W. Precht

Afferent projections to the cerebellar flocculus in the pigmented rat demonstrated by retrograde transport of horseradish peroxidase.

SummaryThe horseradish peroxidase (HRP) retrograde transport method was used to identify brainstem afferents to the cerebellar flocculus in the pigmented rat. Injections of the enzyme were made through recording microelectrodes, making it possible to localize the injection site by physiological criteria. Clearly, the largest number of afferents arise from the bilateral vestibular and perihypoglossal nuclei and from the contralateral dorsal cap (of Kooy) of the inferior olive. Additionally, a substantial number arise bilaterally from: (1) the nucleus reticularis tegmenti pontis (NRTP); (2) several of the cranial motor nuclei including the abducens, retrofacial and facial nuclei and the nucleus ambiguus; (3) the rostral part of the lateral reticular nucleus (subtrigeminal nucleus); (4) the raphe pontis and raphe magnus and (5) neurons intercalated among the medial longitudinal fasciculus (MLF) just rostral to the hypoglossal nucleus and another group rostral to the abducens nucleus.The basilar pontine nuclei contained a large number of lightly labeled neurons in all flocculus injections which were discretely located within the dorsolateral, lateral and medial divisions. These areas were labeled bilaterally but with a slight contralateral preponderance. Injection into the flocculus, but involving the adjacent ventral paraflocculus, produced a heavier labeling of pontine neurons with a slightly different distribution. Therefore, we tentatively conclude that the flocculus receives input from these pontine visual centers (dorsolateral, lateral and medial nuclei), perhaps through collateral projections from neurons projecting to the paraflocculus.The present study demonstrates strong similarities between the rat and other species studied (e.g., rabbit, cat, monkey) in terms of the brainstem nuclei projecting to the flocculus. Most noticeable in quantitative terms are the pathways known to mediate vestibular (vestibular and perihypoglossal nuclei) and visual (optokinetic) information (e.g., NRTP). Additionally, we can provide morphological evidence that the midline and paramedian pontine tegmentum, identified in the cat and monkey as containing saccade-related neurons, send large numbers of projections to the rat flocculus. Given these similarities, the rat may be a suitable animal model in which to study the pathways underlying visual-vestibular interaction and saccadic mechanisms in the flocculus.

Initial, rapid phase of recovery from unilateral vestibular lesion in rat not dependent on survival of central portion of vestibular nerve

The behavioral effects of vestibular endorgan lesions were compared with those of vestibular ganglion lesions in the albino rat. No differences in head tilt angle or spontaneous eye nystagmus beat frequency were noted between the two groups during the first 36 h after the lesion was made. Of rats studied beyond 36 h, 2/7 with lesions restricted to the endorgans and 2/3 with ganglion lesions showed pronounced secondary increases in head tilt and tonic eye deviation, but not eye nystagmus. Single units were recorded in the ganglion acutely, as well as 1,2, and 14 days after an endorgan lesion was made. Practically no resting activity could be recorded in the ganglion acutely (2-7 h) after endorgan damage, and the resting activity at subsequent times was slight. It is concluded that an intact vestibular ganglion isolated from the sensory periphery is of no functional significance during the first 36 h, when the largest decreases in magnitude of the behavioral signs of unilateral labyrinthectomy occur in the rat. A slight significance at later times is not ruled out.

Horizontal optokinetic ocular nystagmus in the pigmented rat

Horizontal optokinetic nystagmus was elicited in rats by rotation of a pattern of bright dots projected onto a cylinder surrounding the animal. Eye position was measured with the electromagnetic search coil technique. Optokinetic stimuli consisted either of velocity steps of pattern rotation or sinusoidal oscillations. Closed-loop gain (slow phase eye velocity/pattern velocity) of steady-stage step responses in binocular vision ranged between 0.8 and 1.0 for pattern velocities up to 20-40 degrees/s and decreased thereafter. Open-loop gain (steady-state slow phase velocity/retinal slip velocity) was dependent on retinal slip velocity and decreased linearly in double logarithmic plot from about 30 (at 0.5 degree/s) to about 9 (at 5 degrees/s). For retinal slip velocities larger than 5 degrees/s open-loop gain decayed faster and reached about 1 at 30 degrees/s. Step response profiles showed a gradual increase in slow phase eye velocity reaching steady-state after a time period roughly proportional to stimulus velocity. Initial slow phase velocity measured within 500 ms after stimulus onset reached between 2 and 4 degrees/s and was largely independent of stimulus amplitudes above 10 degrees/s. Occasionally rats showed fast rises in slow phase eye velocity at the onset of the step response profiles. Primary and secondary optokinetic afternystagmus were present. Duration of primary afternystagmus was largely independent of stimulus amplitude and lasted 8.0 +/- 4 s. Closed-loop gain of steady-state step responses in monocular vision was, for temporonasal stimuli, similar to that measured in binocular condition while for nasotemporal stimulation gain was much smaller even at low stimulus velocities. Sinusoidal modulation of slow phase velocity was linearly dependent on stimulus velocity; the linear range decreased as frequency of stimulation increased. Slow phase velocity gain was relatively constant (ca 0.8) between 0.05 and 0.3 Hz and showed only a small tendency to decrease at larger stimulus frequencies. Phase-lag increased strongly with stimulus frequency and could be fitted by assuming a response time delay of 100 ms. The results show that the rats optokinetic system is qualitatively similar to that found in another lateral-eyed species, namely the rabbit. At a quantitative level, however, both fast and slow optokinetic response dynamics appear to be better developed in the rat than in the rabbit.(ABSTRACT TRUNCATED AT 400 WORDS)

Vestibular nerve and nuclei unit responses and eye movement responses to repetitive galvanic stimulation of the labyrinth in the rat

SummaryTwo-second cathodal current pulses were applied at one-minute intervals at a point external to the round window in the ear of each albino rat subject. Responses were recorded in the vestibular nerve ganglion, the vestibular nuclei (single units), or in the eye movements (search coil recording method) of anaesthetized, decerebrated, or alert rats. The unit responses to the galvanic stimuli were characterized and compared with responses to galvanic and rotational stimuli reported in the literature. The main focus of the study, however, was effects of stimulus repetition. In both the vestibular nerve and vestibular nuclei recordings, the responses of many units were substantially larger or smaller at the end of a 13-pulse stimulus train than at the beginning. In the vestibular nuclei, but not in the nerve, there was a slight bias towards a decrease in response magnitude, with 10/88 units showing decreases great enough to be considered as reflecting an habituation process. In contrast, the eye movement responses showed more consistent response decrements, especially in the alert condition, but also in the other conditions (none of the unit recordings were done in alert rats). It is concluded that some of the modifications underlying habituation of the vestibuloocular reflex probably occur in portions of the neuronal reflex pathways that are downstream from the vestibular nuclei.

The horizontal optokinetic nystagmus in the cat

Summary1)Horizontal optokinetic eye nystagmus (OKN) and afternystagmus (OKAN) were recorded in the alert cat (head restrained) in response to velocity steps and sinusoidal optokinetic stimuli. 2)A strong dependency of OKN performance on stimulus pattern was found: responses were most regular and gain was high over a large range of stimulus velocities when the stimulus consisted of a high-contrast random dot pattern. 3) Following velocity steps, OKN showed a small amplitude fast rise in slow phase velocity (SPV) which was followed by a slow build-up to steady state. The amplitude of the initial jump in SPV increased with stimulus amplitude up to 30°/s and saturated afterwards. The plateau level of initial SPV ranged from 5 to 15°/s. 4) The slow build-up of SPV showed non-linearities, i.e. the time to steady state increased with stimulus amplitude and the slow rise of SPV was irregular. In most animals steady state SPV showed no signs of response saturation for step amplitudes up to 60–80°/s or more. The open-loop gain (steady state SPV/ retinal slip velocity) dependend on retinal slip velocity and decreased from 46 at 0.5°/s to 0.4 at about 60°/s. 5) OKAN I and II were consistently observed and occasionally OKAN III was noted. OKAN I durations (mean 13.8 +- 5.1 s) and OKAN II amplitudes were independent of stimulus magnitude. Initial SPV of OKAN I was typically the same as that of OKN, i.e. no fast fall was observed. Cessation of pattern rotation in light, however, produced a fast initial decay of SPV. 6) A least square fitting of OKAN time course was performed with various time functions. The SPV of OKAN I and II was best fitted with a damped sine wave, indicating that cat optokinetic system behaves like a second order underdamped system. 7) Sinusoidal stimuli produced strong response non-linearities. At a given frequency gain decreased with increasing stimulus amplitudes. Gain correlated best with stimulus acceleration. In addition, strong stimuli produced characteristic response distortions. 8) In the visual-vestibular conflict situation vectorial summation of VOR and OKN was observed only with small stimuli.

Increased projection of ascending dorsal root fibers to vestibular nuclei after hemilabyrinthectomy in the frog

SummaryThe projections from brachial, ascending dorsal root fibers were studied autoradiographically in controls and chronically (four months) hemilabyrinthectomized frogs. Comparison showed that projections into the partially denervated vestibular nuclear complex of chronically hemilabyrinthectomized animals were far more dense than in control animals. In the cerebellar granular layer, no obvious difference in the extent of dorsal root projections was observed between both groups of animals. Cerebellar areas such as the auricular lobe and the dorsal rim, which normally receive many terminals from vestibular but not from dorsal root afferents, were not invaded by dorsal root fibers in chronically hemilabyrinthectomized frogs.

An electrophysiological study of pathways mediating optokinetic responses to the vestibular nucleus in the rat

Summary1)Intra-and extracellular responses of neurons in the pretectum (Pt), the nucleus reticularis tegmenti pontis (NRTP), the prepositus hypoglossal complex (NPH) and the vestibular nuclei (VN) were recorded during orthodromic/antidromic stimulation of their afferent/efferent fibers.2)In the Pt, many neurons were excited by stimulation of the contralateral optic nerve (ONc). Comparison of the latencies of evoked presynaptic action potentials and EPSPs yielded a time difference corresponding to one synaptic delay. Forty five per cent of these monosynaptically driven neurons were also excited antidromically from the ipsilateral NRTP.3)In the NRTP, ONc and Pt stimulations evoked disynaptic and monosynaptic EPSPs, respectively. Thirty six per cent of NRTP neurons orthodromically driven from ONc and/or ipsilateral Pt stimulation were also antidromically invaded from either the contralateral (67%) or the ipsilateral (33%) flocculus but never from both.4)In the NPH, both ipsilateral Pt and NRTP stimulations excited type II neurons monosynaptically. In addition, EPSPs evoked by Pt stimulation could be mediated to the NPH via a disynaptic route involving the NRTP.5)In the VN, type II neurons were excited by ipsilateral Pt stimulation. When comparing the latencies of action potentials and EPSPs evoked by Pt stimulation in the NPH and in VN type II neurons respectively, a short, possibly monosynaptic connection, may be postulated between the NPH and the VN.6)Our results suggest that vestibular neurons may be optokinetically driven from the contralateral eye both via Pt-NPH connections and Pt-NRTP-NPH paths. They also confirm the existence of a transcerebellar route from the Pt via the NRTP to the ipsior contralateral flocculi.

Anatomy and Physiology of the Optokinetic Pathways to the Vestibular Nuclei in the Rat

Neurons in the vestibular nuclei (Vn) have been shown in a variety of species to respond not only to vestibular but also to pure optokinetic stimuli, i.e. rotation of large visual patterns (cf. ref. Precht, 1981). Vestibular and optokinetic inputs are synergistic and expand the working range of Vn (Keller, Precht, 1979). In this context, the vestibular nuclei may be considered as an important premotor structure having direct and indirect access to ocular and spinal motoneurons. At the behavioral level the importance of the transvestibular optokinetic path is stressed by the findings that optokinetic nystagmus (OKN) and afternystagmus (OKAN) are severely affected by bilateral (Cohen et al., 1973) and unilateral (Maioli et al., 1982) labyrinthectomies for long periods of time. More specifically, the transvestibular path has been considered part of the velocity storage mechanism or indirect path which, after the initial fast rise (direct path), provides the additional slower rise of OKN slow phase velocity to steady state values during prolonged stimulation (Cohen, et al. 1977).

Recovery of some vestibuloocular and vestibulospinal functions following unilateral labyrinthectomy

Publisher Summary Numerous studies on the mechanisms of functional recovery from unilateral peripheral vestibular lesions have been performed, making the vestibular lesion model one of the best-studied neural recovery models. Undoubtedly, the long-lasting interest in this model was initiated by the observation that the severe symptoms, which are seen immediately after the removal of one vestibular labyrinth, abate rather quickly with time. Besides the presence of an easily demonstrable recovery of function, the vestibular lesion model offers several other features, which make it an attractive model for the study of neural plasticity. Following unilateral labyrinthectomy, the central nervous system has to cope with two major groups of motor deficits. Firstly, the symmetrical tonic influence exerted by the vestibular receptors via vestibular afferents on posture of the head, body and in the resting condition is altered by unilateral withdrawal of resting activity in vestibular nerve fibers. The second and equally important task of the nervous system consists in the adjustment of the lesion-induced gain deficits of vestibular reflexes during the head and body movements. This chapter discusses on the behavioral and single unit work performed in mammals, mainly the rat and cat. Emphasis is on the results obtained with the vestibuloocular system since it has been studied in much detail, particularly, as far as its dynamic performance is concerned. However, the vestibulospinal system will be dealt with in conjunction with the recovery of head posture after lesion.

On the role of vestibulo-ocular reflex plasticity in recovery after unilateral peripheral vestibular lesions

SummaryAlthough adaptive plasticity is a wellknown feature of the vestibulo-ocular reflex (VOR), deficits in VOR performance after unilateral labyrinthectomy are poorly compensated in a large percentage of cats. To assess whether VOR plastic capabilities are affected by labyrinthectomy, forced oscillation in front of a patterned surround was imposed in unilaterally labyrinthectomized cats. This experimental paradigm has been shown to be very effective in inducing adaptive VOR gain changes in intact animals. We demonstrate that plasticity of VOR gain is still present both in acute and chronic stages following vestibular lesions. By contrast, forced oscillation did not significantly alter the lesion-induced asymmetry of responses. We conclude that VOR gain control mechanisms are not used to their fullest possible extent in a large percentage of animals suffering unilateral vestibular damage.