Yukari Ohki

Excitatory input to burst neurons from the labyrinth and its mediating pathway in the cat: location and functional characteristics of burster-driving neurons

Summary1. Spikes of single neurons were recorded extracellularly in the cat prepositus hypoglossi nucleus and the underlying reticular formation and were identified as type II neurons by horizontal rotation. Among these neurons, those activated by contralateral vestibular nerve stimulation with short latencies (1.5–3.0 ms) were selected for further study. 2. A class of these identified neurons was antidromically activated from the contralateral excitatory burst neuron (EBN) area immediately rostral to the abducens nucleus. Systematic tracking for antidromic stimulation revealed a wide distribution of effective spots in and near the EBN area, with varied latencies and thresholds, suggesting terminal branching in that area. The same neurons were also antidromically activated from the contralateral inhibitory burst neuron (IBN) area, the region near the midline, and the nucleus reticularis tegmenti pontis. 3. These neurons exhibited a characteristic firing pattern related to nystagmus: with contralateral rotation the firing rate gradually increased during the slow phase (type II response) and further steeply increased in a burst fashion before and during the contraversive quick phase. Since the time of occurrence of burst activity in these neurons was similar to that of contralateral ENBs and IBNs that received their axonal projection, it is suggested that they send excitatory input to burst neurons, and can thus be called burster-driving neurons (BDNs). 4. Intracellular study revealed that stimulation of the BDN area produced monosynaptic EPSPs in contralateral EBNs. The monosynaptic connection of BDNs with EBNs was confirmed by detecting unitary extracellular synaptic currents of EBNs with the spike-triggered averaging technique. 5. In contrast to BDNs, another class of nystagmus-related type II neurons in the prepositus hypoglossi and medullary reticular formation showed a discharge pattern similar to that of abducens motoneurons on the same side. None of them was antidromically activated from the contralateral pontine reticular formation including the EBN area. Some neurons responded anti-dromically to stimulation of the ipsilateral dorsomedial pontine reticular formation. 6. In conclusion, the input from the horizontal canal during rotation reaches the contralateral prepositus hypoglossi nucleus and the underlying reticular formation through the vestibular nuclei, and a class of neurons in these structures (BDNs) responds to the canal input in a burst fashion following a tonic type II activity. The axons of BDNs cross the midline and monosynaptically excite EBNs on the side of the canal stimulated. The burst activity of BDNs at the quick phase is suggested to contribute to generation of spike burst of EBNs and IBNs.

Functional identification of last-order interneurones of skin reflex pathways in the cat forelimb segments.

Premotor neurones mediating skin reflex actions onto cat forelimb motoneurones at T1 were identified by observing their monosynaptic effects on motoneurones by means of spike-triggered averaging. Both excitatory and inhibitory premotor neurones, with mono- or polysynaptic inputs from skin afferents, were identified at C7 to rostral C8, and were found mostly in laminae V-VI. They received excitatory inputs from corticospinal and rubrospinal tract fibres.

A physiological and morphological study of premotor interneurones in the cutaneous reflex pathways in cats

Implantation of an HRP-pellet into motor nuclei at the T1 segment resulted in retrograde labelling of laminae V-VII neurones densely at C6-C8 in cats. Electrophysiological experiments showed the presence of neurones in this region that received inputs from skin afferents and cortico- and rubrospinal tracts, sent axons descending in the lateral funiculus, and distributed in the motor nucleus at T1. Termination in T1 motor nuclei was also verified by intra-axonal staining with HRP. The input and output properties of these neurones indicated that they can be premotor neurones of skin reflex pathways to T1 motoneurones.

Inhibitory input to pause neurons from pontine burst neuron area in the cat

Extra- and intracellular recordings were made from pontine pause neurons (PNs) in the cat. Spontaneous spikes of PNs were suppressed after single shock stimulation of excitatory burst neuron (EBN) area immediately rostral to the abducens nucleus. The most effective stimulation site for the suppression was the region where long-lead burst neurons (LLBNs) were predominantly located. Intracellular recordings from PNs showed that stimulation of the LLBN area induced short-latency inhibitory postsynaptic potentials (IPSPs) in PNs and that steep hyperpolarization of PNs associated with quick phases of nystagmus occurred prior to an abrupt change in abducens nerve activity. Results suggest that a pause of PN spikes associated with quick phases is, at least in part, produced by inhibitory action mediated through LLBNs.

Candidate premotor neurones of skin reflex pathways to T1 forelimb motoneurones of the cat

This study explored the locations and input output properties of a large population of putative premotor neurones of skin reflex pathways in the cat. These neurones, interneurones excited by forelimb skin afferents and antidromically from the T1 motor nucleus (MN) and/or the lateral funiculus (LF, C8/T1 border), termed antidromic cells, were extracellularly recorded at C6-8. Selection of this site was based on data showing that cells retrogradely HRP labelled from the T1 MN were most numerous in C6-8 and the observation that transection of LF at the C8/T1 border abolished most skinevoked postsynaptic potentials of T1 motoneurones. Antidromic cells were located in laminae IV–V, VI and VII. The latencies of antidromic excitation ranged from 0.4 to 1.8 ms, with a tendency for laminae IV–V cells to show longer latencies than laminae VI and VII cells. Latency of skinevoked excitation ranged from 0.6 ms (IV–V cells), 0.8 ms (VI) and 1.4 ms (VII) to greater than 5 ms. The sum of the ortho and antidromic latencies (estimated central latency) of individual cells explained the central latencies of skinevoked postsynaptic potentials in T1 motoneurones. Skin-evoked firing responses (average of eight to ten cells) were earliest and largest in laminae IV–V antidromic cells, and latest and smallest in lamina VII cells. The antidromic cells also received inputs from muscle afferents and descending tracts. The following three results support the suggestion that the sampled antidromic cells are mostly premotor neurones. (1) Projection to the T1 MN via LF was verified in six laminae IV–VII antidromic cells, as tested with threshold mapping for antidromic excitation. (2) Three skinexcited axons of the middle LF projected to T1 MN, as revealed by intra-axonal staining (HRP). (3) PHA-L injection in laminae I–V of C8 anterogradely labelled terminals in lamina IX and LF axons at T1. It is suggested that last order neurones of skin reflex pathways to T1 motoneurones are widely distributed in laminae IV–VII of C6-8 and consist of a variety of neurones with different locations and input patterns.

Two stages of excitability change of pontine pause neurons in the cat

Spikes of single pontine pause neurons (PNs) in the cat were recorded extracellularly and identified by observing cessation of tonic discharges before and during the fast phase of vestibular nystagmus. PNs were antidromically activated from the excitatory burst neuron area. Antidromic spikes were abolished or their peak latencies were prolonged during particular periods within a nystagmic cycle. The excitability change in PNs indicated by alteration of the antidromic responses occurred with two stages; i.e. a moderate decrease in excitability before the onset of the fast phase and an abrupt, intense decrease starting form the beginning of the fast phase.

Characterization of premotor interneurones by their input patterns--application of principal component analysis to cat cervical interneurones.

Principal component analysis of input patterns of cat C6-C8 interneurones (300 cells) revealed that identified premotor interneurones (11 cells) activated from skin afferents and projecting to T1 motoneurones possessed a special input pattern, characterized by restricted distribution on the plane of the first (Prin 1) versus second (Prin 2) principal component (high positive values of both components). These premotor neurones were located mostly in laminae V-VI. Among other laminae V-VI cells descending in the lateral funiculus to T1 similar to such premotor neurones, there were cells distributed similarly on the Prin 1-2 plane. Further, a majority of interneurones antidromically activated from the T1 motor nucleus at low thresholds also showed a distribution on the plane similar to the premotor neurones. We suggest that premotor neurones of this input pattern constitute a major group among laminae V-VI premotor neurones projecting to T1.