mechanism is proposed for hopping motion control of biped robot in unpredictable environment.Based on analysis of robot motion and biological observation of animal's control mechanism,the motion control task is divided into two simple parts:motion sequence control and output force control.Inspired by a two-level CPG model,a two-level CPG control mechanism is constructed to coordinate the drivers of robot joint,while various feedback information are introduced into the control mechanism.Interneurons within the control mechanism are modeled to generate motion rhythm and pattern promptly for motion sequence control; motoneurons are modeled to control output forces of joint drivers in real time according to feedbacks.The control system can perceive changes caused by unknown perturbations and environment changes according to feedback information,and adapt to unpredictable environment by adjusting outputs of neurons.The control mechanism is applied to a biped hopping robot in unpredictable environment on simulation platform,and stable adaptive motions are obtained.     

Influence of Vibration on Motor Responses in the Upper Arm Muscles under Stationary Conditions and during Voluntary and Evoked Arm Movements under Limb Unloading Conditions

Locomotion of mammals, including humans, is based on the rhythmic activity of spinal cord circuitries. The functioning of these circuitries depends on multimodal afferent information and on supraspinal influences from the motor cortex. Using the method of transcranial magnetic stimulation (TMS) of arm muscle areas in the motor cortex, we studied the motor evoked potentials (MEP) in the upper arm muscles in stationary conditions and during voluntary and vibration-evoked arm movements. The study included 13 healthy subjects under arm and leg unloading conditions. In the first series of experiments, with motionless limbs, the effect of vibration of left upper arm muscles on motor responses in these muscles was evaluated. In the second series of experiments, MEP were compared in the same muscles during voluntary and rhythmic movements generated by left arm m. triceps brachii vibration (the right arm was stationary). Motionless left arm vibration led to an increase in MEP values in both vibrated muscle and in most of the non-vibrated muscles. For most target muscles, MEP was greater with voluntary arm movements than with vibration-evoked movements. At the same time, a similar MEP modulation in the cycle of arm movements was observed in the same upper arm muscles during both types of arm movements. TMS of the motor cortex significantly potentiated arm movements generated by vibration, but its effect on voluntary movements was weaker. These results indicate significant differences in the degree of motor cortex involvement in voluntary and evoked arm movements. We suppose that evoked arm movements are largely due to spinal rather than central mechanisms of generation of rhythmic movements.     

Age-Related Features of the Voluntary Control of the Upright Posture

The capability for the unconscious control of the upright posture in elderly people is impaired, which increases the risk of falls and traumata. The impairment of the unconscious control of posture is partially compensated by the fixation of voluntary attention on the maintenance of an appropriate posture. Elderly people fall predominantly during the performance of movements that demand additional voluntary postural control, for example, unstable support conditions. Thus, voluntary postural control assumes importance in elderly persons. Since it is unclear whether this function changes with age, the aim of this work was to study age-related features of the learning voluntary postural control using visual feedback by center-of-pressure position. The results of the study showed that voluntary postural control is a complex multicomponent process that includes, at least, the following functions: selection of a strategy of postural control, its actualization, and precision of its regulation. With aging, strategy selection in healthy people impairs, but both elderly and middle-aged people can learn this function as successfully as the young. At the same time, despite the absence of an initial deficit in the accuracy of postural setting in elderly people, training of this function becomes substantially more difficult with age.     

Resonance tuning in a neuro-musculo-skeletal model of the forearm

In rhythmic movements, humans activate their muscles in a robust and energy efficient way. These activation patterns are oscillatory and seem to originate from neural networks in the spinal cord, called central pattern generators (CPGs). Evidence for the existence of CPGs was found for instance in lampreys, cats and rats. There are indications that CPGs exist in humans as well, but this is not proven yet. Energy efficiency is achieved by resonance tuning: the central nervous system is able to tune into the resonance frequency of the limb, which is determined by the local reflex gains. The goal of this study is to investigate if the existence of a CPG in the human spine can explain the resonance tuning behavior, observed in human rhythmic limb movement. A neuro-musculo-skeletal model of the forearm is proposed, in which a CPG is organized in parallel to the local reflexloop. The afferent and efferent connections to the CPG are based on clues about the organization of the CPG, found in literature. The model is kept as simple as possible (i.e., lumped muscle models, groups of neurons are lumped into half-centers, simple reflex model), but incorporates enough of the essential dynamics to explain behavior—such as resonance tuning—in a qualitative way. Resonance tuning is achieved above, at and below the endogenous frequency of the CPG in a highly non-linear neuro- musculo-skeletal model. Afferent feedback of muscle lengthening to the CPG is necessary to accomplish resonance tuning above the endogenous frequency of the CPG, while feedback of muscle velocity is necessary to compensate for the phase lag, caused by the time delay in the loop coupling the limb to the CPG. This afferent feedback of muscle lengthening and velocity represents the Ia and II fibers, which—according to literature—is the input to the CPG. An internal process of the CPG, which integrates the delayed muscle lengthening and feeds it to the half-center model, provides resonance tuning below the endogenous frequency. Increased co-contraction makes higher movement frequencies possible. This agrees with studies of rhythmic forearm movements, which have shown that co-contraction increases with movement frequency. Robustness against force perturbations originates mainly from the CPG and the local reflex loop. The CPG delivers an increasing part of the necessary muscle activation for increasing perturbation size. As far as we know, the proposed neuro-musculo-skeletal model is the first that explains the observed resonance tuning in human rhythmic limb movement.     

Saccades Improve Postural Control: A Developmental Study in Normal Children

Introduction

Dual-task performance is known to affect postural stability in children. This study focused on the effect of oculomotor tasks like saccadic eye movements on postural stability, studied in a large population of children by recording simultaneously their eye movements and posture.

Materials and Methods

Ninety-five healthy children from 5.8 to 17.6 years old were examined. All children were free of any vestibular, neurological, ophtalmologic and orthoptic abnormalities. Postural control was measured with a force platform TechnoConcept®, and eye movements with video oculography (MobilEBT®). Children performed two oculomotor tasks: fixation of a stable central target and horizontal saccades. We measured the saccade latency and the number of saccades during fixation as well as the surface, length and mean velocity of the center of pressure.

Results

During postural measurement, we observed a correlation between the age on the one hand and a decrease in saccade latency as well as an improvement in the quality of fixation on the other. Postural sway decreases with age and is reduced in the dual task (saccades) in comparison with a simple task of fixation.

Discussion - Conclusion

These results suggest a maturation of neural circuits controlling posture and eye movements during childhood. This study also shows the presence of an interaction between the oculomotor system and the postural system. Engaging in oculomotor tasks results in a reduction of postural sway.     

Contribution of the frontal lobe secondary motor area to organization of postural components in human voluntary movement

A total of 225 patients with local verified brain lesions were investigated with a view to identifying the brain regions contributing to organizing postural aspects of voluntary movement. Impaired postural adjustment movements associated with voluntary deep breathing were found in patients with damage primarily to the posterior section of the frontal lobe inferior convolution. Impaired activation of leg and trunk muscles accompanying arm movements were revealed in patients with damage chiefly to the posterior section of the superior convolution of the lobe, including the accessory motor area. It was deduced that postural movements differing in their functional purpose are controlled, like other learned tasks, by different sections of the secondary motor zone of the frontal lobe of the brain.Institute for Information Transmission Studies, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 20, No. 1, pp. 7–15, January–February, 1988.     

On the definition of central pattern generator and its sensory control

A central pattern generator (CPG) is defined here as a neural network responsible for the production of the timing cues of a rhythmic motor output pattern in the isolated CNS. For the intact animal, model considerations show that this term is neither clearly delimited from the concept of a reflex chain nor from the concept of a pattern generator with functional principles different from those of the CPG. Therefore, it cannot be concluded from the existence of a CPG in the isolated nervous system that this CPG also provides the decisive timing cues in the intact animal. Consequences for the study of the neural basis of rhythmic movements are shown.     

The investigation of locomotor activity control system in humans under the air-stepping conditions during passive and voluntary leg movements

The degree of activation of the central stepping program during passive leg movement was studied in healthy subjects under unloading conditions; the excitability of spinal motoneurons was studied during passive and voluntary stepping movements. Passive stepping movements with characteristics maximally close to those during voluntary stepping were accomplished by the experimenter. The bursts of muscular activity during voluntary and imposed stepping movements were compared. In addition, the influence on the leg movement of artificially created loading onto the foot was studied. The excitability of spinal motoneurons was estimated by the amplitude of modulation of the m. soleus H reflex. Changes in the H reflex (Hoffmann’s reflex) after fixation of the knee and hip joints were also studied. In most subjects, passive movements were accompanied by bursts of electromyographic (EMG) activity in the hip muscles (sometimes in shank muscles); the timing of the EMG burst during the step cycle coincided with the burst’s timing during voluntary stepping. In many cases, the bursts in EMG activity exceeded the activity of homonymous muscles during voluntary stepping. Simulation of foot loading influenced significantly the distal part of the moving extremity during both voluntary and passive movements, which was expressed in the appearance of movements in the ankle joint and an increase in the phasic EMG activity of the shank muscles. The excitability of motoneurons during passive movements was higher than during voluntary movements. Changes and modulation of the H reflex throughout the step cycle were similar without restriction of joint mobility and without hip joint mobility. Fixation of the knee joint was of great importance. It is supposed that imposed movements activate the same mechanisms of rhythm generation as supraspinal commands during voluntary movements. During passive movements, presynaptic inhibition depends mostly on the afferent influences from the moving leg rather than on the central commands. Under the conditions of “air-stepping,” the afferent influences from the foot pressure receptors are likely to interact actively with the central program of stepping and to determine the final activity pattern irrespective of the movement type (voluntary or passive).     

Intersegmental coordinating system of the lamprey central pattern generator for locomotion

Summary In the lamprey,Ichthyomyzon unicuspis, the wave of activity required for normal swimming movements can be generated by a central pattern generator (CPG) residing in the spinal cord. A constant phase coupling between spinal segments can be organized by intersegmental coordinating neurons intrinsic to the cord. The rostral and caudal segmental oscillators of the CPG have different preferred frequencies when separated from each other. Therefore the system must maintain the segmental oscillators of the locomotor CPG at a single common frequency and with the proper relative timing. Using selective lesions and a split-bath, it is demonstrated that the coordinating system is comprised of at least 3 subsystems, short-axon systems in the lateral and medial tracts and a long axon system in the lateral tracts. Each alone can sustain relatively stable coordinated activity.Abbreviations CPG central pattern generator - NMDA N-methyl-D-aspartate - VR ventral root     

Serotonin regulates rhythmic whisking

Many rodents explore their environment by rhythmically palpating objects with their mystacial whiskers. These rhythmic whisker movements ("whisking"; 5-9 Hz) are thought to be regulated by an unknown brainstem central pattern generator (CPG). We tested the hypothesis that serotonin (5-HT) inputs to whisking facial motoneurons (wFMNs) are part of this CPG. In response to exogenous serotonin, wFMNs recorded in vitro fire rhythmically at whisking frequencies, and selective 5-HT2 or 5-HT3 receptor antagonists suppress this rhythmic firing. In vivo, stimulation of brainstem serotonergic raphe nuclei evokes whisker movements. Unilateral infusion of selective 5-HT2 or 5-HT3 receptor antagonists suppresses ipsilateral whisking and substantially alters the frequencies and symmetry of whisker movements. These findings suggest that serotonin is both necessary and sufficient to generate rhythmic whisker movements and that serotonergic premotoneurons are part of a whisking CPG.     

The investigation of control mechanisms of stepping rhythm in human in the air-stepping conditions during passive and voluntary leg movements

In unloading condition the degree of activation of the central stepping program was investigated during passive leg movements in healthy subjects, as well as the excitability of spinal motoneurons during passive and voluntary stepping movement. Passive stepping movements with characteristics maximally approximated to those during voluntary stepping were accomplished by experimenter. The comparison of the muscle activity bursts during voluntary and imposed movements was made. In addition to that the influence of artificially created loading onto the foot to the leg movement characteristics was analyzed. Spinal motoneuron excitability was estimated by means of evaluation of amplitude modulation of the soleus H-reflex. The changes of H-reflexes under the fixation of knee or hip joints were also studied. In majority of subjects the passive movements were accompanied by bursts of EMG activity of hip muscles (and sometimes of knee muscles), which timing during step cycle was coincided with burst timing of voluntary step cycle. In many cases the bursts of EMG activity during passive movements exceeded activity in homonymous muscles during voluntary stepping. The foot loading imitation exerted essential influence on distal parts of moving extremity during voluntary as well passive movements, that was expressed in the appearance of movements in the ankle joint and accompanied by emergence and increasing of phasic EMG activity of shank muscles. The excitability of motoneurons during passive movements was greater then during voluntary ones. The changes and modulation of H-reflex throughout the step cycle without restriction of joint mobility and during exclusion of hip joint mobility were similar. The knee joint fixation exerted the greater influence. It is supposed that imposed movements activate the same mechanisms of rhythm generation as a supraspinal commands during voluntary movements. In the conditions of passive movements the presynaptic inhibition depend on afferent influences from moving leg in the most degree then on central commands. It seems that afferent inputs from pressure receptors of foot in the condition of "air-stepping" actively interact with central program of stepping and, irrespective of type of the performing movements (voluntary or passive), form the final pattern activity.     

Feedback equilibrium control during human standing

Equilibrium maintenance during standing in humans was investigated with a 3-joint (ankle, knee and hip) sagittal model of body movement. The experimental paradigm consisted of sudden perturbations of humans in quiet stance by backward displacements of the support platform. Data analysis was performed using eigenvectors of motion equation. The results supported three conclusions. First, independent feedback control of movements along eigenvectors (eigenmovements) can adequately describe human postural responses to stance perturbations. This conclusion is consistent with previous observations (Alexandrov et al., 2001b) that these same eigenmovements are also independently controlled in a feed-forward manner during voluntary upper-trunk bending. Second, independent feedback control of each eigenmovement is sufficient to provide its stability. Third, the feedback loop in each eigenmovement can be modeled as a linear visco-elastic spring with delay. Visco-elastic parameters and time-delay values result from the combined contribution of passive visco-elastic mechanisms and sensory systems of different modalities     

Analysis of the gait generation principle by a simulated quadruped model with a CPG incorporating vestibular modulation

This study aims to understand the principles of gait generation in a quadrupedal model. It is difficult to determine the essence of gait generation simply by observation of the movement of complicated animals composed of brains, nerves, muscles, etc. Therefore, we build a planar quadruped model with simplified nervous system and mechanisms, in order to observe its gaits under simulation. The model is equipped with a mathematical central pattern generator (CPG), consisting of four coupled neural oscillators, basically producing a trot pattern. The model also contains sensory feedback to the CPG, measuring the body tilt (vestibular modulation). This spontaneously gives rise to an unprogrammed lateral walk at low speeds, a transverse gallop while running, in addition to trotting at a medium speed. This is because the body oscillation exhibits a double peak per leg frequency at low speeds, no peak (little oscillation) at medium speeds, and a single peak while running. The body oscillation autonomously adjusts the phase differences between the neural oscillators via the feedback. We assume that the oscillations of the four legs produced by the CPG and the body oscillation varying according to the current speed are synchronized along with the varied phase differences to keep balance during locomotion through postural adaptation via the vestibular modulation, resulting in each gait. We succeeded in determining a single simple principle that accounts for gait transition from walking to trotting to galloping, even without brain control, complicated leg mechanisms, or a flexible trunk.     

Mechanisms of Locomotory Speed Change: The Pteropod Solution

Three primary factors contribute to locomotory speed changesin the pteropod mollusk Clione limacina. (1) An increase incycle frequency of locomotory appendages is associated witha baseline depolarization and enhancement of postinhibitoryrebound in central pattern generator (CPG) interneurons, anda reorganization of the CPG through recruitment of additionalinterneurons. (2) An increase in the force of appendage movementsis generated through enhancement of activity of active motoneurons,recruitment of additional motoneurons and peripheral modulationof swim musculature. (3) Changes in biomechanical aspects ofappendage movements are presumably achieved, at least in part,through changes in the activity of motoneurons and swim muscle.All changes associated with non-startle swim acceleration areproduced by a serotonergic arousal system that acts at all threelevels of the swimming system: CPG interneurons, motoneuronsand swim musculature.     

The contribution of sensory inputs to the pattern generation of breathing

Current concepts of the basic neural control system and its modulation by afferent inputs are reviewed. It is emphasized that, in analogy with locomotion, the central pattern generator (CPG) for automatic metabolic respiration does not depend on any afferent feedback from receptors sensitive to the movements of the "pump," or the streams of pumped air, for its production of a rhythmic motor output provides the CPG receives some "drive" inputs above threshold and adequate bias. The operation for a variety of reflexes and feedback loops is of fundamental importance, however, for adapting the breathing pattern to the varying requirements for gas exchange to the many behavioural, nonmetabolic demands on the breathing apparatus which are competing with its primary metabolic control functions. The presentation is focussed also on available evidence that the respiratory CPG exerts powerful modulations on the transmission in these reflex pathways controlling the pattern of breathing and adjusting it to the various metabolic and behavioural demands. Mechanisms for "gating," "phasic gain changes," and "phase-dependent reflex reversal" are exemplified.     

Effects of coupled upper limbs movements on postural stabilisation

The preference for in-phase association of coupled cyclic limbs movements is well described (mirror-symmetrical patterns) and this is demonstrated by the ease of performing in-phase movements compared to anti-phase ones. The hypothesis of this study is that the easiest movement patterns are those with minor postural activity. The aim of this study was to describe postural activity in standing subjects in the sagittal and frontal planes during the execution of three upper limbs tasks (single arm, in-phase, anti-phase) at four different frequencies (from 0.6 to 1.2 Hz).We employed six infrared cameras for recording kinematics information, a force platform for measuring forces exerted on the ground, and a system for surface electromyography (SEMG). Outcome measures were: upper limb range of movement and relative-phase, centre of pressure displacement (COP), screw torque (T_z) exerted on the ground, and SEMG recordings of postural muscles (adductor longus, gluteus medius, rectus femoris, and biceps femoris).Our results show that in both the planes the in-phase task resulted in less COP displacement, torque production, and postural muscles involvement than the anti-phase and single arm tasks. This reduced need of postural control could explain the ease of performing in-phase coupled limb movements compared with anti-phase movements.     

Effects of postural muscle fatigue on the relation between segmental posture and movement.

The purpose of this study was to examine whether fatigue of postural muscles might influence the coordination between segmental posture and movement. Seven healthy adults performed series of fifteen fast wrist flexions and extensions while being instructed to keep a dominant upper limb posture as constant as possible. These series of voluntary movements were performed before and after a fatiguing submaximal isometric elbow flexion, and also with or without the help of an elbow support. Surface EMG from muscles Delto?deus anterior, Biceps brachii, Triceps brachii, Flexor carpi ulnaris, Extensor carpi radialis were recorded simultaneously with wrist, elbow and shoulder accelerations and wrist and elbow displacements. Fatigue was evidenced by a shift of the elbow and shoulder muscles EMG spectra towards low frequencies. Kinematics of wrist movements and corresponding activations of wrist prime-movers, as well as the background of postural muscle activation before wrist movement were not modified. There were only slight changes in timing of postural muscle activations. These data indicate that postural fatigue induced by a low-level isometric contraction has no effect on voluntary movement and requires no dramatic adaptation in postural control.