Topographic studies on medial reticular nucleus.

Several distinct classes of neurons have been identified in the medial reticular nucleus of the medulla and pons and in proximity thereto. Neurons projecting down the spinal cord comprised the principal class with two subclasses according as the neurons did or did not receive monosynaptic inputs from the fastigial nuclei of the cerebellum. Two other classes were recognized accordings as they projected to the cerebellum or rostrally to the mesencephalon. Topographic planar maps giving the location of these neurons have been constructed by exploring the nucleus with series of microelectrode tracks in parasagittal or in transverse planes. The different classes of neurons were not arranged in large discrete nuclei. In part they appeared to be randomly distributed, but many colonies of one or another class of neurons could be recognized with 3-11 neurons in zones with dimensions of a millimeter or so. Because of the limitations of sampling by microelectrode tracks at spacings of 0.5 mm, single colonies might have an actual population of 100 or more. Many of the class of neurons projecting to the cerebellum were in the region of the perihypoglossal nucleus. However, almost as many were located deep in the medial reticular nucleus. None was found at the pontine level. Reticulospinal neurons with fast axonal conduction velocities tended to be located dorsally to those with slow velocities. Correlation with the findings of Ito et al. leads to the conjecture that the neurons with fast axons are excitatory, while those with slow axons are primary inhibitory neurons. There is a brief reference to the problems raised by the admixture of the various neuronal classes, there being discrete colonies immersed in a scattered arrangement of all classes.

Somatotopic studies on red nucleus: spinal projection level and respective receptive fields.

The somatotopic inputs into red nucleus (RN) neurons have been studied with special reference to their level of projection in the spinal cord. As inputs we employed either volleys in predominantly cutaneous nerves of forelimb and hindlimb or cutaneous mechanoreceptor discharges evoked by taps to footpads of forelimb and hindlimb. There has been physiological confirmation of the anatomical findings that RD neurons projecting to the lumbar cord are located in the ventrolateral zone of the pars magnocellularis, whereas in the dorsomedial zone are RN neurons with cervical but not lumbar projection. Somatotopically there was found to be a differentiation of input to RN neurons according as they projected to the lumbar or only to the cervical cord. This finding was presented in the form both of tables and of somatotopic maps. As expected, this discrimination was more restrictive for the more selective inputs from pad taps than for nerve inputs. Nevertheless, forelimb inputs often had a considerable excitatory and inhibitory action on lumbar-projecting RN neurons, and vice versa for cervical-projecting neurons. There were two notable somatotopic findings that suggest specificities of connectivities. First, despite the large convergence of IP neurons onto RN neurons (about 50-fold), the degree of somatotopic discrimination was about the same for interpositus and RN neurons with two testing procedures: between inputs from forelimb and hindlimb; and between inputs from pads on one foot. Second, although there was in the interpositus nucleus a considerable topographical admixture of neurons with dominant forelimb or hindlimb inputs, the axonal projections of these neurons were apparently unscrambled on the way to the target RN neurons, so as to deliver the somatotopic specificities observed for two classes of RN neurons; those projecting down the spinal cord beyond L2 level, and those projecting to C2 but not L2. Finally, there is a general discussion of motor control with reference to the pathway; pars intermedia of anterior lobe of cerebellum leads to interpositus nucleus leads to red nucleus leads to rubrospinal tract leads to spinal motoneurons.

Responses of red nucleus neurons to antidromic and synaptic activation.

An account is given of the responses of 432 red nucleus (RN) neurons with axons projecting down the spinal cord. Almost half were in an initial series of 18 experiments on anesthetized cats, and the remainder were in a second series of 12 experiments on decerebrate unanesthetized cats. The differences between the two series were of little significance. All recording was from single neurons using extracellular glass microelectrodes that were inserted throught the right superior colliculus and directed to the right red nucleus at a standard orientation. Identification of RN neurons was both by location, checked by subsequent histology, and by antidromic invasion from the spinal cord. The spinal stimulating electrodes were placed in proximity to the left rubrospinal tract at C2 and L2 segmental levels. Axonal conduction velocities were calculated from the latency differential between the L2 and C2 antidromic responses and were usually in the range 60-130 m/s, 97% of all neurons located in the red nucleus had axons projecting to the C2 level, and 37% projected to the L2 level. The responses of 229 RN neurons were observed with stimulation applied to the contralateral (left) interpositus nucleus. In 10 (5%) there were antidromic responses to both interpositus and C2 stimulation, a finding in good agreement with the anatomical description of rare axon collaterals from rubrospinal fibers to the interpositus nucleus. In 209 (91%), there was a clear monosynaptic excitation. The impulse generation was at a latency usually of 1.0-1.8 ms, which a modal value of 1.4 ms. The afferent inputs to RN neurons were provided by stimulation either of predominantly cutaneous nerves in all four limbs or of cutaneous mechanoreceptors of the contralateral forelimb and hindlimb...

The differential effect of cooling on responses of cerebellar cortex.

1. Responses of the cerebellar cortex in anaesthetized cats were evoked by mossy fibre and/or climbing fibre inputs, and the effects of graded cooling of the cerebellar cortex were investigated. Cooling was applied either globally by flooding the exposed cortex with cooled Ringer Locke, or in later experiments locally be passing cooled fluid through a silver tube in contact with the cerebellar cortex. The cortical temperature was continuously monitored by a thermistor inserted to a depth of 0.5 mm close to the recording site. 2. In the granular layer the cooling caused a large increase in the diphasic P1N1 wave generated by the afferent mossy fibre volley. The waves generated by synaptic excitation and discharge of granule cells, N2P2, were not diminished until the temperature fell towards 20 degrees C. In contrast the N3 wave of the molecular layer was largest with cooling in the range of 35 to 25 degrees C, often several times larger than at 38 to 40 degrees C. Associated with the enhanced N3 wave there was an enhanced N4 wave, which indicates an increased discharge by Purkyne cells. 3. Climbing fibre inputs generate a negative field potential in the molecular layer due to the powerful excitation of Purkyne cells. In contrast to the N3 potential this climbing fibre wave was largest at the higher temperatures 35-40 degrees C and declined progressively with cooling, being usually suppressed at moderate coolings of 31-27 degrees C. Intracellular recording revealed that the diminution was due both to the elimination of all but the first impulses of the normal burst discharge of the climbing fibre impulses and to the diminution of the synaptic excitation of a single climbing fibre impulse. 4. It is shown that the negative potentials produced in the molecular layer by combinations of mossy fibre and climbing fibre inputs can be very effectively distinguished by this differential effect of cooling. 5. The effects of cooling even to a severe level are immediately recoverable on warming. Repeated cooling has no untoward effects and there is no sign of the hysteresis reported for the cuneate nucleus. 6. There is a discussion of the factors that could cause cooling to differentiate between the actions of the mossy fibre and climbing fibre impulses on Purkyne cells.

Reticulospinal neurons with and without monosynaptic inputs from cerebellar nuclei.

An account is given of the responses of 557 medial reticular neurons with axons projecting down the spinal cord. All 30 experiments were on decerebrated unanesthetized cats paralyzed by Flaxedil. Recording from single neurons was by extracellular glass microelectrodes. Identification was first by location (confirmed by subsequent histology) in the medial reticular nucleus of medulla or pons, and second by antidromic activation from cord stimulation at C2 and L2 segmental levels. Axonal conduction velocities were calculated from the latency differential between L2 and C2 antidromic responses, and were usually in the range of 90-140 m/s; but about 25% were slower, ranging down to 30 m/s. Stimulation by electrodes in the ipsilateral and contralateral fastigial nuclei differentiated reticulospinal neurons into two classes according to whether they did or did not receive monosynaptic inputs, the respective populations of fully investigated neurons being 270 and 174. The fastigioreticular neurons were distinguished by a higher background frequency with mean values of 28 as against 15/s. There were also significant diffences in both the excitatory and inhibitory responses to afferent volleys from forelimb and hindlimb nerves. Comparison of the respective latency histograms showed that the responses of neurons with a fastigial input had an excess of latencies in the ranges that can be correlated with the latency histograms observed for fastigial responses. Thus, there is evidence for the effectiveness of the fastigial input and so for the pathway with monosynaptic linkage: Purkinje cells of cerebellar vermis yields fastigial neurons yields medial reticular neurons projecting down the spinal cord. Adequate stimulation of cutaneous receptors by pad taps and air-jet stimulation of hairy skin in a disppointingly small action when compared with fastigical responses. Explanations of this deficiency are suggested. Another discrpancy from the fastigial responses is that the medial reticular neurons have much wider receptive fields with little discrimination between ipsilateral and contralateral and between forelimb and hindlimb. Stimulation of the ipsilateral tegmental tract was tested on 183 reticulospinal neurons, 112 being with fastigial inputs. In about half there was a powerful monosynaptic excitation, which would identify such neurons as being on the pathway from mesencephalic and diencephalic centers to the spinal cord. There is a general discussion of transmission across successive synaptic relays, where specificity is sacrificed to integration.

Medial recticular neurons projecting Rostrally.

1. In the latter part of investigations on the medial reticular neurons, stimulation was applied to the ipsilateral tegmental tract in the upper pontine level. Of the 426 neurons in this series, 56 had rostrally projecting axons as evidenced by their antidromic responses. 2. Of these 56 neurons, 41 were activated by limb nerve stimulation, usually very strongly, and so qualify as units in the reticular activating system. Fastigial stimulation monosynaptically excited 14 of these reticular activating neurons, so providing and ascending pathway from the cerebellar vermis. 3. Axonal branching has been demonstrated by antidromic testing in four reticular neurons with ascending axons. In two there were branches to the fastigial nucleus and in two, down the spinal cord. 4. The latency histograms of the excitatory and inhibitory responses resembled those for reticular neurons with descending axons except for the poverty of inhibitory responses in neurons not receiving a fastigial projection.