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PubMed1993

Stretch reflexes of individual parkinsonian patients studied during changes in clinical rigidity following medication.

Stretch reflexes were elicited in flexor carpi radialis (FCR) of healthy subjects and patients with Parkinson's disease by forcible ramp and hold extensions of the wrist joint. Individual patients were studied off treatment when rigidity was detected clinically at the joint and throughout the clinical response to anti-parkinsonian medication that abolished or reduced their rigidity. In this way the possible effects of inter-subject variability upon the relationship between reflex behaviour and rigidity were eliminated. The long-latency (M2) stretch reflexes of the patient group were increased on average compared to those of healthy subjects. However, in the large majority of individual patients there were no significant correlations between the amplitudes of their M2 or total (short-latency (M1) + M2) reflex activities, recorded off and on treatment, and the accompanying changes in clinically assessed rigidity. These results suggest that parkinsonian rigidity cannot be uniquely attributed to the increased reflex responsiveness measured by the present laboratory techniques. However, the techniques used to test reflex function in our study differed in several respects (e.g., background activity, stretching wave form) from those employed during clinical assessment of rigidity so that the balance of reflex mechanisms may have varied in the two situations. Therefore, these results cannot be taken as definitive evidence against a reflex origin of rigidity.

Aged↗
PubMed1983

Depressive effect of morphine on the sympathetic reflex elicited by stimulation of unmyelinated hindlimb afferent nerve fibers in anesthetized cats.

The effect of morphine on somatosympathetic reflexes was analyzed in central nervous system (CNS) intact, anesthetized cats. Sympathetic efferent nerve activity was recorded via an inferior cardiac nerve. Electrical train pulse stimulation of the superficial peroneal afferent nerve produced an A reflex of about 60 ms latency (evoked by myelinated fiber excitation) and a C reflex of about 360 ms latency (evoked by unmyelinated fiber excitation), both of which have central reflex pathways through the medulla oblongata. An intravenous injection of morphine (2 mg/kg), while not affecting the A reflex, markedly depressed the C reflex. The degree of this depressive effect was somewhat dependent on the stimulus intensity applied to the somatic afferent nerves. Naloxone (0.2 mg/kg, i.v.) antagonized the depressive effects of morphine on the C reflex. The present results suggest that the somatosympathetic C reflex can be a useful indicator of central morphine effects in anesthetized animals.

Afferent Pathways↗
PubMed1992

The depressive effect of intrathecal clonidine on the spinal flexor reflex is enhanced after sciatic nerve section in rats.

The effect of intrathecal (i.t.) alpha 2-adrenoceptor agonist, clonidine, on the spinal nociceptive flexor reflex was studied in decerebrate, spinalized, unanesthetized rats with intact sciatic nerves or in rats in which the sciatic nerve had been ipsilaterally sectioned. In rats with intact nerves i.t. clonidine caused a dose-dependent biphasic effect on flexor reflex excitability. At low dose (10 ng) the effect of clonidine was purely facilitatory, whereas with 50-100 ng clonidine the initial facilitation was often followed by reflex depression. Long-lasting, strong reflex depression was observed after i.t. injection of high doses of clonidine (1 and 10 micrograms). Four to 18 days after sciatic nerve section, the depressive effect of clonidine on the flexor reflex was dramatically enhanced. Depression was frequently observed already with doses of 5 and 10 ng, and maximal depression was reached at 100 ng and 1 micrograms in axotomized rats. The facilitatory effect of low doses of clonidine on the reflex was also observed, although somewhat less frequently than in normals. The depressive effect of clonidine on the flexor reflex was reversed by the selective alpha 2-receptor antagonist, atipamezole (20 micrograms, i.t.), in rats with both intact and sectioned sciatic nerves. The present results revealed an increased sensitivity and effectiveness of the depression of spinal reflex mechanisms by i.t. clonidine after sciatic nerve section, which is opposite to the decreased sensitivity to i.t. morphine after axotomy that we observed previously.(ABSTRACT TRUNCATED AT 250 WORDS)

Adrenergic alpha-Antagonists↗
PubMed1988

Different effects of norepinephrine and serotonin on the electrical toe stimulation-induced reflexes in the rat spinal cord.

1. Electrical toe stimulation was shown to elicit reflex potentials in the ipsilateral common peroneal nerve in the rat. 2. This reflex consisted of the spinal reflex and the spino-bulbo-spinal reflex (SBS reflex). 3. Administration of norepinephrine into the subarachnoid space enhanced the spinal reflex but depressed the SBS reflex. On the other hand, serotonin enhanced both reflexes. 4. These different effects of norepinephrine and serotonin suggest different control of nociceptive reactions by these biogenic amines.

Animals↗
PubMed1985

Injury-induced plasticity of the flexor reflex in chronic decerebrate rats.

The hindlimb-flexor-withdrawal reflex elicited by stimulation of the skin of the hindpaw has been examined in chronic decerebrate rats. This flexor reflex manifests as a typical phasic avoidance response when measured either behaviourally in the decerebrate rat or electrophysiologically in the decerebrate-spinal preparation. Once the threshold of the cutaneous flexor-reflex afferents in the skin have been exceeded a brief burst of activity with only a short afterdischarge occurs in the flexor motoneurones. The response to sustained stimuli adapts rapidly. In the absence of any treatment to the hindlimb the threshold, duration and responsiveness of the reflex remains stable when tested repeatedly. Thermal or chemical stimuli of sufficient intensity to produce tissue injury and prolonged local inflammation in a hindpaw of the chronic decerebrate rat result in marked and long-lasting (several weeks) alterations in the ipsilateral withdrawal reflex. The mechanical threshold necessary to elicit the reflex by stimulation of the hindpaw falls so that light touch or brush can now elicit a response instead of the firm pressure or pinch required pre-injury. Suprathreshold stimuli to the inflamed skin generate a sustained oscillating pattern of flexion in contrast to the brief flicking movement found in control animals. Electromyographic recordings from the hamstring flexor muscles ipsilateral to the inflamed hindpaw show decreased mechanothresholds, increased spontaneous activity, prolonged afterdischarges to brief stimuli and a slowly adapting tonic response to sustained stimulation. Populations of single cutaneous mechanoreceptive C-primary afferents recorded both from untreated decerebrate rats and from rats with an inflamed hindpaw are indistinguishable in terms of their response properties. There is no difference in threshold, spontaneous activity or afterdischarge between the two populations. The possible mechanisms responsible for the conversion of the high threshold phasic flexor reflex into a low threshold tonic reflex are discussed as are the possible implications for sensory disorders that accompany chronic injury in man.

Animals↗
PubMed1989

Late flexion reflex in paraplegic patients. Evidence for a spinal stepping generator.

We demonstrated previously that electrical stimulation of the Flexor Reflex Afferents (FRA) induces a late flexion reflex with a central conduction time longer than 100 msec. Its latency is prolonged by increasing the intensity or the duration of the stimulation. This late reflex is therefore similar to the late flexion reflex observed in acute spinal cat with DOPA. Some findings suggest that in man the late flexion reflex could be inhibited at a premotoneuronal level by contralateral FRA stimulation. In relation to the late flexion reflex, a late contralateral facilitation of soleus monosynaptic reflex (MSR) was observed. Rhythmical activity was observed in only one patient who had an exceptional form of spinal myoclonus. This myoclonus could be modulated by FRA stimulation. These facts show that the reflex organization in paraplegic patients is similar to the one described in acute spinal cat with DOPA and therefore suggest that a spinal stepping generator could exist in humans.

Electric Stimulation↗
PubMed1995

Postural modulation of the soleus H reflex in young and old subjects.

The influence of different static postures on the soleus H reflex was assessed in 15 old (mean age = 76.3 years) and 10 young (mean age = 24.2 years) subjects. H reflex and M wave recruitment curves were obtained under 2 randomly administered conditions: (1) standing; and (2) prone. Once in place, the recording and stimulating electrodes were not removed until the completion of testing, to ensure that exact placement was maintained. A 1 msec current pulse was given transcutaneously to elicit the H reflex and M response. Static postural sway area (cm2) was assessed on a Kistler force platform using custom software (sample rate = 50 Hz/15 sec trials). Results demonstrated that the young subjects reduced the amplitude of the H reflex from the prone (Hmax/Mmax = 73.6%) to the standing (Hmax/Mmax = 59.9%) condition, whereas the old subjects did not (prone = 32.4%, standing = 38.2%). However, within the old group, 2 subgroups emerged--those who depressed the reflex similar to the young subjects (O-D, n = 6) and those who did not depress the reflex (O-ND, n = 9). Furthermore, there were significant differences in postural sway scores between the young and old, between the O-D and O-ND, but not between the O-D and young groups. These results suggest differences in the manner in which young and old subjects modulate the soleus H reflex when standing, and support the view that modulation of the stretch reflex may be important in the control of static posture.

Adult↗
PubMed2005

Ventricular reflex interactions during transmural myocardial ischemia.

The role of left ventricular receptors with sympathetic afferent fibers in the reflex response to myocardial ischemia is controversial, particularly in the canine model. Previous experiments have shown that reflex excitatory responses mediated by left ventricular sympathetic afferents can be detected in sinoaortic denervated and vagotomized dogs during transmural myocardial ischemia. The purpose of these experiments was to determine if reflex excitatory responses occur in dogs with intact left ventricular vagal afferents. Experiments were performed in 27 chloralose-anesthetized dogs with sinoaortic denervation. Responses of efferent renal sympathetic nerve activity, arterial, and left atrial pressures to transmural and non-transmural inferoposterior myocardial ischemia were measured before and after interruption of left ventricular sympathetic afferents by stellectomy. The adequacy of sympathetic deafferentation was assessed by measurement of renal nerve responses to epicardial bradykinin. Prior to stellectomy, excitatory responses were observed in 10 animals and inhibitory responses in 9 animals. The remaining animals had no responses and were excluded from analysis. In the excitatory group, reflex increases in renal nerve activity during both transmural and non-transmural inferoposterior ischemia were abolished by stellectomy and not replaced by inhibitory responses. In the inhibitory group, non-transmural inferoposterior ischemia elicited greater reflex decreases in renal nerve activity when left ventricular sympathetic afferents were intact. After stellectomy, transmural ischemia elicited greater reflex inhibition of renal nerve activity. Renal nerve responses to epicardial bradykinin were abolished by stellectomy. These results indicate that reflex excitatory responses mediated by left ventricular receptors with sympathetic afferent fibers can be elicited in dogs with intact vagal afferents. These excitatory responses are most apparent during transmural myocardial ischemia. In dogs with inhibitory responses to coronary occlusion, activation of sympathetic afferents during transmural ischemia appears to attenuate reflex inhibitory responses mediated by left ventricular vagal afferents.

Afferent Pathways↗
PubMed2006

Enteropancreatic reflexes mediating the pancreatic enzyme response to nutrients.

The observation that in dogs electrical stimulation of the vagus nerve elicited a strong secretory activity of the pancreas, prompted I. P. Pavlov in 1888 to conclude that the pancreatic secretory response to nutrients is mediated by enteropancreatic reflexes involving the vagus nerves. It took, however, more than 90 years until by studying the latency of pancreatic amylase response to exogenous and endogenous stimuli for the first time experimental evidence was provided for the actual existence of cholinergic vago-vagal enteropancreatic reflexes. Follow-up studies, based on stepwise extrinsic denervation of the pancreas, ruled out possible splanchnic pathways for enteropancreatic reflexes. In more recent years, experiments utilizing specific antagonists demonstrated a physiological role for both cholinergic M1 and cholecystokinin (CCK) receptors within the enteropancreatic reflex. At least a significant portion of the cholinergic fibres of the enteropancreatic reflex end on muscarinic receptors of the subtype M1. CCK, the most important hormone stimulating pancreatic enzyme secretion, appears to act at least in part on CCK receptors located on vagal afferent nerves, which in turn elicit a vago-vagal reflex, implying that CCK exerts its effect on the pancreas at least in part through vago-vagal reflexes. Furthermore, pharmacological blockade of CCK receptors totally abolished the early pancreatic amylase response to intestinal nutrients, suggesting that the activation of (probably vagal) CCK receptors is essential to run the enteropancreatic reflex.

Animals↗
PubMed2004

A C-fiber reflex inhibition induced by electroacupuncture with different intensities applied at homotopic and heterotopic acupoints in rats selectively destructive effects on myelinated and unmyelinated afferent fibers.

The aims of this study were to explore the C-fiber reflex inhibition induced by electroacupuncture with different intensities applied at homotopic or heterotopic acupoints, and to determine the influence selectively destroyed myelinated and unmyelinated afferent fibers on the C-fiber reflex inhibition. In the ipsilateral local acupoint, the general behavior of the C-fiber reflex can be depressed by electroacupuncture below the threshold of Adelta-fiber activation. Electroacupuncture stimulation within the intensity of Adelta-fiber activation applied to the ipsilateral limb pretreated with cobra venom did not elicit inhibition of C-fiber reflex in rats with demyelinated sciatic nerve. However, heterotopic electroacupuncture below the threshold of Adelta-fiber activation was totally ineffective. In contralateral heterotopic acupoints, the C-fiber reflex can be depressed only by electroacupuncture with stimulating intensities exceeding thresholds of Adelta and C-fiber activation. Electroacupuncture stimuli applied to capsaicin-pretreated limb in the intensities of threshold of Adelta-fiber and treble thresholds of C-fiber activation produced only a little inhibition of C-fiber reflex. Inhibitory intensity was roughly similar to that induced by the stimulation with intensity for the activation of Adelta-fiber in normal animal. In the spinalized animals transections at T6-T7 segments, regardless of intensities, the homotopic electroacupuncture stimulation only induced moderate depression of C-fiber reflex similar to that of Adelta-fiber activation; whereas, no matter what intensities of application, the inhibitory effects of C-fiber reflex disappeared totally by using heterotopic noxious electroacupuncture in these animals.

Acupuncture Points↗
PubMed2005

Effects of static flexion-relaxation on paraspinal reflex behavior.

BACKGROUND: Static trunk flexion working postures and disturbed trunk muscle reflexes are related to increased risk of low-back pain. Animal studies conclude that these factors may be related; passive tissue strain in spinal ligaments causes subsequent short-term changes in reflex. Although studies have documented changes in the myoelectric onset angle of flexion-relaxation following prolonged static flexion and cyclic flexion we could find no published evidence related to the human reflex response of the trunk extensor muscles following a period of static flexion-relaxation loading. METHODS: Eighteen subjects maintained static lumbar flexion for 15 min. Paraspinal muscle reflexes were elicited both before and after the flexion-relaxation protocol using pseudorandom stochastic force disturbances while recording EMG. Reflex gain was computed from the peak value of the impulse response function relating input force perturbation to EMG response using time-domain deconvolution analyses. FINDINGS: Reflexes showed a trend toward increased gain after the period of flexion-relaxation (P < 0.055) and were increased with trunk extension exertion (P < 0.021). Significant gender differences in reflex gain were observed (P < 0.01). INTERPRETATIONS: Occupational activities requiring extended periods of trunk flexion contribute to changes in reflex behavior of the paraspinal muscles. Results suggest potential mechanisms by which flexed posture work may contribute to low-back pain. Significant gender differences indicate risk analyses should consider personal factors when considering neuromuscular behavior.

Adaptation, Physiological↗
PubMed2004

Galvanic evoked vestibulospinal and vestibulocollic reflexes in stroke.

OBJECTIVE: Following stroke, the startle reflex, mediated via the reticulospinal tract, is often facilitated. Vestibulospinal reflexes are another bulbospinal reflex, abnormalities of which may contribute to impaired body posture and stance following stroke. We recorded galvanic evoked vestibulospinal and vestibulocollic reflexes to assess whether these showed similar changes to those for startle following stroke affecting the pons and above. METHODS: Twenty-four stroke subjects (aged 40-82) were studied in the vestibulospinal part of the study, 21 stroke subjects (aged 40-81 years) were studied in the vestibulocollic part, including 18 studied in both. Transmastoid galvanic (DC) current was used to stimulate the vestibular nerve. Vestibulocollic responses were recorded from the sternocleidomastoid muscles and vestibulospinal responses from over soleus in standing subjects. RESULTS: Vestibulocollic reflex amplitudes and latencies showed no significant differences between the two sides. Similarly short latency (SL) and medium latency (ML) vestibulospinal reflexes did not differ significantly in frequency, latency or amplitude between the affected and unaffected legs. CONCLUSIONS: Vestibular reflexes are not facilitated by stroke at or above the pontine level. The exaggeration of startle by stroke may be specific to this reflex.

Adult↗
PubMed2006

The blink reflex in "chronic migraine".

OBJECTIVES: Activation of the trigeminovascular system and sensitisation of brainstem trigeminal nuclei are thought to play an important role in migraine. The aim of this study was to investigate the blink reflex and its habituation in patients with "chronic migraine". METHODS: We studied 35 patients suffering from "chronic migraine" (IHS classification criteria) outside and during a spontaneous attack, and 35 control subjects. An EMG device with a specific habituation test program was used to elicit and record blink reflex responses and to randomly repeat stimulations at different time intervals so as to induce habituation. RESULTS: The R(1) and R(2) latencies, amplitudes and areas of the basal blink reflex were similar in patients studied both outside and during an attack as well as in control subjects, whereas the blink reflex habituation responses were markedly reduced in patients studied outside an attack. The percent changes in the R(2) areas from the baseline values, obtained when stimuli were delivered at time intervals of 10, 5, 4, 3, 2 and 1s, were statistically different (p<0.01-p<0.001) from those of the same patients studied during a migraine attack and of those of control subjects. There was a significant correlation between decreased habituation of the blink reflex and a higher frequency of attacks. The stimulus intensities of the blink reflex (multiples of the detection threshold intensities) were significantly lower (p<0.001) on the side affected, or more severely affected, by headache in patients studied during a migraine attack. CONCLUSIONS: The decreased habituation of the blink reflex outside an attack reflects abnormal excitability in "chronic migraine", which normalizes during the attacks. The inverse correlation between the frequency of attacks and habituation responses confirms the abnormal excitability induced by the high frequency of attacks. Central sensitisation mechanisms (allodynia) may explain the lower detection thresholds observed on the side affected by headache in patients during the attacks. SIGNIFICANCE: The blink reflex and its habituation may help shed light on the subtle neurophysiological changes that occur in migraine patients between and during attacks.

Adult↗
PubMed2004

Aglycemia and ischemia depress monosynaptic and polysynaptic reflexes in neonatal rat spinal cord in vitro by involving different types of 5-hydroxytryptamine receptors.

Involvement of 5-hydroxytryptaminergic (5-HT) system for aglycemia- and ischemia-induced depression of the spinal reflexes was evaluated. The monosynaptic (MSR) and polysynaptic (PSR) reflex potentials were elicited in the ventral root by stimulating the corresponding dorsal root in an isolated spinal cord from neonatal rat. Superfusion of aglycemic (glucose-free) or ischemic (glucose- and O(2)-free) solution produced a time-dependent depression of the spinal reflexes and abolished them within 35 min. The time required to produce 50% depression of the reflexes (T-50) was around 19 and 13 min for aglycemia and ischemia, respectively. Spiperone (5-HT(2A) antagonist) and ketanserin, (5-HT(2A/2C) antagonist) blocked the aglycemia-induced depression of the reflexes completely while ondansetron (5-HT(3) antagonist) attenuated it partially (as abolition times were around 50 min). Ischemia-induced depression was blocked up to 50 min by ketanserin or ondansetron but not by spiperone. In the presence of ketanserin or ondansetron, the reflexes were abolished by 60 min while in spiperone the reflexes were abolished by 30 min. In 5,7-dihydroxytryptamine treated rats, aglycemia depressed the reflexes by 45 min (greater than control, P<0.05) while the time for ischemia-induced depression was not different from the control response. The results indicate that aglycemia involves mainly 5-HT(2) receptors while ischemia involves 5-HT(3) receptors.

Action Potentials↗
PubMed2006

Soleus H-reflex modulation in response to change in percentage of leg loading in standing after incomplete spinal cord injury.

Body weight support (BWS) is frequently used to retrain standing and walking in persons post spinal cord injury (SCI), but the effects of leg unloading using BWS on H-reflex excitability in this population are unknown. The purpose of the study was to assess the effect of two different loading conditions on soleus H-reflex while standing in persons with motor incomplete SCI (i-SCI) and non-injured persons. Eight persons with motor i-SCI (mean age 50.25 years) and five non-injured persons (mean age 48.6 years) participated in the study. Specific control of the loading conditions was achieved using a BWS system. To compare reflex modulation at 40% versus 0% BWS, soleus H-reflexes were evoked during standing and normalized to the maximum M wave amplitude. These studies revealed that (1) H-reflex excitability is significantly greater after SCI and is exhibited even during quiet standing; and (2) no significant modulation in reflex excitability was observed by change in loading conditions in either the non-injured or the i-SCI subjects. These findings suggest that non-injured persons and persons with i-SCI respond similarly to bilateral limb unloading during standing with no change in H-reflex amplitude. Our results suggest that BWS of up to 40% does not produce detectable changes in the excitability of the soleus H-reflex in persons with i-SCI.

Adult↗
PubMed2006

Ptychodiscus brevis toxin-induced depression of spinal reflexes involves 5-HT via 5-HT3 receptors modulated by NMDA receptor.

The involvement of 5-hydroxytryptaminergic (5-HT) system for the Ptychodiscus brevis toxin (PbTx)-induced depression of spinal reflexes was evaluated. The reflex potentials were recorded at ventral root by stimulating the corresponding dorsal root in neonatal rat spinal cord in vitro. Superfusion of PbTx (2.8-84microM) depressed the monosynaptic (MSR) and polysynaptic (PSR) reflexes in a concentration-dependent manner. The depression of the reflexes was maximal with 84microM of the toxin. Ondansetron (0.1microM), a 5-HT(3) receptor antagonist, blocked the PbTx-induced depression of MSR and PSR. Spiperone (a 5-HT(2A) antagonist) or ketanserin (5-HT(2A/2C) antagonist and also at 5-HT(1B/1D)) failed to block the PbTx-induced depression of the reflexes. The 5-HT concentration of the cords was increased by four-fold after exposure to PbTx (28microM) and the increase was not seen in the cords pretreated with dl-2 amino-5-phosphonovaleric acid (APV, a NMDA receptor antagonist). Superfusion of 5-HT or phenylbiguanide (PBG, a 5-HT(3) receptor agonist) also produced depression of the spinal reflexes in a concentration-dependent manner. The 5-HT-induced depression of reflexes was blocked by ondansetron but not by spiperone. The results demonstrate that the PbTx-induced depression of spinal reflexes involves 5-hydroxytryptamine via 5-HT(3) receptors modulated by NMDA receptor.

Animals↗
PubMed2006

Influence of posture and stimulus parameters on post-activation depression of the soleus H-reflex in individuals with chronic spinal cord injury.

In non-disabled (ND) individuals, reflexes are modulated by influences related to physiologic state (e.g., posture, joint position, load) and activation history. Repeated activation of the H-reflex results in post-activation depression (PAD) of the response amplitude. The modulation associated with physiologic state and activation history is suppressed or abolished in individuals with spinal cord injury (SCI). While posture is known to affect H-reflex amplitude and PAD in non-disabled individuals, the effect of posture on PAD in SCI individuals is not known. Further, while the amount of PAD is also known to be influenced by the stimulus rate and by the amplitude of the evoked reflex, the interaction of posture with stimulus parameters has not been previously investigated in either group. We investigated differences in PAD of the soleus H-reflex between SCI subjects and ND subjects during sitting versus supported standing. Subjects were tested using paired conditioning-test stimulus pulses of 2.5s and 5s interpulse intervals (ISI) and with stimulus intensity adjusted to evoke reflex responses of 20% and 40% of the maximum motor response. We found standing posture to be associated with significantly less PAD in SCI subjects compared to ND subjects. In both groups, shorter ISIs and smaller reflex amplitudes were associated with greater PAD of the H-reflex. These results indicate that postural influences on post-activation modulation, while present, are impaired in individuals with chronic incomplete SCI.

Adolescent↗
PubMed2005

Computation of inverse functions in a model of cerebellar and reflex pathways allows to control a mobile mechanical segment.

The command and control of limb movements by the cerebellar and reflex pathways are modeled by means of a circuit whose structure is deduced from functional constraints. One constraint is that fast limb movements must be accurate although they cannot be continuously controlled in closed loop by use of sensory signals. Thus, the pathways which process the motor orders must contain approximate inverse functions of the bio-mechanical functions of the limb and of the muscles. This can be achieved by means of parallel feedback loops, whose pattern turns out to be comparable to the anatomy of the cerebellar pathways. They contain neural networks able to anticipate the motor consequences of the motor orders, modeled by artificial neural networks whose connectivity is similar to that of the cerebellar cortex. These networks learn the direct biomechanical functions of the limbs and muscles by means of a supervised learning process. Teaching signals calculated from motor errors are sent to the learning sites, as, in the cerebellum, complex spikes issued from the inferior olive are conveyed to the Purkinje cells by climbing fibers. Learning rules are deduced by a differential calculation, as classical gradient rules, and they account for the long term depression which takes place in the dendritic arborizations of the Purkinje cells. Another constraint is that reflexes must not impede voluntary movements while remaining at any instant ready to oppose perturbations. Therefore, efferent copies of the motor orders are sent to the interneurones of the reflexes, where they cancel the sensory-motor consequences of the voluntary movements. After learning, the model is able to drive accurately, both in velocity and position, angular movements of a rod actuated by two pneumatic McKibben muscles. Reflexes comparable to the myotatic and tendinous reflexes, and stabilizing reactions comparable to the cerebellar sensory-motor reactions, reduce efficiently the effects of perturbing torques. These results allow to link the behavioral concepts of the equilibrium-point "lambda model" [J Motor Behav 18 (1986) 17] with anatomical and physiological features: gains of reflexes and sensori-motor reactions set the slope of the "invariant characteristic," and efferent copies set the "threshold of the stretch reflex." Thus, mathematical and physical laws account for the raison d'etre of the inhibitory nature of Purkinje cells and for the conspicuous anatomical pattern of the cerebellar pathways. These properties of these pathways allow to perform approximate inverse calculations after learning of direct functions, and insure also the coordination of voluntary and reflex motor orders.

Algorithms↗