Modulation of the soleus muscle H reflex caused by ballistic voluntary arm movements in humans

In healthy humans, we studied the influence of conditioning voluntary arm movements on the H reflex induced by transcutaneous stimulation of the tibial nerve and recorded from the soleus muscle. We examined the effects of flexion and extension of the forearm, as well as of finger clenching performed with the maximum rate. Conditioning arm movements were self-induced or realized upon presentation of a visual signal (light flash). We found that the pattern of changes in the H reflex is determined by the position of the subject’s body in the course of tests. The ipsilateral arm flexion in the elbow joint in the standing position resulted in depression of the H reflex lasting about 100 msec from the beginning of the movement, while the effect observed in the lying position (on the couch with the feet hanging free in the air) looked like a facilitation of the reflex lasting about 100 to 200 msec. The direction and dynamics of modifications of the H reflex under conditions of the use of different conditioning movements (forearm flexions/extensions and finger clenching of the ipsilateral arm, as well as contralateral forearm flexions in the elbow joint) were rather similar. We also showed that the observed facilitation of the H reflex began earlier than the voluntary arm movement (40 to 50 msec prior to the beginning). We hypothesize that these conditioning influences result from the action of central motor commands and represent the factor related to anticipatory postural rearrangements. Such rearrangements are directed toward the maintenance of equilibrium of the body in the course of a future movement. These commands depend significantly on the spatial position of the subject’s body. Neirofiziologiya/Neurophysiology, Vol. 40, No. 2, pp. 147–154, March–April, 2008.     

Changes in the H Reflex Preceding Voluntary Movements of the Contralateral Lower Limb in Humans

In electromyographic studies on healthy subjects, we recorded the H reflex from the right m. soleus and measured changes in the magnitude of this reflex response related to voluntary movements of the contralateral lower limb performed according to a visual signal. The effects of back and plantar flexions of the contralateral foot of the tested subject in the lying and standing positions were examined. Changes in the H reflex magnitude began to be recorded 60 to 90 msec prior to voluntary movements of the contralateral limb. When the subject was in the lying position, these changes looked like facilitation of the H reflex at both types of movement of the contralateral foot. When the subject stood, facilitation preceded back flexion of the foot of this extremity, while plantar flexion was preceded by inhibition of the tested H reflex. Our results show that the pattern of preliminary changes in the muscle tone of one of the lower limbs is determined by the type of future movement of another limb and peculiarities of the support function realized by this limb.     

Modulation of soleus H-reflex during dorsal and plantar flexions in the human ankle joint

Recording of the H-reflex was used to study the changes in the reflex excitability of soleus motoneurons during dorsal and plantar flexions of the ipsilateral and contralateral feet performed with different strengths by 15 healthy subjects. The dorsiflexion of the ipsilateral foot was accompanied by the “classic” reciprocal inhibition of the soleus motoneurons, the degree of the inhibition being directly proportional to the strength of the contraction of pretibial muscles and depending on the presence of foot support. The plantar flexion of the ipsilateral foot was accompanied by changes in reflex excitability, which were inversely proportional to the strength of the flexion. This was apparently related to the activation of a mechanism protecting the muscle against excessive contraction. The dorsal and plantar flexions of the contralateral foot were accompanied by similar changes in the reflex excitability of soleus motoneurons, namely, an increase in the case of weak contraction and a decrease in the case of strong contraction. However, the increase in reflex excitability during contralateral dorsiflexion was smaller and its decrease began at a weaker contraction than in the case of contralateral plantar flexion. The changes in the reflex excitability of soleus motoneurons during movements of the contralateral foot, which were also strength-dependent, confirmed the presence of cross-projections that are likely to be part of the generator of the central pattern of lower limb movement coordination.     

Facilitation and Inhibition of the <Emphasis Type="Italic">Soleus</Emphasis> H Reflex Related to Movements of the Upper Limb in Humans

In studies on healthy volunteers, we recorded an EMG discharge from the m. soleus corresponding to the H reflex evoked by transcutaneous stimulation of the n. tibialis comm. Changes in the magnitude of this reflex related to realization of brief voluntary movements of the ipsilateral upper limb were examined. The subjects were in a prone position. Fast flexion-extension of the forearm resulted first in 100- to 200-msec-long facilitation of the H reflex begun 30–40 msec before the appearance of EMG activity in the m. biceps brachii; this feature is indicative of the central nature of this effect related to the action of motor programs initiating the forearm movement. Facilitation of the H reflex was followed by its inhibition lasting several seconds. Within an interval corresponding to the maximum suppression of the H response, we tested the effect of additional conditioning stimulation of the n. peroneus comm. Occlusion of the inhibitory effects indicates that the same inhibitory neurons mediate the influences from both the peroneal input and the pathways transmitting inhibitory influences from the neuronal systems controlling upper limb muscles. Contractions of the ipsilateral m. biceps brachii evoked by direct electrical stimulation of the latter also resulted in inhibition of the soleus H reflex, which was rather similar in its time course to the above-mentioned inhibitory effects. There was no inhibition of the reflex after stimulations of the cutaneous receptors and n. medianus. These findings allow us to suppose that long-lasting inhibition of the H reflex induced by voluntary movements of the upper limb results from afferent influences from the receptors of contracting muscles. Such effects can be realized via the propriospinal pathways or long reflex arcs.     

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.     

Effects of Contralateral Afferent Stimulation on H Reflex in Humans

We studied changes of the H reflex recorded from the m. soleus, which were evoked by conditioning transcutaneous stimulation of the n. tibialis and n. peroneous comm. of the contralateral leg. In both cases, rather similar two-phase changes in the amplitude of the tested H reflex were observed. After a latent period (50 to 60 msec), the reflex was facilitated for about 300 msec, with the maximum at an about 100-msec-long interval. Then, facilitation was replaced by inhibition; the time course of the latter at test intervals longer than 500 msec could be satisfactorily approximated by a logarithmic curve. The mean durations of inhibition calculated with the use of a least-square technique were 4.0 and 2.7 sec in the cases of stimulation of n. tibialis and n. peroneous comm., respectively. Facilitation of the reflex was initiated with the intensity of conditioning stimulation corresponding to the threshold for excitation of cutaneous receptors. Facilitation could also be evoked by electrical stimulation of the skin in the contralateral popliteal dimple outside the projections of the above-mentioned nerves. Inhibition of the H reflex was evoked only with greater intensities of transcutaneous stimulation of the contralateral nerves corresponding to activation of low-threshold afferents of the above-mentioned nerves. The examined inhibition of the H reflex is probably of a presynaptic nature because it was not eliminated by tonic activation of the motoneurons of the tested muscle evoked by voluntary sole flexion. Long-lasting contralateral presynaptic inhibition can play a noticeable role in redistribution of the tone of skeletal muscles in the course of the motor activity. Neirofiziologiya/Neurophysiology, Vol. 37, No. 4, pp. 372–378, July–August, 2005.     

Using voluntary motor commands to inhibit involuntary arm movements

A hallmark of voluntary motor control is the ability to stop an ongoing movement. Is voluntary motor inhibition a general neural mechanism that can be focused on any movement, including involuntary movements, or is it mere termination of a positive voluntary motor command? The involuntary arm lift, or ‘floating arm trick’, is a distinctive long-lasting reflex of the deltoid muscle. We investigated how a voluntary motor network inhibits this form of involuntary motor control. Transcranial magnetic stimulation of the motor cortex during the floating arm trick produced a silent period in the reflexively contracting deltoid muscle, followed by a rebound of muscle activity. This pattern suggests a persistent generator of involuntary motor commands. Instructions to bring the arm down voluntarily reduced activity of deltoid muscle. When this voluntary effort was withdrawn, the involuntary arm lift resumed. Further, voluntary motor inhibition produced a strange illusion of physical resistance to bringing the arm down, as if ongoing involuntarily generated commands were located in a ‘sensory blind-spot’, inaccessible to conscious perception. Our results suggest that voluntary motor inhibition may be a specific neural function, distinct from absence of positive voluntary motor commands.     

The role of co-activation in strength and force modulation in the elbow of children with unilateral cerebral palsy

To study the role of coactivation in strength and force modulation in the elbow joint of children and adolescents with cerebral palsy (CP), we investigated the affected and contralateral arm of 21 persons (age 8-18) with spastic unilateral CP in three tasks: maximal voluntary isokinetic concentric contraction and passive isokinetic movement during elbow flexion and extension, and sub-maximal isometric force tracing during elbow flexion. Elbow flexion-extension torque and surface electromyography (EMG) of the biceps brachii (BB) and triceps brachii (TB) muscles were recorded. During the maximal contractions, the affected arm was weaker, had decreased agonist and similar antagonist EMG amplitudes, and thus increased antagonist co-activation (% of maximal activity as agonist) during both elbow flexion and extension, with higher coactivation levels of the TB than the BB. During passive elbow extension, the BB of the affected arm showed increased resistance torque and indication of reflex, and thus spastic, activity. No difference between the two arms was found in the ability to modulate force, despite increased TB coactivation in the affected arm. The results indicate that coactivation plays a minor role in muscle weakness in CP, and does not limit force modulation. Moreover, spasticity seems particularly to increase coactivation in the muscle antagonistic to the spastic one, possibly in order to increase stability.     

Locomotor-Like Rotation of Either Hip or Knee Inhibits Soleus H Reflexes in Humans

Human soleus H reflexes are depressed with passive movement of the leg. We investigated the limb segment origin of this inhibition. In the first experiment, H reflexes were evoked in four subjects during (1) passive pedaling movement of the test leg at 60 rpm; (2 and 3) pedaling-like flexion and extension of the hip and the knee of the test leg separately; and (4) stationary controls. In the second experiment, with the test leg stationary, the same series of movements occurred in the opposite leg. Rotation of the hip or the knee of the test leg significantly reduced mean reflex amplitudes (p > 0.01) to levels similar to those for whole-leg movement (mean H reflexes: stationary, 71%; test leg pedaling movement, 10%; knee rotation, 15%; hip rotation, 13% [all data are given as percentages of M_max]). The angle of the stationary joint did not significantly affect the results. Rotation of the contralateral hip significantly reduced mean reflex magnitudes. Rotation of the contralateral knee had a similar effect in three of the four subjects. We infer that a delimited field of receptors induces the movement conditioning of both the ipsilateral and contralateral spinal paths. It appears that somatosensory receptor discharge from movement of the hip or knee of either leg induces inhibition as the foundation for the modulation of H reflexes observed during human movement.     

Analysis of spike discharge of a neuron population in the cat motor cortex during a conditioned change of posture reflex

Spike responses of area 4 neurons in the projection area of the contralateral forelimb to acoustic stimulation (1 sec), which became the conditioned stimulus after training, and to dropping of the platform beneath the test limb, which served as reinforcing stimulus, were studied in trained and untrained cats. Responses only of those neurons which were activated during a passive movement caused by dropping of the platform were studied. In trained animals the number of these neurons which responded to the conditioned stimulus if a reflex occurred was 100%, and in the absence of conditioned-reflex movements to the conditioned stimulus it was 70%, much greater than the number of neurons responding to the same acoustic stimulus in untrained animals (45%). On peristimulus histograms of responses of the test neuron population in untrained and trained animals to acoustic stimulation (in the absence of movements) only the initial spike response with a latent period of under 50 msec and a duration of up to 100 msec could be clearly distinguished. In the presence of reflex movement multicomponent spike responses were observed: an initial spike response and early and late after-responses linked with performance of conditioned-reflex limb flexion. Early after-responses 100–200 msec in duration, appearing after a latent period of 100–150 msec, were linked to the time of application of the conditioned stimulus, whereas the appearance and duration of late after-responses were determined by the time of onset of conditioned-reflex movement. The magnitude of the neuronal response to reinforcement in trained animals does not depend on the appearance of the conditioned movement.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 17, No. 1, pp. 93–102, January–February, 1985.     

Characteristics and lateral dominance of hand grip and elbow flexion powers in young male adults

This study aimed to clarify the characteristics and the lateral dominance of hand grip power and elbow flexion power. The subjects were 15 healthy young males (mean age 22.1+/-0.7 yr, mean height 171.3+/-3.4 cm, mean mass 64.5+/-4.1 kg). All subjects were right-handed. Peak power was measured by both hands with 6 different loads of 20%-70% of maximum voluntary contraction. The maximum voluntary contraction of hand grip movement and elbow flexion movement was significantly larger in the dominant hand. Peak power of the dominant hand was larger in all loads in hand grip movement and in loads of 20% and 30% of maximum voluntary contraction in elbow flexion movement. In short, lateral dominance was confirmed. Peak power was significantly larger in hand grip movement than in elbow flexion movement in both hands. Peak velocity decreased with increasing loads in both movements, but peak power increased until about 50% of maximum voluntary contraction and then decreased. The peak power ratio of the dominant hand to the nondominant hand was significantly larger in hand grip movement than in elbow flexion movement in all loads and the peak power ratio in elbow flexion movement was more marked in light loads. In conclusion, both powers showed lateral dominance. Lateral dominance is more marked in hand grip power.     

Effects of fatiguing unilateral plantar flexions on corticospinal and transcallosal inhibition in the primary motor hand area

Background

Corticospinal excitability of the primary motor cortex (M1) representing the hand muscle is depressed by bilateral lower limb muscle fatigue. The effects of fatiguing unilateral lower limb contraction on corticospinal excitability and transcallosal inhibition in the M1 hand areas remain unclear. The purpose of this study was to determine the effects of fatiguing unilateral plantar flexions on corticospinal excitability in the M1 hand areas and transcallosal inhibition originated from the M1 hand area contralateral to the fatigued ankle.

Methods

Ten healthy volunteers (26.2 ± 3.8 years) participated in the study. Using transcranial magnetic stimulation, we examined motor evoked potentials (MEPs) and interhemispheric inhibition (IHI) recorded from resting first dorsal interosseous (FDI) muscles before, immediately after, and 10 min after fatiguing unilateral lower limb muscle contraction, which was consisted of 40 unilateral maximal isometric plantar flexions intermittently with a 2-s contraction followed by 1 s of rest.

Results

We demonstrated no significant changes in MEPs in the FDI muscle ipsilateral to the fatigued ankle and decrease in IHI from the M1 hand area contralateral to the fatigued ankle to the ipsilateral M1 hand area after the fatiguing contraction. MEPs in the FDI muscle contralateral to the fatigued ankle were increased after the fatiguing contraction.

Conclusions

These results suggest that fatiguing unilateral lower limb muscle contraction differently influences corticospinal excitability of the contralateral M1 hand area and IHI from the contralateral M1 hand area to the ipsilateral M1 hand area. Although fatiguing unilateral lower limb muscle contraction increases corticospinal excitability of the ipsilateral M1 hand area, the increased corticospinal excitability is not associated with the decreased IHI.     

Convergence in Reflex Pathways from Multiple Cutaneous Nerves Innervating the Foot Depends upon the Number of Rhythmically Active Limbs during Locomotion

Neural output from the locomotor system for each arm and leg influences the spinal motoneuronal pools directly and indirectly through interneuronal (IN) reflex networks. While well documented in other species, less is known about the functions and features of convergence in common IN reflex system from cutaneous afferents innervating different foot regions during remote arm and leg movement in humans. The purpose of the present study was to use spatial facilitation to examine possible convergence in common reflex pathways during rhythmic locomotor limb movements. Cutaneous reflexes were evoked in ipsilateral tibialis anterior muscle by stimulating (in random order) the sural nerve (SUR), the distal tibial nerve (TIB), and combined simultaneous stimulation of both nerves (TIB&SUR). Reflexes were evoked while participants performed rhythmic stepping and arm swinging movement with both arms and the leg contralateral to stimulation (ARM&LEG), with just arm movement (ARM) and with just contralateral leg movement (LEG). Stimulation intensities were just below threshold for evoking early latency (<80 ms to peak) reflexes. For each stimulus condition, rectified EMG signals were averaged while participants held static contractions in the stationary (stimulated) leg. During ARM&LEG movement, amplitudes of cutaneous reflexes evoked by combined TIB&SUR stimulation were significantly larger than simple mathematical summation of the amplitudes evoked by SUR or TIB alone. Interestingly, this extra facilitation seen during combined nerve stimulation was significantly reduced when performing ARM or LEG compared to ARM&LEG. We conclude that locomotor rhythmic limb movement induces excitation of common IN reflex pathways from cutaneous afferents innervating different foot regions. Importantly, activity in this pathway is most facilitated during ARM&LEG movement. These results suggest that transmission in IN reflex pathways is weighted according to the number of limbs directly engaged in human locomotor activity and underscores the importance of arm swing to support neuronal excitability in leg muscles.     

Changes in movement final position associated with agonist and antagonist muscle fatigue

We have tested the hypothesis that agonist and antagonist muscle fatigue could affect the final position of rapid, discrete movements. Six subjects performed consecutive elbow flexion and extension movements between two targets, with their eyes closed prior to, and after fatiguing the elbow extensor muscles. The results demonstrate that elbow extension movements performed in the post-test period systematically undershot the final position as compared to pre-test movements. However, attainment of the aimed final position in elbow flexion movements was unaffected by fatiguing of the extensor muscles. Undershoot of the final position obtained in extension movements was associated with agonist muscle fatigue, a result that was expected from the point of view of current motor control theories, and that could be explained by a reduced ability of the shortening muscle to exert force. On the other hand, the absence of the expected overshoot of the final position when the antagonist is fatigued, indicates the involvement of various reflex and/or central mechanisms operating around the stretched muscle that could contribute to returning the limb to the standard final position after a brief prominent overshoot.     

Role of agonist and antagonist muscle strength in performance of rapid movements

Six subjects performed rapid self-terminated elbow movements under different mechanical conditions prior to, and 5 weeks after an elbow extensor strengthening programme. Despite the large difference in the strengths of elbow flexors and extensors, the pretest did not demonstrate significant differences between the movement time of flexion and extension movements performed under the same mechanical conditions. The results obtained in the posttest demonstrated a decrease in movement time (i.e. an increase in movement speed) in both elbow flexion and extension movements under some mechanical conditions. In addition, flexion movements demonstrated a relative increase in the acceleration time (acceleration time as a proportion of the movement time). It was concluded that the strength of both the agonist and antagonist muscles was important for the performance of rapid movements. Stronger agonists could increase the acceleration of the limb being moved, while stronger antagonists could facilitate the arrest of the limb movement in a shorter time, providing a longer time for acceleration.     

Effect of long-duration spaceflight on postural control during self-generated perturbations.

This report is the first systematic evaluation of the effects of prolonged weightlessness on the bipedal postural control processes during self-generated perturbations produced by voluntary upper limb movements. Spaceflight impacts humans in a variety of ways, one of which is compromised postflight postural control. We examined the neuromuscular activation characteristics and center of pressure (COP) motion associated with arm movement of eight subjects who experienced long-duration spaceflight (3--6 mo) aboard the Mir space station. Surface electromyography, arm acceleration, and COP motion were collected while astronauts performed rapid unilateral shoulder flexions before and after spaceflight. Subjects generally displayed compromised postural control after flight, as evidenced by modified COP peak-to-peak anterior-posterior and mediolateral excursion, and pathlength relative to preflight values. These changes were associated with disrupted neuromuscular activation characteristics, particularly after the completion of arm acceleration (i.e., when subjects were attempting to maintain upright posture in response to self-generated perturbations). These findings suggest that, although the subjects were able to assemble coordination modes that enabled them to generate rapid arm movements, the subtle control necessary to maintain bipedal equilibrium evident in their preflight performance is compromised after long-duration spaceflight.     

Transcranial Direct Current Stimulation Improves Ipsilateral Selective Muscle Activation in a Frequency Dependent Manner

Failure to suppress antagonist muscles can lead to movement dysfunction, such as the abnormal muscle synergies often seen in the upper limb after stroke. A neurophysiological surrogate of upper limb synergies, the selectivity ratio (SR), can be determined from the ratio of biceps brachii (BB) motor evoked potentials to transcranial magnetic stimulation prior to forearm pronation versus elbow flexion. Surprisingly, cathodal transcranial direct current stimulation (c-TDCS) over ipsilateral primary motor cortex (M1) reduces (i.e. improves) the SR in healthy adults, and chronic stroke patients. The ability to suppress antagonist muscles may be exacerbated at high movement rates. The aim of the present study was to investigate whether the selective muscle activation of the biceps brachii (BB) is dependent on altering frequency demands, and whether the c-tDCS improvement of SR is dependent on task frequency. Seventeen healthy participants performed repetitive isometric elbow flexion and forearm pronation at three rates, before and after c-tDCS or sham delivered to ipsilateral left M1. Ipsilateral c-tDCS improved the SR in a frequency dependent manner by selectively suppressing BB antagonist excitability. Our findings confirm that c-tDCS is an effective tool for improving selective muscle activation, and provide novel evidence for its efficacy at rates of movement where it is most likely to benefit task performance.     

State of monosynaptic reflex (H reflex) arc during voluntary muscle contraction in humans

Effect of an isometric voluntary contraction of the soleus muscle (5–6 sec in duration) on the H reflex was studied in persons aged 25–45 years. A sharp enhancement of the H reflex at a dynamic phase of fast contraction (in the force increase period) and its less pronounced increase at a static phase of contraction were shown. When a paired stimulation of the nerve was used (interstimulus interval, 500 msec), a voluntary contraction of the muscle fully abolished the first stimulus-related suppression of the second H reflex. When the muscle contracted against the background of vibration, the suppressed H reflex recovered up the the level observed in the resting muscle, but did not reach the level characteristic for the contracting muscle. The findings show that the two pathways controlling the H reflex (descending and afferent) function independently, and apparently there exist at least two corresponding groups of interneurons causing a presynaptic inhibition of the H reflex.Neirofiziologiya/Neurophysiology, Vol. 25, No. 5, pp. 365–371, September–October, 1993.     

Muscle power output properties using the stretch-shortening cycle of the upper limb and their relationships with a one-repetition maximum bench press

The purpose of this study was to examine the output properties of muscle power by the dominant upper limb using SSC, and the relationships between the power output by SSC and a one-repetition maximum bench press (1 RM BP) used as a strength indicator of the upper body. Sixteen male athletes (21.4+/-0.9 yr) participated in this study. They pulled a load of 40% of maximum voluntary contraction (MVC) at a stretch by elbow flexion of the dominant upper limb in the following three preliminary conditions: static relaxed muscle state (SR condition), isometric muscle contraction state (ISO condition), and using SSC (SSC condition). The velocity with a wire load via a pulley during elbow flexion was measured accurately using a power instrument with a rotary encoder, and the muscle power curve was drawn from the product of the velocity and load. Significant differences were found among all evaluation parameters of muscle power exerted from the above three conditions and the parameters regarding early power output during concentric contraction were larger in the SSC condition than the SR and ISO conditions. The parameters on initial muscle contraction velocity when only using SSC significantly correlated with 1 RM BP (r=0.60-0.62). The use of SSC before powerful elbow flexion may contribute largely to early explosive power output during concentric contraction. Bench press capacity relates to a development of the above early power output when using SSC.     

Lateral facilitation of Hoffmann-reflexes prior to voluntary movement in a choice reaction time task.

Hoffmann reflexes (H reflexes) were elicited from both legs simultaneously in human subjects at varying intervals after a reaction signal (RS) in a binary choice reaction time task. A left light RS required a rapid plantar flexion of the left foot and a right light RS required a similar rapid response of the right foot. A large faciliataion of reflex amplitude occurred only in the muscle involved in the movement (right of left soleus). The timing of the facilitation indicated that a decision about the status of the RS occurred within 200 msec and probably was completed somewhat earlier. Furthermore, the facilitation of the H reflexes was shown to be closely linked with the organization required for the contractions of the responding muscle. The results are considered in the light of hypothesized mechanisms regulating voluntary movement.