No abstract

Our goal was to study potential substrates for cortical modulation of vestibular reflexes in the cat. In initial experiments, injections of wheat-germ-agglutinate-horseradish-peroxidase into Deiters' nucleus and the rostral descending nucleus revealed bilateral colonies of retrogradely filled neurons in cortical areas 6, 2, and 3a (about 60 cells per colony). In cats anesthetized with chloralose-urethane, we stimulated areas 2 and 3a with trains of pulses while recording from ipsilateral vestibular-nucleus neurons, which were characterized by their responses to sinusoidal tilts and tested for the presence of antidromic responses to stimulation of the upper cervical cord. A majority of the neurons was affected by cortical stimulation, showing either facilitation, inhibition, or a mixture of the two. Stimulation in area 2 was more effective than stimulation in area 3a. Despite the anatomic presence of direct cortico-vestibular projections, properties of facilitation and inhibition suggest that both were evoked by polysynaptic pathways. Cortical effects were broadly distributed to vestibular neurons without regard to responses of these neurons to sinusoidal tilts. There was no significant difference between effects on lateral and medial vestibulospinal tract neurons, but, as a group, vestibulospinal neurons were much more likely to be affected by cortical stimulation than neurons not antidromically activated from the C2 segment. We conclude that, by their influence on vestibulospinal neurons, neurons in cortical areas 2 and 3a should be able to modulate, in behaving animals, vestibular reflexes acting on the neck and limbs.

1. To investigate the type of vestibular signals that neurons in the caudal parts of the vestibular nuclei transmit to the cerebellum and spinal cord, we studied their responses to natural vestibular stimulation in vertical planes in decerebrate cats with the caudal cerebellum removed. Most neurons were in the caudal half of the descending vestibular nucleus, the remainder at corresponding levels of the medial nucleus or the medial-descending border. 2. Dynamics of the responses of spontaneously firing neurons were studied with sinusoidal tilts delivered at 0.05-1 Hz near the plane of body rotation that produced maximal modulation of the neuron's activity (response vector orientation). For most neurons the predominant vestibular input could be identified as coming from otolith organs (46%) or vertical semicircular canals (37%). Some neurons had otolith+canal convergence (9%) and others either had such converging input or some other form of central processing (8%). 3. Gain and phase of the responses of otolith neurons were comparable with values obtained in earlier studies on Deiters' nucleus and the rostral descending nucleus. Many canal neurons had a steeper gain slope and more advanced phase than observed previously for more rostral neurons. This may be due to more irregular afferent input to many neurons or to the absence of the vestibulocerebellum. 4. Response vector orientations of canal neurons were closely bunched near the planes of the ipsilateral vertical canals. The small number of contralaterally projecting vectors showed evidence of convergence between the two contralateral vertical canals. As is the case elsewhere in the vestibular nuclei, there was no evidence of convergence from bilateral vertical canals. Response vector orientations of otolith neurons were restricted to the roll quadrants; the majority pointed ipsilaterally. 5. Antidromic stimulation with an electrode in the restiform body or with several electrodes in the dorsal half of the white matter of the upper cervical cord was used to identify neurons projecting to the cerebellum and spinal cord, respectively. A substantial number of spontaneously firing neurons projected to the cerebellum, but there were few spontaneously active vestibulospinal neurons. The properties of the vestibular input to cerebellar-projecting neurons were the same as those of the population as a whole, but the effect of tilt on vestibulospinal neurons appeared weak or absent. 6. Many neurons were inhibited by stimulation of the restiform body. We suggest that this is mainly due to stimulation of the axons of vestibulocerebellar Purkinje cells. 7. Our results demonstrate a robust vertical vestibular input to the caudal parts of the vestibular nuclei.(ABSTRACT TRUNCATED AT 400 WORDS)

The morphometry, histochemistry, and biomechanical relationships of rectus capitis muscles were examined in adult cats. This family of muscles contained six members on the dorsal, ventral, and lateral aspects of the upper cervical vertebral column. Three dorsal muscles (rectus capitis posterior major, medius, and minor) formed a layered complex spanning from C1 and C2 to the skull. Rectus capitis posterior major was composed predominantly of fast fibers, but the other two deeper muscles contained progressively higher proportions of slow fibers. One ventral muscle, rectus capitis anterior major, was architecturally complex. It originated from several cervical vertebrae and appeared to be divided into two different heads. In contrast, rectus capitis anterior minor and rectus capitis lateralis were short, parallel-fibered muscles spanning between the skull and C1. The ventral muscles all had nonuniform distributions of muscle-fiber types in which fast fibers predominated. Dorsal and ventral muscle groupings usually had cross-sectional areas of 0.5 cm2 or more, reflecting a potential capacity to generate maximal tetanic force in excess of 9 N. Biomechanical analyses suggested that one muscle, rectus capitis lateralis, had its largest moment in lateral flexion, whereas the other muscles had large, posturally dependent moment arms appropriate for actions in flexion-extension. The observation that most rectus muscles have relatively large cross-sectional areas and high fast-fiber proportions suggests that the muscles may have important phasic as well as postural roles during head movement.

The purpose of this study was to examine the effect of bilateral labyrinthectomy on quiet stance in the freely-standing cat. Since loss of the vestibular end organs produces marked deficits in motor behaviour, including ataxia and problems with balance, we hypothesized that labyrinthectomized animals would show impairment in quantitative measures of stance. Stance was quantified in terms of the ground reaction forces under each limb and the tonic electromyographic (EMG) activity of selected muscles. Animals were labyrinthectomized by drilling into the vestibule and removing the vestibular epithelium. Following lesion, animals were able to stand unsupported on the force platform within 2 days. To our surprise, the lesioned animals showed little change in stance parameters from the control, pre-lesion state. Thus, our hypothesis of changes in stance parameters was not supported. There was no change in the distribution of vertical forces under the limbs and no increase in sway, as measured by the area of excursion of the centre of pressure over time. The horizontal plane forces, which were diagonally directed prior to lesion, became more laterally directed and larger in amplitude. The change in direction persisted even after the animals had fully compensated for the lesion, but the force amplitudes returned to control values within 10-12 days. The change in horizontal force direction was similar to that observed in normal animals that were required to stand with their paws closer than preferred in the sagittal plane (unpublished observations). EMG activity changed in some muscles but not others, and usually transiently.(ABSTRACT TRUNCATED AT 250 WORDS)