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Spinal branching of corticospinal axons in the cat.

Branching patterns of single corticospinal (CS) neurons were studied in the cat by activating these neurons antidromically from various regions of the spinal cord. 1. One hundred and ninety-three neurons were activated antidromically by microstimulation in the gray substance of the cervical cord and the majority of them were found in the forelimb area of the pericruciate cortex. 2. Branches to the lower levels of the spinal cord were found for 30% of the neurons projecting to the cervical gray matter. 3. The remaining 70% sent axons only to the cervical gray matter and some of them sent multiple branches to several segments in the cervical cord. 4. Only a few CS neurons located outside of the forelimb area could be activated from the cervical cord, but all of them also sent branches to the lower levels of the spinal cord. Neurons projecting to both the cervical cord and the lower levels were intermingled in the cortex with those projecting only to the cervical cord. 5. CS neurons activated from a given area of the cervical cord were often clustered together in a small area of the cortex, although some of these CS neurons sent their other branches to other parts of the spinal cord and neurons projecting to other parts were also intermingled among them. 6. The functional significance of multiple axonal branching of CS neurons is discussed in relation to cortical motor functions.

Animals

The effect of morphine on the activity evoked in ventrolateral tract axons of the cat spinal cord.

The effect of morphine on the activity in ventrolateral tract axons was studied in intercollicularly decerebrate cats with and without spinal section. Activity was elicited by electrical stimulation of Adelta- and C-fibres in the sural nerves. In spinal animals, morphine injected intravenously in a dose as low as 0.5 mg/kg reduced the post-stimulus discharge of impulses recorded in ventrolateral tract axons below the site of transection. The depression was not only abolished but reversed by levallorphan and naloxone. Pretreatment with reserpine did not diminish the effect of morphine. The effect of morphine was considerably weaker in decerebrate cats. Reversible block of the spinal cord produced by cold revealed that morphine reduced inhibition from the brain stem controlling the impulse transmission to ventrolateral tract axons. It is concluded that a spinal effect contributes to the analgesic action of morphine.

Afferent Pathways

Excitatory nature of dopamine in the nigro-caudate pathway.

Inputs from the substantia nigra (SN) to the caudate nucleus (Cd) and the effects of electrophoretic dopamine on Cd neurones were studied in cats anaesthetized with pentobarbitone with intracellular techniques. Single shock electrical stimulation of the SN or the medial forebrain bundle (MFB) induced monosynaptic EPSPs in Cd neurones. Dopamine depolarized Cd neurones and chlorpromazine suppressed the SN or MFB induced EPSPs. Appropriate controls indicate that the drug effects were not artifactual. Some recorded neurones were identified by procion yellow dye injection as the medium-size intrinsic Cd neurones.

Animals

Medial septal projections to the dentate gyrus of the rat: electrophysiological analysis of distribution and plasticity.

Previous electrophysiological experiments in rabbits have suggested that medial septal stimulation activates dentate granule cells and evokes an associated negative field response at the granule cell layer, without an associated "dendritic" response. Anatomical studies have suggested that septal input to the granule cells may be to stratum moleculare, or close to the cell layer, or may not exist at all. The present experiments confirmed in rats anaesthetised with urethane that medial septal stimulation elicits single action potentials from cells in the granule layer. The associated negative field potential was maximal in the granule cell layer and there was no sign of a separate dendritic potential. The fibres responsible for this potential travel to the dorsal hippocampus in the fornix superior rather than the fimbria, taking the same course as the fibres which contribute to the dense cholinesterase staining just above the granule cell layer. Stimulation at 100 Hz for 1 s of either medial septal, or perforant path, input to the dentate granule cell layer produced long term potentiation of the subsequent evoked field responses to the stimulated pathway. The responses to the non-stimulated pathway were unchanged. Paired pulse stimulation produced both homosynaptic and heterosynaptic potentiation. These data suggest that medial septal input synapses close to granule cell bodies and produces a negative field potential which is a combination of dendritic and population spike potentials. Medial septal input also appeared to produce direct activation of hilar neurones, some of which may be basket cells or other interneurones. The data also show that long term potentiation is specific to this input, perhaps dependent on presynaptic mechanisms. Paired pulse potentiation, at least in the heterosynaptic case appears to depend on postsynaptic mechanisms.

Animals

Reflex pathways from group II muscle afferents. 1. Distribution and linkage of reflex actions to alpha-motoneurones.

The interneuronally mediated reflex actions evoked by electrical stimulation of group II muscle afferents in low spinal cats have been reinvestigated with intracellular recording with motoneurones to knee flexors and ankle extensors. The results of Eccles and Lundberg (1959) have been confirmed and extended. There was wide convergence from flexors and extensors of group II excitation to flexor and group II inhibition to extensor motoneurones. Some quantitative differences in the effect from the different nerves are described. Latency measurements suggest that the minimal linkage is disynaptic in the excitatory interneuronal pathways and trisynaptic in the inhibitory pathways. Disynaptic group II EPSPs were found in 14% of the ankle extensor motoneurones but were much more common in unanaesthetized high spinal cats (Wilson and Kato 1965). From these results and corresponding ones on flexors (Holmqvist and Lundberg 1961) it is postulated that secondary afferents in addition to the weak monosynaptic connexions (Kirkwood and Sears 1975) have disynaptic excitatory pathways and trisynaptic inhibitory pathways to both flexor and extensor motoneurones. It is proposed that the group II actions of the flexor reflex pattern characterizing the anaesthetized low spinal cat are due to suppression of the inhibitory pathway to flexor motoneurones and the excitatory pathway to extensor motoneurones. In some ankle extensor motoneurones the disynaptic group II EPSPs occurred in combination with IPSPs from the FRA (including group II and III muscle afferents). The possibility is considered that these group II EPSPs are mediated by an interneuronal group II pathway with little or no input from group III muscle afferents but probably from extramuscular receptors. In other ankle extensor motoneurones group II EPSPs were combined with EPSPs from group III muscle afferents, cutaneous afferents and joint afferents. It is postulated that these group II EPSPs are mediated by an interneuronal pathway from the FRA which also supply interneuronal pathways giving inhibition to extensor or/and flexor motoneurones and excitation to flexors as postulated by Eccles and Lundberg (1959) and Holmqvist and Lundberg (1961).

Animals

Reflex pathways from group II muscle afferents. 3. Secondary spindle afferents and the FRA: a new hypothesis.

A hypothesis is forwarded regarding the role of secondary spindle afferents and the FRA (flexor reflex afferents) in motor control. The hypothesis is based on evidence (cf. Lundberg et al. 1987a, b) summarized in 9 introductory paragraphs. Group II excitation. It is postulated that subsets of excitatory group II interneurones (transmitting disynaptic group II excitation to motoneurones) may be used by the brain to mediate motor commands. It is assumed that the brain selects subsets of interneurones with convergence of secondary afferents from muscles whose activity is required for the movement. During movements depending on coactivation of static gamma-motoneurones impulses in secondary afferents may servo-control transmission to alpha-motoneurones at an interneuronal level. The large group II unitary EPSPs in interneurones are taken to indicate that, given an adequate interneuronal excitability, impulses in single secondary afferents may fire the interneurone and produce EPSPs in motoneurones; interneuronal transmission would then be equivalent to that in a monosynaptic pathway but with impulses from different muscles combining into one line. It is postulated that impulses in the FRA are evoked by the active movements and that the role of the multisensory convergence from the FRA onto the group II interneurones is to provide the high background excitability which allows the secondary spindle afferents to operate as outlined above. The working hypothesis is put forward that a movement governed by the excitatory group II interneurones is initiated by descending activation of these interneurones, but is maintained in a later phase by the combined effect of FRA activity evoked by the movement and by spindle secondaries activated by descending activation of static gamma-motoneurones. As in the original "follow up length servo" hypothesis (Rossi 1927; Merton 1953), we assume that a movement at least in a certain phase can be governed from the brain solely or mainly via static gamma-motoneurones. However, our hypothesis implies that the excitatory group II reflex connexions have a strength which does not allow transmission to motoneurones at rest and that the increase in the gain of transmission during an active movement is supplied by the movement itself. Group II inhibition. It is suggested that the inhibitory reflex pathways like the excitatory ones have subsets of interneurones with limited group II convergence. When higher centres utilize a subset of excitatory group II interneurones to evoke a given movement, there may mobilize inhibitory subsets to inhibit muscles not required in the movement.(ABSTRACT TRUNCATED AT 400 WORDS)

Afferent Pathways

Autogenetic effects of muscle contraction on extensor gamma motoneurones in the cat.

The effect of isometric twitch contractions of hind limb muscles on the discharge of gamma motoneurones has been studied in decerebrated cats with thoracic spinal cord section. Contractions of flexor digitorum longus (FDL) or gastrocnemiussoleus (GS) strongly inhibited the background discharge of their homonymous gamma motoneurones. In contrast, contraction of FDL generally caused less inhibition, or did not affect the discharge, of the synergist GS gamma motoneurones. Both the autogenetic and synergist inhibition were considerably weaker in decerebrated cats with intact spinal cords. The inhibition lasted 20 to 50 ms and occurred principally during the rising phase of contraction. It could be followed or terminated by a weaker period of facilitation during relaxation of the muscle. Shortening the muscle so that no active tension developed during contraction either abolished or substantially curtailed the inhibition. We discuss the probability that the inhibition is a spinal segmental reflex brought about by impulses generated in Ib afferents of tendon organs. The inhibition is under tonic inhibitory control in the decerebrated cat when the spinal cord is intact. Unlike the Ib inhibition of alpha motoneurones that of gamma motoneurones has a particularly strong autogenetic component.

Animals