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Biomedical subjects

H Hultborn

Publications and source records attributed to H Hultborn.

At least 37 records · Page 2Linked to original sources
PubMed1994

Transmission in a locomotor-related group Ib pathway from hindlimb extensor muscles in the cat.

It has been previously shown that phasic stimulation of group I afferents from ankle and knee extensor muscles may entrain and/or reset the intrinsic locomotor rhythm; these afferents are thus acting on motoneurones through the spinal rhythm generators. It was also concluded that the major part of these effects originates from Golgi tendon organ Ib afferents. Transmission in this pathway to lumbar motoneurones has now been investigated during fictive locomotion in spinal cats injected with nialamide and L-DOPA, and in decerebrate cats with stimulation of the mesencephalic locomotor region. In spinal cats injected with nialamide and L-DOPA, it was possible to evoke long-latency, long-lasting reflexes upon stimulation of high threshold afferents before spontaneous fictive locomotion commenced. During that period, stimulation of ankle and knee extensor group I afferents evoked oligosynaptic excitation of extensor motoneurones, rather than the "classical" Ib inhibition. Furthermore, a premotoneuronal convergence (spatial facilitation) between this group I excitation and the crossed extensor reflex was established. During fictive locomotion, in both preparations, the transmissions in these groups I pathway was phasically modulated within the step cycle. During the flexor phase, the group I input cut the depolarised (active) phase in flexor motoneurones and evoked EPSPs in extensor motoneurones; during the extensor phase the group I input evoked smaller EPSPs in extensor motoneurones and had virtually no effect on flexor motoneurones. The above results suggest that the group I input from extensor muscles is transmitted through the spinal rhythm generator and more particularly, through the extensor "half-centre". The locomotor-related group I excitation had a central latency of 3.5-4.0 ms. The excitation from ankle extensors to ankle extensors remained after a spinal transection at the caudal part of L6 segment; the interneurones must therefore be located in the L7 and S1 spinal segments. Candidate interneurones for mediating these actions were recorded extracellularly in lamina VII of the 7th lumbar segment. Responses to different peripheral nerve stimulation (high threshold afferents and group I afferents bilaterally) were in concordance with the convergence studies in motoneurones. The interneurones were rhythmically active in the appropriate phases of the fictive locomotor cycle, as predicted by their response patterns. The synaptic input to, and the projection of these candidate interneurones must be fully identified before their possible role as components of the spinal locomotor network can be evaluated.

Animals↗
PubMed1994

Voltage-dependent excitation of motoneurones from spinal locomotor centres in the cat.

Lumbar motoneurones were recorded intracellularly during fictive locomotion induced by stimulation of the mesencephalic locomotor region in decerebrate cats. After blocking the action potentials using intracellular QX-314, and by using a discontinuous current clamp, it is shown that the excitatory component of the locomotor drive potentials behaves in a voltage-dependent manner, such that its amplitude increases with depolarisation. As the input to motoneurones during locomotion is comprised of alternating excitation and inhibition, it was desirable to examine the excitatory input in relative isolation. This was accomplished in spinalised decerebrate cats treated with nialamide and L-dihydroxy-phenylalanine (L-DOPA) by studying the excitatory post-synaptic potentials (EPSPs) evoked from the "flexor reflex afferents" (FRA) and extensor Ib afferents, both of which are likely to be mediated via the locomotor network. As expected, these EPSPs also demonstrate a voltage-dependent increase in amplitude. In addition, the input to motoneurones from the network for scratching, which is thought to share interneurones with the locomotor network, also results in voltage-dependent excitation. The possible underlying mechanisms of NMDA-mediated excitation and plateau potentials are discussed: both may contribute to the observed effect. It is suggested that this nonlinear increase in excitation contributes to the mechanisms involved in the production of the high rates of repetitive firing of motoneurones typically seen during locomotion, thus ensuring appropriate muscle contraction.

Action Potentials↗
PubMed1994

Disynaptic reciprocal inhibition of ankle extensors in spastic patients.

The soleus Hoffmann-reflex (H-reflex) was conditioned by a preceding stimulation of the common peroneal nerve in 74 healthy control subjects and 39 patients with spasticity in the lower extremities due to multiple sclerosis. At a conditioning-test interval of 1-3 ms a decrease of the size of the soleus H-reflex was seen in the healthy subjects. The decrease was most likely caused by disynaptic reciprocal Ia inhibition (Crone et al., 1987). In the spastic patients a similar short-latency inhibition was rarely seen. On the contrary, in several patients a facilitation was seen at a conditioning-test interval of 3-4 ms. A short-latency inhibition as pronounced as in healthy subjects was seen in four patients. These four patients did not differ from the other patients regarding the degree of spasticity or any other clinical parameter. However, they all used an external peroneal nerve stimulator daily as a walking aid. It is suggested that the lack of short-latency reciprocal inhibition reflects a deficient control of the interneurons which mediate this inhibitory spinal mechanism between antagonistic muscles in man. This might contribute to the pathophysiology of spasticity and it might be related to the frequent occurrence of co-contraction of functionally antagonistic muscles during gait in spastic patients. The existence of a pronounced reciprocal inhibition in patients receiving frequent stimulation of the peroneal nerve may suggest that regular activation of peripheral nerves is of importance for the maintenance of the activity in spinal pathways.

Adult↗
PubMed1993

H-reflexes are smaller in dancers from The Royal Danish Ballet than in well-trained athletes.

The size of the maximal H-reflex (Hmax) was measured at rest and expressed as a percentage of the maximal M-response (Mmax) in 17 untrained subjects, 27 moderately trained subjects, 19 well-trained subjects and 7 dancers from the Royal Danish Ballet. The Hmax/Mmax was significantly larger in the moderately and well-trained subjects than in the untrained subjects but smaller in the ballet dancers. It is therefore suggested that both the amount and the type of habitual activity may influence the excitability of spinal reflexes.

Adolescent↗
PubMed1992

Spinal locomotor activity in acutely spinalized cats induced by intrathecal application of noradrenaline.

In this study we present a method of intrathecal drug administration in the acute spinal cat. A thin tube introduced into the subarachnoid space below the spinal cord was used for local drug administration. Noradrenaline reduced short-latency and provoked long-latency reflexes following high threshold afferent stimulation. Topical application of noradrenaline to the spinal cord was also able to induce and maintain locomotion. Our results strengthen the notion that noradrenergic systems play an important role in motor control. In addition, the present report introduces a protocol in the acute spinal cat, which combine the benefits associated with direct application of drugs as used in vitro experiments with the advantages of using the well investigated in vivo cat spinal cord preparation.

Animals↗
PubMed1992

Induction of fos expression by activity in the spinal rhythm generator for scratching.

Fos expression was evaluated immunohistochemically in L7-S1 spinal segments after inducing fictive scratching in paralysed, unanaesthetized, decerebrate cats. The activity was induced by cutaneous stimulation of the pinna on one side and recorded from peripheral nerves. A cumulative duration of scratching of 60 to 90 min was effective in inducing fos expression. Most Fos-positive neurones were found in the dorsolateral part of the ventral horn and in the intermediate region of the spinal cord on the scratching side. In sham-operated animals the finding of Fos-positive neurones in these areas was very rare.

Animals↗
PubMed1992

Central facilitation of Ia inhibition during tonic ankle dorsiflexion revealed after blockade of peripheral feedback.

Recent studies have reported that no increase of the disynaptic reciprocal inhibition can be observed during tonic voluntary dorsiflexion of the foot as compared to rest, when the size of the control H-reflex is kept constant. Other studies have, however, shown that a voluntary contraction evokes a strong and long-lasting depression of the synaptic transmission from Ia afferents to motoneurones, most likely secondary to activation of these afferents during the contraction (post-activation depression). It was thought that this effect could also interfere with the demonstration of a central facilitation of the reciprocal inhibition during movement. The amount of disynaptic Ia reciprocal inhibition from the pretibial flexors to the soleus H-reflex was therefore estimated in normal human subjects at rest and during voluntary tonic dorsiflexion before, during and after blocking the peripheral feedback from the investigated muscles. It was observed that the reciprocal inhibition measured during dorsiflexion increased during occlusion of the blood supply to the leg, reaching a maximum of inhibition after 30 min of ischaemia. After release of the ischaemia the inhibition gradually decreased to its pre-ischaemic level. It is therefore suggested that the brain facilitates transmission in the Ia disynaptic reciprocal pathway during tonic voluntary dorsiflexion of the foot, but that this facilitation is normally not observed due to a post-activation depression following the peripheral feedback activation during the movement.

Adult↗
PubMed1992

Neuromodulation of vertebrate motor neuron membrane properties.

The short-term function of motor neurons is to integrate synaptic inputs converging onto the somato-dendritic membrane and to transform the net synaptic drive into spike trains. A set of voltage-gated ion channels determines the electro-responsiveness and thereby the motor neuron's input-output function. In addition, several of the decisive ion channels are transmitter controlled, which results in a flexible control of the input-output relationship.

Animals↗
PubMed1990

Synaptic effects on recruitment gain: a mechanism of importance for the input-output relations of motoneurone pools?

The present theoretical analysis concerns quantitative aspects of the graduation of motoneuronal recruitment. It is pointed out that variations in the relative distribution of synaptic input to low- and high-threshold cells will serve to alter the ultimate magnitude of the threshold differences between them. These threshold differences determine the pool's 'recruitment gain', i.e. the ease with which a variation of excitatory drive to the pool may alter the number of active motoneurones. In this context, not only the command signal itself but also any steady synaptic 'background' activity is of importance, whereby the background effects on recruitment gain depend on the presence of differences in intra-pool distribution between these synapses and those of the command signal. Given the appropriate intra-pool distribution, even inhibitory background effects might serve to increase the recruitment gain. It is stressed that synaptic effects on recruitment gain should be taken into account when analyzing the input-output relations of motoneurone pools.

Models, Neurological↗
PubMed1990

Sensitivity of monosynaptic test reflexes to facilitation and inhibition as a function of the test reflex size: a study in man and the cat.

In parallel experiments on humans and in the cat it was investigated how the sensitivity of monosynaptic test reflexes to facilitation and inhibition varies as a function of the size of the control test reflex itself. In man the monosynaptic reflex (the Hoffmann reflex) was evoked in either the soleus muscle (by stimulation of the tibial nerve) or the quadriceps muscle (by stimulation of the femoral nerve). In the decerebrate cat monosynaptic reflexes were recorded from the nerves to soleus and medial gastrocnemius muscles; they were evoked by stimulation of the proximal ends of the sectioned L7 and S1 dorsal roots. Various excitatory and inhibitory spinal reflex pathways were used for conditioning the test reflexes (e.g. monosynaptic Ia excitation, disynaptic reciprocal inhibition, cutaneous inhibition, recurrent inhibition, presynaptic inhibition of the Ia fibres mediating the test reflex). It was shown that the additional number of motoneurones recruited in a monosynaptic test reflex by a constant excitatory conditioning stimulus was very much dependent on the size of the test reflex itself. This dependency had the same characteristic pattern whatever the conditioning stimulus. With increasing size of the test reflex the number of additionally recruited motoneurones first increased, then reached a peak (or plateau) and finally decreased. A similar relation was also seen with inhibitory conditioning stimuli. The basic physiological factors responsible for these findings are discussed. Finally, the implications for the interpretation of experiments in man with the H-reflex technique are considered.

Adolescent↗
PubMed1989

Facilitation of quadriceps motoneurones by group I afferents from pretibial flexors in man. 2. Changes occurring during voluntary contraction.

The facilitation of the quadriceps (Q) H reflex evoked by stimulation of group I afferents from pretibial flexors exhibits biphasic changes during Q voluntary contraction. At short conditioning-test intervals the facilitation is increased, whereas it is decreased at longer intervals and/or at high conditioning stimulus intensities and/or when the contraction strength is increased. The spatial facilitation at a premotoneuronal level observed at rest on combined stimulation - common peroneal nerve (CPN) and femoral nerve (FN) - regularly disappeared during contraction. It is argued that the increase in facilitation at the onset of the weakest Q contraction reflects a descending facilitation of the interneurones mediating the CPN-induced excitation to Q MNs. An occlusion of descending and peripheral excitation in these interneurones can easily account for the depression of the facilitation when the conditioning stimulus intensity and/or the strength of the contraction is increased. However, occlusion cannot explain all the results and inhibition of the neurones mediating excitation of MNs, i.e. disfacilitation of the MNs, probably contributes to the decrease in facilitation observed during contraction.

Adult↗
PubMed1989

Possible functions of transmitter-controlled plateau potentials in alpha motoneurones.

An increasing number of vertebrate central neurones has been shown to possess complex membrane properties. However, the functional significance of such properties is unclear. The aim of the present paper is to review some old and new findings in this field from this laboratory. First, a bistability in alpha motoneurones in reduced preparations is described. Thereafter we present some new data on a bistable behaviour in motor units in unrestrained intact animals during posture. Finally, the possible role of motoneuronal bistability in locomotion and in spasticity is discussed. Recently a bistable firing behaviour in motoneurones was described in the unanaesthetized decerebrate cat. This behaviour is generated by a plateau potential, which causes long-lasting excitability increase and can be initiated and terminated by short-lasting synaptic excitation and inhibition respectively, and is contingent upon activity in descending noradrenergic and serotonergic systems. In an in vitro preparation of the turtle spinal cord the plateau potential was shown to be serotonin dependent and generated by a voltage-dependent non-inactivating calcium conductance. In order to elucidate possible functional consequences of a bistable firing behaviour in the intact animal, the firing pattern of individual soleus motor units was studied by means of chronic EMG registration in awake unrestrained rats during quiet standing. Implanted electrodes allowed the delivery of excitatory and inhibitory stimulus trains to the motoneurones. It was found that short-lasting synaptic stimulation could induce maintained shifts between two stable levels of motoneurone firing frequencies, as in the decerebrate cat. Spontaneous shifts between the same two levels were also present. It seems most likely that plateau potentials are responsible for this bistable firing property in intact animals. The role of plateau potentials in locomotion is difficult to study. At present there are no clear indications of the utilization of plateau potentials in locomotion in intact animals. However, "clamped frequency" bursts which are observed in fictive locomotion in spinal cats might be explained by plateaus. The existence of plateau potentials in motoneurones may also be of importance in spasticity. Therefore, the development of spasticity in two spinalized cats was followed for 3 weeks. Acute experiments demonstrated plateau potentials in some motoneurones in this preparation.

Action Potentials↗
PubMed1988

Distribution of recurrent inhibition within a motor nucleus. II. Amount of recurrent inhibition in motoneurones to fast and slow units.

The maximal recurrent inhibition was studied by intracellular recording from 43 triceps surae motoneurones (tentatively type-identified by the biophysical properties of the neurones) and from 67 medial gastrocnemius motoneurones (type-identified by the muscle unit properties; fast fatiguing, FF; fatigue resistant, FR and slow, S). Maximal homonymous recurrent IPSP (RIPSP) and input resistance (RN) were measured at 'resting' membrane potential and close to firing threshold. The 'synaptic current' at the peak of the RIPSP was estimated (RIPSP/RN). At 'resting' membrane potential the RIPSPs increased in the order FF less than FR less than S (0.9, 1.4, and 2.5 mV, respectively). This order was preserved when the 'current' (RIPSP/RN) rather than voltage was considered, although the overall range was much reduced. When investigated close to firing threshold there were no significant differences in synaptic 'current'. This apparent paradox may be explained by systematic differences in firing threshold between motor unit types; it increased in the order S less than FR less than FF (4.6, 8.9 and 13.5 mV, respectively).

Animals↗
PubMed1988

Distribution of recurrent inhibition within a motor nucleus. I. Contribution from slow and fast motor units to the excitation of Renshaw cells.

The relation between the size of a monosynaptic reflex (MSR) to triceps surae and the resulting Renshaw cell discharge was used to evaluate the contribution from slow and fast motor units to the excitation of Renshaw cells. It is, however, difficult to interpret these results in terms of excitation contributed by slow and fast motor units because of the following reasons. First, the size of the MSR recorded in ventral roots is not linearly related to the number of recruited motor units, since larger motor axons contribute more to the size of the MSR than smaller ones. Second, the number of spikes evoked in a Renshaw cell burst is not linearly related to the excitatory input because Renshaw cell discharge saturates in the case of large responses. The contribution of small, early-recruited motoneurones to Renshaw cell excitation is consequently overestimated. Procedures were introduced to deal with these problems. It is concluded that the last-recruited motor units (probably 'fast twitch, fast fatiguing') on average contribute four times as much excitation to Renshaw cells as the first recruited ('slow twitch') motor units.

Animals↗
PubMed1988

Maintained changes in motoneuronal excitability by short-lasting synaptic inputs in the decerebrate cat.

1. During investigation of the tonic stretch reflex in the unanaesthetized decerebrate cat we observed that a short train of impulses in Ia afferents from the soleus muscle (or its synergists) may cause a prolonged activity in the soleus muscle as judged by EMG and tension recordings. This excitability increase, which outlasted the stimulus train, could stay virtually constant during long periods (even minutes), but could be terminated at any time by a train of impulses in, for example, the peroneal nerve. 2. Gradation of the strength of stimulation and the duration of the train of impulses show that the amount of maintained excitability increase depends-within some limits-on the total amount of Ia impulses. 3. In paralysed preparations a short train of impulses in Ia afferents from any part of the triceps surae, caused a maintained increase of the efferent activity in the nerves to triceps surae and a maintained increase of the triceps surae monosynaptic test reflex. These experiments demonstrate the existence of a central mechanism (in the spinal cord and/or the brain stem), which is responsible for the maintained excitability increase seen in motoneurones to the homonymous and synergic muscles. 4. In acute spinal preparations it was not possible to demonstrate any long-lasting excitability increase by a train of Ia impulses. Following intravenous administration of the serotonin precursor 5-hydroxytryptophan, mimicking the tonic activity of these pathways in the decerebrate state, it was again possible to elicit the long-lasting excitability increase by a train of impulses in Ia afferents. A subsequent I.V. injection of methysergide (a serotonin receptor blocker) abolished the long-lasting excitability increase. This set of experiments demonstrates that the basic mechanism responsible for the maintained excitability increase is located at segmental level, and involves serotonergic systems. 5. It was demonstrated that activation of several ipsilateral and crossed reflex pathways by trains of impulses in cutaneous or high-threshold muscle afferents could trigger a maintained excitability increase of those motoneurone pools which were activated by the stimulation. Trains of stimuli to facilitatory regions in the brain stem could also cause a long-lasting excitability increase of motoneurones. Furthermore, activation of all reflex pathways which mediate postsynaptic inhibition to a motor nucleus (including recurrent inhibition via Renshaw cells) could terminate the prolonged excitability increase of that particular motor nucleus.(ABSTRACT TRUNCATED AT 400 WORDS)

5-Hydroxytryptophan↗
PubMed1988

Bistability of alpha-motoneurones in the decerebrate cat and in the acute spinal cat after intravenous 5-hydroxytryptophan.

1. In the preceding paper (Crone, Hultborn, Kiehn, Mazieres & Wigström, 1988) it was shown that a short-lasting synaptic excitation ('on' stimulus) of extensor motoneurones (primarily triceps surae) in the decerebrate cat often resulted in a maintained excitability increase, which could be reset by a short-lasting inhibitory stimulus train ('off' stimulus). In the present experiments intracellular recording from triceps surae motoneurones and the electroneurogram (ENG activity) from triceps surae nerve branches were performed in parallel. 2. Sustained firing of individual triceps surae motoneurones was most often recorded in parallel with the maintained ENG activity following a synaptic 'on' stimulus. When the motoneurone was silenced, by a hyperpolarizing current through the microelectrode, there was no sign of on-going synaptic excitation during the maintained ENG activity following an 'on' stimulus. It was therefore suggested that voltage-dependent intrinsic properties of the motoneurones themselves could be responsible for the maintained firing. 3. In confirmation of this hypothesis it was found that short-lasting depolarizing current pulses through the recording microelectrode could trigger a self-sustained firing in the motoneurone provided that the bias current (i.e. the holding potential) was kept within certain limits. Hyperpolarizing current pulses terminated the firing. When the spike-generating mechanism was inactivated (by long-lasting excessive depolarization) similar depolarizing and hyperpolarizing current pulses could initiate and terminate plateau potentials in the motoneurones. By grading the depolarizing current pulses it was found that the plateau potentials were of all-or-none character, typically around 10 mV in amplitude. The two levels of excitability which can be triggered by short-lasting excitation and inhibition of the motoneurones is referred to as 'bistable' behaviour of the motoneurones. 4. After an acute spinal transection, in the unanaesthetized cat, the bistable behaviour of the motoneurones disappeared. However, it reappears following intravenous injection of the serotonin precursor 5-hydroxytryptophan (50-120 mg/kg). 5. Individual triceps surae motor units were recorded by selective EMG electrodes during tonic stretch reflexes in the decerebrate preparations. Based on an analysis of their firing pattern during lengthening and shortening (or vibration) of the muscle it is suggested that plateau potentials in motoneurones are recruited during the tonic stretch reflex. Furthermore, it is argued that a quantitatively important part of the depolarization of motoneurones during the tonic stretch reflex indeed originates from these plateau potentials.(ABSTRACT TRUNCATED AT 400 WORDS)

5-Hydroxytryptophan↗
PubMed1988

Plateau potentials in alpha-motoneurones induced by intravenous injection of L-dopa and clonidine in the spinal cat.

1. Intracellular recordings were made from lumbar alpha-motoneurones in unanaesthetized decerebrate acute spinal cats. The response of motoneurones to direct current pulse injection or synaptic excitation was investigated following intravenous injection of L-beta-3,4-dihydroxyphenylalanine (L-DOPA, 20-120 mg/kg) alone, nialamide (10-50 mg/kg) and L-DOPA or clonidine (0.5-1 mg/kg). 2. The response properties of motoneurones were tested with rectangular and triangular current waveforms. Before L-DOPA treatment motoneuronal firing during a rectangular current pulse is characterized by an initial high firing frequency which rapidly decreases to a lower steady-state firing which is maintained only for the duration of the pulse. Following administration of L-DOPA an acceleration in firing frequency is apparent following the initial adaptation seen with rectangular current pulses. A transient after-depolarization or an after-discharge often followed the termination of the pulse. The frequency-current relation in response to a triangular current injection changed from a clockwise to a counter-clockwise hysteresis after L-DOPA treatment (i.e. after L-DOPA the firing frequency was higher for any given current during the descending phase than during the ascending phase of the triangular waveform). 3. Firing acceleration during and self-sustained firing after rectangular current pulses and counter-clockwise hysteresis of firing frequency with triangular current pulses are causally related to the presence of plateau potentials, which can be directly visualized after inactivation of the spikes. Plateau potentials in motoneurones could be generated by short-lasting intracellular depolarizing current pulses or brief excitatory synaptic inputs and terminated by short-lasting hyperpolarizing current pulses or brief inhibitory synaptic inputs. Plateau potentials were demonstrated in flexor and extensor motoneurones. 4. All bistable properties described in the preceding paragraphs following L-DOPA administration could also be seen after administration of the alpha-receptor agonist clonidine. 5. Slow rhythmic oscillations of the membrane potential (7.5-10 Hz) were seen superimposed on plateau potentials in a few cells after administration of L-DOPA and clonidine. The oscillations had an amplitude in the range 10-20 mV and represent the expression of an intrinsic property of the motoneurone. 6. It is demonstrated that plateau potentials in the motoneurones contribute to the late long-lasting reflexes observed in L-DOPA-treated spinal cats. 7. It is concluded that L-DOPA (and clonidine) change the response properties of the motoneurones in an analogous way to 5-hydroxy-DL-tryptophan (5-HTP).(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗