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H Hultborn

Publications and source records attributed to H Hultborn.

At least 73 records · Page 4Linked to original sources

Input-output relations in the pathway of recurrent inhibition to motoneurones in the cat.

1. The output from Renshaw cells caused by a phasic motor volley was investigated when these neurones were submitted to a background firing secondary to a tonic motor discharge elicited by a repetitive stimulation of muscle group I afferents. It was invariably found in individual Renshaw cells that tonic excitation produced an increase in the additional ouput caused by the phasic motor volley. The curves displaying this increase exhibited a significant 'jump' when the output resulting from the combined tonic and phasic motor discharges ranged between 2 and 5 spikes during the first 10 msec following the phasic volley. 2. The whole pool of Renshaw cells was also considered by assessing the amount of recurrent inhibition in motoneurones following a phasic motor volley. Similarly it was found that additional recurrent inhibition elicited by a phasic motor volley was enhanced when the Renshaw cells received a tonic excitatory input. 3. The Renshaw cell discharges elicited by stimulation of two different nerves were compared when a conditioning stimulus was previously applied to only one of them. The results strongly suggest that a preceding volley caused a decrease in synaptic efficacy at the terminals of the recurrent collaterals. 4. The firing produced by current injected through the recording micro-electrode was investigated in one intracellularly recorded Renshaw cell. It was found that the current-frequency curve was linear for 'steady-state' firing but displayed a clearcut sigmoid shape for the earliest intervals before the final adaptation. 5. It is demonstrated that this sigmoid input-output relation in individual Renshaw cells is sufficient to explain how the controls acting on these neurones may change the gain in the recurrent pathway when Renshaw cells are fired by a phasic motor discharge. When Renshaw cells are fired by a longlasting tonic motor discharge the linear input-output relation in individual cells should not cause any modifications of the gain in the recurrent pathway. A change in this gain secondary to the effects of segmental and supraspinal control systems is, however, possible if the motor discharge creates a subliminal fringe within the pool of Renshaw cells.

Animals↗

The neuronal pathway subserving the pupillary light reflex.

The pupillary light reflex was investigated using electrical stimulation along the pathway and recording in the short ciliary nerves. The discharge of single units in the ciliary ganglion was compared during diffuse light stimuli and electrical stimuli. It was concluded that the early reflex discharge in the short ciliary nerves following electrical stimulation on the optic tract is due to excitation of fibres active during the pupillary light reflex. The light reflex is conveyed by slow (less than 10 m/sec) optic tract fibres which synapase in the medial part of the pretectal area. In turn, pretectal neurones with conduction velocities of about 6 m/sec pass to the Edinger-Westphal complex from which the preganglionic 'pupilloconstrictor' neurones originate. Latency measurements show that there are synapses in the pretectal region and the Edinger-Westphal nucleus. The amplitude of the 'pupilloconstrictor' responses in the short ciliary nerves can be used as a measure of the excitability in the pathway of the pupillary light reflex under various conditions, e.g. following conditioning stimuli of other interacting pathways. In addition to the 'pupilloconstrictor' response there is also another short-latency discharge in the short ciliary nerves following stimulation of the posterior commissure and the Edinger-Westphal nucleus. That discharge is presumably due to activation of fibres which cause accommodation of the lens.

Animals↗

Cerebellar influence on parasympathetic neurones innervating intra-ocular muscles.

Previous work has suggested that cerebellum may control the size of the pupil and transmission in the pathway mediating the pupillary light reflex. In this work we found that electrical stimulation of the cerebellar nuclei (nucleus fastigius and nucleus interpositus bilaterally) evoked a discharge in the short ciliary nerves which innervate the intraocular muscles. The latency was 5.7 msec, which is short enough for a direct excitatory connexion from cerebellar efferents onto the preganglionic neurones in the Edinger-Westphal nucleus. These neurones are controlling the ciliary muscle of the lens system as well as the constrictor muscle of the pupil and it was therefore important to elucidate whether both of these groups participate in the discharge. Studies on interaction between the response to cerebellar stimulation and 'pupilloconstrictor' responses from optic tract fibres as well as recording from individual cells in the ciliary ganglion (identified with diffuse light stimuli) led to the conclusion that the short latency excitation is more or less confined to units controlling accommodation. Cerebellar inhibition (or disfacilitation) of longer latency affects both pupilloconstrictor and accommodation units.

Accommodation, Ocular↗

Reciprocal inhibition during the tonic stretch reflex in the decerebrate cat.

1. The aim of this study was to investigate post-synaptic reciprocal Ia inhibition during the stretch reflex; particularly the extent to which an increased Ia excitation of the Ia inhibitory interneurones will be counteracted by recurrent inhibition from motor axon collaterals. For this purpose we investigated depression of monosynaptic test reflexes antagonist flexors (reciprocal inhibition) during static stretch of quadriceps or triceps surae in unanaesthetized decerebrate cats. 3. With increasing stretch of the extensor muscle there was first a linear augmentation of reciprocal inhibition, but along with the stretch reflex in the extensor a plateau appeared in the inhibition of the flexors, although the extensor stretch reflex (judged by the e.m.g.) increased with further stretching. Within the range of stretching of triceps surae which gave increased stretch reflexes the plateau in the reciprocal inhibition was usually maintained, while during stretching of quadriceps a second phase of augmenting reciprocal inhibition often appeared. Stretch beyond the level which increased the stretch reflex activity gave augmenting reciprocal inhibition both in case of quadriceps and triceps surae. 3. Excitability measurements from central terminals of Ia afferents revealed that the increasing reciprocal inhibition during increasing stretch reflex activity in quadriceps was associated with a primary afferent depolarization in knee flexor Ia afferents; there was no corresponding effect in ankle flexor Ia afferents during stretch reflexes in triceps surae. 4. The primary afferent depolarization evoked in knee flexor Ia afferents by electrical nerve stimulation was then compared with the presynaptic inhibition of knee flexor monosynaptic test reflexes produced by the same stimuli. The results suggest that the second phase of increasing reciprocal inhibition in knee flexors is due to presynaptic inhibition and accordingly that the depth of post-synaptic reciprocal inhibition remains constant at different degrees of stretch reflex activity in both knee and ankle extensors. 5. It is postulated that during increasing stretch reflex activity the increment in Ia excitation and recurrent inhibitio; on to the Ia inhibitory interneurones almost exactly balance each other. It is suggested that recurrent inhibition of Ia inhibitory interneurones may serve as a segmental autoregulatory mechanism to keep 'alpha-gamma-linked reciprocal inhibition' at a constant depth during different levels of agonist activity.

Animals↗

Convergence on interneurones mediating the reciprocal Ia inhibition of motoneurones. I. Disynaptic Ia inhibition of Ia inhibitory interneurones.

Interneurones identified as mediating the disynaptic reciprocal Ia inhibition of motoneurones (referred to as "Ia inhibitory interneurones") were recorded in the lumbar spinal cord of the cat. It was revealed that the Ia inhibitory interneurones themselves receive disynaptic Ia inhibition. The muscles from which this inhibition is evoked are strictly antagonistic to those supplying their Ia excitation. Similar to the Ia inhibition in motoneurones the Ia inhibition in the Ia inhibitory interneurones is decreased when preceded by an antidromic stimulation of ventral roots. Furthermore, transmission of Ia inhibition to the Ia inhibitory interneurones is facilitated from ipsilateral and contralateral primary afferents as well as several supraspinal pathways analogous to earlier findings for the Ia inhibition of motoneurones. The pattern and control of the Ia inhibition of motoneurones and of Ia inhibitory interneurones display so striking similarities that it is suggested that identical interneurones are responsible. The conclusion thus emerges that "opposite" Ia inhibitory interneurones (i.e. interneurones monosynaptically connected to antagonistic muscles) are mutually inhibiting each other. The functional significance of this organization is discussed.

Afferent Pathways↗

Convergence on interneurones mediating the reciprocal Ia inhibition of motoneurones. II. Effects from segmental flexor reflex pathways.

Interneurones identified as mediating the disynaptic reciprocal Ia inhibition of motoneurones (referred to as "Ia inhibitory interneurones") were recorded in the lumbar spinal cord of the cat. Volleys in ipsilateral and contralateral high threshold muscle afferents, cutaneous and high threshold joint afferents evoked a mixture of polysynaptic excitation and inhibition. These effects were ascribed to pathways activated by flexor reflex afferents (FRA) and in addition a specific ipsilateral low threshold cutaneous pathway. Ia inhibitory interneurones excited monosynaptically from flexor nerves received stronger net excitation by volleys in ipsilateral FRA than did extensor coupled interneurones, while the opposite pattern was seen from the contralateral FRA. These patterns are similar to those found in flexor and extensor motoneurones respectivey. The FRA inhibition in Ia inhibitory interneurones was partly mediated by "opposite" Ia inhibitory interneurones, i.e. those which are mediating the Ia inhibition of Ia inhibitory interneurones. The extent to which the FRA inhibition is transmitted by Ia inhibitory interneurones was roughly estimated by its susceptibility to recurrent depression by antidromic ventral root stimulation. The main conclusion is that most segmental pathways seem to evoke their effects in parallel to motoneurones and Ia inhibitory interneurones which are monosynaptically linked to the same muscle. The functional importance of this conclusion is discussed in a following report.

Animals↗

Convergence on interneurones mediating the reciprocal Ia inhibition of motoneurones. III. Effects from supraspinal pathways.

Supraspinal effects were investigated in interneurones identified as mediating the disynaptic reciprocal Ia inhibition of motoneurones (referred to as Ia inhibitory interneurones). It was revealed that volleys in the vestibulospinal tract may evoke mono- and disynaptic EPSPs in interneurones monosynaptically excited from extensor muscles, i.e. extensor coupled Ia inhibitory interneurones. Flexor coupled interneurones instead received disynaptic inhibition. Volleys in the rubrospinal tract evoked a dominating polysynaptic excitation, usually mixed with inhibition, in flexor as well as extensor coupled interneurones. Disynaptic rubrospinal EPSPs and IPSPs were also revealed. The pyramidal tract also gives rise to a dominating polysynaptic excitation, usually mixed with inhibition, in flexor as well as extensor coupled Ia inhibitory interneurones. Rubrospinal and pyramidal volleys were shown to facilitate transmission in various segmental reflex pathways to the Ia inhibitory interneurones. A detailed comparison reveals a striking parallelism of segmental and supraspinal effects on alpha-motoneurones and Ia inhibitory interneurones connected to the same muscles. This considerably strengthens the hypothesis of an "alpha-gamma-linkage in the reciprocal inhibition".

Animals↗

Electrical constants of neurons of the red nucleus.

Membrane electrical constants have been studied in neurons of the red nucleus (RN) of the cat which were identified antidromically from the spinal cord. For each cell, the input resistance was determined from the membrane potential changes to current steps and was found to be 2.5+/-0.9 Momega in twenty five RN cells studied. In addition, linear summation of the membrane responses induced by two current pulses was demonstrated. 2. From the membrane voltage transients to current steps, the first membrane time constant, tau0, and second time constant, tau1, were determined as 5.6+/-1.0 msec and 0.6+/-0.2 msec, respectively. The ratio of the amplitudes of two exponential functions, E1/E0, was 0.18+/-0.05. A linear relation was found between the ratio of these amplitudes of exponential functions and that of the two time constants. 3. The cable parameter (electrotonic length, L) of the combined soma and dendrites of the RN neutrons was estimated as 1.1 from membrane transient data using the relation developed by Rall (1969). 4. By using this parameter, an attempt was made to estimate the location of the two excitatory inputs on the soma-dendritic membrane of RN cells.

Animals↗

Facilitation from contralateral primary afferents of interneuronal transmission in the Ia inhibitory pathway to motoneurones.

The action of volleys in contralateral primary afferents on transmission in the Ia inhibitory pathways to motoneurones was investigated with intracellular recording from motoneurones. Ia IPSPs in flexor as well as most extensor motoneurones were regularly facilitated by volleys in contralateral high threshold muscle, cutaneous and joint afferents in spinal cats under chloralose anaesthesia. In decerebrate cats with a low pontine lesion transmission in Ia inhibitory pathways was not facilitated but rather depressed by volleys in these afferents. The recurrent effects from motor axon collaterals were investigated on inhibitory transmission from different contralateral afferents to motoneurones. Previous investigations have shown that the interneurones mediating the reciprocal Ia inhibition receive recurrent inhibition via motor axon collaterals and Renshaw cells. Now a strong positive correlation was revealed between recurrent depression of IPSPs evoked from different contralateral afferents and facilitation of Ia IPSPs by the same afferent volleys. These results suggest that the recurrent depression of IPSPs from different contralateral primary afferents depends on their excitatory convergence onto the Ia inhibitory interneurones, which then partly mediate the IPSP evoked in the motoneurone from these afferents.

Animals↗

Electrophysiological study of formation of new synapses and collateral sprouting in red nucleus neurons after partial denervation.

Intracellular recording was made from cat's red nucleus after chronic lesion of the nucleus interpositus (IP) of the cerebellum and the properties of the corticorubral EPSPs were examined. 2. It was found that the time to peak of the corticorubral ESPSs induced from cerebral peduncle (CP EPSP) of chronic cats was much faster than that of normal cats. There was no simple decay following the rapid rise and early summit; instead there was another peak as if the slow normal CP EPSPs were superimposed on the fast ones. 3. By stimulating two loci, sensorimotor cortex and the cerebral peduncle, the conduction velocities of the fibers responsible for the newly appeared fast-rising component of the corticorubral EPSPs were measured. They were almost the same as those of normal corticorubral EPSPs.

Action Potentials↗