Relationship between aerobic endurance training and dynamic cerebral blood flow regulation in humans.

The incidence of orthostatic intolerance is elevated in endurance-trained individuals. We sought to test the hypothesis that aerobic endurance training is associated with an attenuated control of the cerebral vasculature. Endurance trained (ET, n = 13) and age-matched untrained (UT, n = 11) individuals (peak O2 consumption, mean ± SEM; 63 ± 1 vs 42 ± 1 mL/min/kg, P < 0.05) were examined while supine and seated upright. Dynamic cerebral autoregulation (CA) was assessed by calculation of the rate of regulation (RoR) from the arterial blood pressure (ABP) and middle cerebral artery (MCA) mean blood velocity (V mean ) responses to a bilateral thigh cuff release, which evoked a transient hypotension. Cerebral oxygenation (oxyhemoglobin; HbO2 ) was determined with near-infrared spectroscopy. When seated upright, cuff release evoked a greater decrease in ABP (P < 0.001), MCA V mean (P = 0.096) and HbO2 (P < 0.001) in ET compared with UT. However, RoR was similar in ET and UT individuals while seated upright (to 0.193 ± 0.039 vs 0.129 ± 0.029/s, P > 0.05), and there was no significant difference in the relative change in RoR from the supine to upright positions (ΔRoR: -65 ± 7 and -69 ± 7%, for ET and UT, respectively). These findings suggest that aerobic endurance training is not associated with an attenuation in dynamic CA.

Difference between horizontal and vertical optokinetic nystagmus in cats at upright position.

UNLABELLED: The slow-phase velocity (SPV) of optokinetic nystagmus (OKN) and optokinetic after nystagmus (OKAN) in response to a velocity step of surround rotation in the horizontal direction is composed of the rapid and slow components in the cat: a rapid rise, a slow rise to a steady state, a rapid fall, and a slow decline to 0 deg/s. The rapid and slow components are attributed to the direct pathway and velocity storage neuronal mechanisms, respectively. The difference between horizontal and vertical OKN has been reported in the monkey at the upright position, but the slow and rapid components have not been distinguished. The present study compared horizontal OKN-OKAN with vertical OKN-OKAN in the cat at the upright position, distinguishing the rapid and slow components. Constant velocity rotation of a random dot pattern at a velocity of 5 to 160 deg/s was used for optokinetic stimulation. THE RESULTS: First, the amplitude of the rapid rise was relatively small in all SPV directions and all stimulus velocities investigated, with a slight upward-SPV preference to the downward-SPV (maximum 6.4, 6.0, and 3.4 deg/s in horizontal, upward, and downward SPV directions, respectively). Second, the steady state velocity was large during horizontal OKN (maximum 69.0 deg/s), small during upward-SPV OKN (12.9 deg/s), and missing (SPV is negligibly small and irregular) during downward-SPV OKN, indicating a large directional difference of OKN. Third, the acceleration of the slow rise decreased with the stimulus velocity at higher stimulus velocities >20 deg/s during both horizontal and upward-SPV OKN, suggesting strong nonlinearity in the velocity charge system. Fourth, the decay time course of the OKAN was described by the time constant of the exponential function, and the time constant was longer during horizontal (mean, 8.3 s at a stimulus velocity of 20 deg/s) than during upward-SPV (5.4 s) OKAN, suggesting that the velocity discharge system is relatively linear compared with the velocity charge system. It is concluded that horizontal OKN-OKAN is much larger than vertical OKN-OKAN in the cat at the upright position, and this directional difference is caused mainly by the directional difference in the velocity storage mechanism, but not in the direct pathway mechanism.

Purkinje cell activity in the middle zone of the cerebellar flocculus during optokinetic and vestibular eye movement in cats.

Based on the inverse dynamics theory, a previous paper reconstructed simple-spike (SS) firing rates of Purkinje cells in the cat's flocculus middle-zone by a linear-weighted summation of eye acceleration, velocity, and position during optokinetic response (OKR). The present study investigated the SS rates during combined optokinetic and vestibular stimuli of the cells recorded in the previous paper. During the sinusoidal vestibuloocular reflex (VOR) in the light (VORL) and in the dark (VORD) the firing modulation was small. During VOR suppression (VORS) by head and visual-pattern rotation in the same direction, the modulation was deep, with the peak coinciding roughly with peak ipsiversive head velocity. During VOR enhancement (VORE), the modulation was deep, with the peak coinciding roughly with peak contraversive head velocity. If we interpret these data in relation to eye and head movements, the cells in the cat were comparable to the horizontal-gaze-velocity Purkinje cells in the monkey that encode a linear summation of eye and head velocity signals. Alternatively, if we interpret the data on the basis of the inverse dynamics theory, the SS rates during VORL, VORS, and VORE were well-fitted by the OKR components of the movements (subtraction of VORD from VORL, VORS, and VORE eye movements, respectively), but not by the whole movements, using the coefficients calculated during OKR. It is concluded that the data are interpretable by both theories when the VOR gain (eye movement/head movement) is close to 1 and the firing is dominated by eye velocity information.

Motor dynamics encoding in the rostral zone of the cat cerebellar flocculus during vertical optokinetic eye movements.

The complex spike (CS) and simple spike (SS) activities of Purkinje cells in the rostral zone of the cerebellar flocculus were recorded in alert cats during optokinetic responses (OKR) elicited by a stimulus sequence consisting of a constant-speed visual pattern movement in one direction for 1 s and then in the opposite direction for 1 s. The quick-phase-free trials were selected. Ninety-eight cells were identified as rostral zone cells by the direction-selective CS activity that was modulated during vertical but not horizontal stimuli. In most of the majority population (88 cells), with an increasing CS firing rate during upward OKR and an increasing SS rate during downward OKR, the inverse dynamics approach was successful and the time course of the SS rate was reconstructed (mean coefficient of determination, 0.70 and 0.72 during upward and downward stimuli, respectively) by a linear weighted superposition of the eye acceleration, velocity, position, and constant terms, at a given time delay (mean 10 ms) from the unit response to the eye-movement response. Standard regression coefficient (SRC) analysis revealed that the contribution of the velocity term (mean SRC 0.98 for upward and 0.80 for downward) to regression was dominant over acceleration (mean SRC 0.018 and 0.058) and position (-0.14 and -0.12) terms. The velocity coefficient during upward stimuli (6.6 spikes/s per degree/s) was significantly (P<0.01) larger than that during downward stimuli (4.9 spikes/s per degree/s). In most of the minority population (10 cells), with both CS and SS firing rates increasing during upward OKR, the inverse dynamics approach was not successful. It is concluded that 1) in the cat rostral zone Purkinje cells, in which the preferred direction is upward for CS and downward for SS, eye velocity and acceleration information is encoded in SS firing to counteract the viscosity and inertia forces, respectively, on the eye during vertical OKR; 2) the eye position information encoded in SS firing is inappropriate for counteracting the elastic force; 3) encoding of eye velocity information during upward OKR is quantitatively different from that during downward OKR: SS firing modulation is larger for upward than for downward OKR of the same amplitude; and 4) encoding of motor dynamics is obscure in cells in which the preferred direction is upward for both CS and SS.

Motor dynamics encoding in cat cerebellar flocculus middle zone during optokinetic eye movements.

We investigated the relationship between eye movement and simple-spike (SS) frequency of Purkinje cells in the cerebellar flocculus middle zone during the optokinetic response (OKR) in alert cats. The OKR was elicited by a sequence of a constant-speed visual pattern movement in one direction for 1 s and then in the opposite direction for 1 s. Quick-phase-free trials were selected. Sixty-six cells had direction-selective complex spike (CS) activity that was modulated during horizontal (preferring contraversive) but not vertical stimuli. The SS activity was modulated during horizontal OKR, preferring ipsiversive stimuli. Forty-one cells had well-modulated activity and were suitable for the regression model. In these cells, an inverse dynamics approach was applied, and the time course of the SS rate was reconstructed, with mean coefficient of determination 0.76, by a linear weighted superposition of the eye acceleration (mean coefficient, 0.056 spikes/s per deg/s(2)), velocity (5.10 spikes/s per deg/s), position (-2.40 spikes/s per deg), and constant (mean 34.3 spikes/s) terms, using a time delay (mean 11 ms) from the unit response to the eye response. The velocity and acceleration terms contributed to the increase in the reconstructed SS rates during ipsilateral movements, whereas the position term contributed during contralateral movements. The standard regression coefficient analyses revealed that the contribution of the velocity term (mean coefficient 0.81) was predominant over the acceleration (0.03) and position (-0.17) terms. Forward selection analysis revealed three cell types: Velocity-Position-Acceleration type (n = 27): velocity, position, and acceleration terms are significant (P < 0.05); Velocity-Position type (n = 12): velocity and position terms are significant; and Velocity-Acceleration type (n = 2): velocity and acceleration terms are significant. Using the set of coefficients obtained by regression of the response to a 5 deg/s stimulus velocity, the SS rates during higher (10, 20, and 40 deg/s) stimulus velocities were successfully reconstructed, suggesting generality of the model. The eye-position information encoded in the SS firing during the OKR was relative but not absolute in the sense that the magnitude of the position shift from the initial eye position (0 deg/s velocity) contributed to firing rate changes, but the initial eye position did not. It is concluded that 1) the SS firing frequency in the cat middle zone encodes the velocity and acceleration information for counteracting the viscosity and inertia forces respectively, during short-duration horizontal OKR and 2) the apparent position information encoded in the SS firing is not appropriate for counteracting the elastic force during the OKR.

An anatomical substrate for the spatiotemporal transformation.

The purpose of the present experiments was to test the hypothesis that the metrics of saccades caused by the activation of distinct collicular sites depend on the strength of their projections onto the burst generators. This study of morphofunctional correlations was limited to the horizontal components of saccades. We evoked saccades by stimulation of the deeper layers of the superior colliculus (SC) in alert, head-fixed cats. We used standard stimulus trains of 350 msec duration, 200 Hz pulse rate, and intensity set at two times saccade threshold in all experiments. Evoked saccades were analyzed quantitatively to determine the amplitude of the horizontal component of their "characteristic vectors". This parameter is independent of eye position and was used as the physiological, saccade-related metric of the stimulation sites. Anatomical connections arising from these sites were visualized after anterograde transport of biocytin injected through a micropipette adjoining the stimulation electrode. The stimulation and injection sites were, therefore, practically identical. We counted boutons deployed in regions of the paramedian pontine reticular formation reported to contain long-lead and medium-lead burst neurons of the horizontal burst generator. Regression analysis of the normalized bouton counts revealed a significant positive correlation with the size of the horizontal component of the characteristic vectors. This data supports a frequent modelling assumption that the spatiotemporal transformation in the saccadic system relies on the graded strength of anatomical projections of distinct SC sites onto the burst generators.

Temporal firing patterns of Purkinje cells in the cerebellar ventral paraflocculus during ocular following responses in monkeys II. Complex spikes.

Many theories of cerebellar motor learning propose that complex spikes (CS) provide essential error signals for learning and modulate parallel fiber inputs that generate simple spikes (SS). These theories, however, do not satisfactorily specify what modality is represented by CS or how information is conveyed by the ultra-low CS firing rate (1 Hz). To further examine the function of CS and the relationship between CS and SS in the cerebellum, CS and SS were recorded in the ventral paraflocculus (VPFL) of awake monkeys during ocular following responses (OFR). In addition, a new statistical method using a generalized linear model of firing probability based on a binomial distribution of the spike count was developed for analysis of the ultra-low CS firing rate. The results of the present study showed that the spatial coordinates of CS were aligned with those of SS and the speed-tuning properties of CS and SS were more linear for eye movement than retinal slip velocity, indicating that CS contain a motor component in addition to the sensory component identified in previous studies. The generalized linear model to reproduce firing probability confirmed these results, demonstrating that CS conveyed high-frequency information with its ultra-low firing frequency and conveyed both sensory and motor information. Although the temporal patterns of the CS were similar to those of the SS when the sign was reversed and magnitude was amplified approximately 50 times, the velocity/acceleration coefficient ratio of the eye movement model, an aspect of the CS temporal firing profile, was less than that of the SS, suggesting that CS were more sensory in nature than SS. A cross-correlation analysis of SS that are triggered by CS revealed that short-term modulation, that is, the brief pause in SS caused by CS, does not account for the reciprocal modulation of SS and CS. The results also showed that three major aspects of the CS and SS individual cell firing characteristics were negatively correlated on a cell-to-cell basis: the preferred direction of stimulus motion, the mean percent change in firing rate induced by upward stimulus motion, and patterns of temporal firing probability. These results suggest that CS may contribute to long-term interactions between parallel and climbing fiber inputs, such as long-term depression and/or potentiation.

Dependence of short-latency ocular following and associated activity in the medial superior temporal area (MST) on ocular vergence.

Motion of a large-field pattern elicits short-latency ocular following responses (OFR) in the monkey, which are mediated at least in part by the medial superior temporal area of the cortex (MST). The magnitude of the OFR is known to be inversely related to viewing distance, and we investigated the dependence of OFR and the associated neuronal activity in the MST on a major cue to viewing distance, ocular vergence, in alert monkeys (Macaca fuscata). The vergence angle, expressed in terms of the apparent viewing distance, ranged from infinity to 16.6 cm (0-6 m(-1)). The magnitude of the initial OFR increased monotonically with increases in convergence at a mean (+/-SD) rate of 19.6+/-4.5%/m(-1) in four monkeys (over the range 0-4 m(-1)). In two monkeys, we recorded the single unit activity of 160 MST neurons that responded to motion of a large-field pattern with directional selectivity. The mean latency (+/-SD) of the MST discharges elicited by large-field motion was 50+/-7.5 ms (n=115), which preceded the onset of OFR by an average of 10+/-9.9 ms. The discharge modulation elicited by large-field motion showed a significant dependence on vergence in 91/160 neurons (57%), 72 of which (79%) increased their firing rate with increasing convergence ("near" neurons), and the remainder increasing their firing rate with decreasing convergence ("far" neurons). However, on average, the sensitivity of these MST neurons to vergence was only about 30% of that shown by the OFR. It could be that only those neurons that are very sensitive to vergence angle contribute to the OFR, but it is also possible that much of the modulation of OFR with vergence occurs downstream from the MST or in alternative pathways (yet to be discovered) that contribute to OFR.

Visual inputs to cerebellar ventral paraflocculus during ocular following responses.

The activity of cerebellar Purkinje cells was recorded in the ventral paraflocculus (VPFL) of alert monkeys during ocular following responses induced by brief movements of the visual scene. The mossy fiber input evoked 'simple-spikes' which increased their activity during either ipsiversive or downward motion of the visual scene. On the other hand, the 'complex-spikes' evoked by the climbing fiber input increased their activity during either contraversive or upward movement. To further define the sources of the visual input to the VPFL, we recorded single units in the dorsolateral pontine nucleus (DLPN), which is known to project to the VPFL as mossy fibers, and in the pretectal nucleus of the optic tract (NOT), which is known to project to the inferior olive whose neurons project to the cerebellum as climbing fibers. In both areas, most neurons were directionally selective, and responded to the moving visual scene with very short latencies (approximately 40-60 ms). These results suggest that both DLPN and NOT neurons deliver information concerning movements of the visual scene to the VPFL through different pathways.

Neuronal mechanisms of two-dimensional orienting movements in the cat. I. A quantitative study of saccades and slow drifts produced in response to the electrical stimulation of the superior colliculus.

To evaluate the metrics of rapid eye movements caused by the activation of distinct collicular microzones, the superior colliculus (SC) was electrically stimulated in alert behaving cats while their heads were restrained. A quantitative study of electrically induced rapid eye movements demonstrated that their amplitude and direction depended on the intensity of stimulation, the electrode location, and the initial position of the eyes, while their duration depended on the intensity of stimulation. When detailed quantitative procedures are employed, properties of saccades produced in response to the electrical stimulation of the feline SC resemble those of saccades elicited in response to the electrical stimulation of a variety of primate brain areas. Besides saccades, electrical stimulation of the feline SC gave rise to slow drifts whose amplitude and direction was also influenced by the initial position of the eyes. Because their size depended on the frequency of stimulation and their time course reflected mechanical properties of the oculomotor plant, induced slow drifts could be due to a more or less direct projection of the SC onto extraocular motoneurons. A model that includes such a variety of connections between the SC and extraocular motoneurons is presented and is shown to produce realistic combinations of fast and slow eye movements when its input is a step function of time. The present findings support the notion that an orbital mechanical factor underlies the eye position sensitivity of slow drifts and saccades evoked in response to the electrical stimulation of the SC.

Site of interaction between saccade signals and vestibular signals induced by head rotation in the alert cat: functional properties and afferent organization of burster-driving neurons.

1. Extracellular spikes of burster-driving neurons (BDNs) were recorded within and immediately below the prepositus hypoglossi nucleus in the alert cat. BDNs were characterized by short-latency activation after stimulation of the contralateral vestibular nerve (latency: 1.4-2.7 ms) and the ipsilateral superior colliculus (latency: 1.7-3.5 ms). Convergence of vestibular and collicular inputs was found in all of 85 BDNs tested. Firing of BDNs increased during contralateral horizontal head rotation and decreased during ipsilateral rotation. A burst of spikes was induced in association with contralateral saccades and quick phases of nystagmus. 2. BDNs showed irregular tonic discharges during fixation. There was no significant correlation between the firing rate during fixation and horizontal or vertical eye position in most BDNs. During horizontal sinusoidal head rotation, the change in firing rate was approximately proportional to and in phase with contralateral head velocity. The phase lag of the response relative to head angular velocity was 13.8 +/- 20.1 degrees (mean +/- SD) at 0.5 Hz and 7.2 +/- 13.5 degrees at 0.2 Hz on the average. The gain was 0.88 +/- 0.25 (spikes/s)/(degrees/s) at 0.5 Hz and 1.19 +/- 0.49 (spikes/s)/(degrees/s) at 0.2 Hz. 3. Quantitative analysis of burst activity associated with saccades or quick phases indicated that the ON direction of BDNs was contralateral horizontal. The number of spikes in the burst was linearly related to the amplitude of the contralateral component of rapid eye movements. The slope of regression line was, on the average, 1.14 +/- 0.48 spikes/deg. There was no significant difference between the mean slopes for saccades and quick phases. The number of spikes depended on the difference between initial and final horizontal eye positions and not on the absolute eye position in the orbit. The mean burst firing rate was proportional to the mean velocity of the contralateral component of rapid eye movements. The slope of the regression line was 0.82 +/- 0.34 (spikes/s)/(degrees/s). Significant correlation was also found between intraburst instantaneous firing rate and instantaneous component eye velocity. 4. Objects presented in the contralateral visual field elicited a brief burst of spikes in BDNs independent of any eye movement. Contralateral saccades to the target were preceded by an early response to the visual stimulus and subsequent response associated with eye movement. 5. Excitation of BDNs produced by stimulation of the ipsilateral superior colliculus was facilitated by contralateral horizontal head rotation. Therefore saccadic signals from the superior colliculus to BDNs may be augmented by vestibular signals during head rotation.(ABSTRACT TRUNCATED AT 400 WORDS)

Orienting-related eye-neck neurons of the medial ponto-bulbar reticular formation do not participate in horizontal canal-dependent vestibular reflexes of alert cats.

Ponto-bulbar reticular formation neurons, including identified reticulospinal neurons, were studied in alert, head-fixed cats. Orienting-related neurons of "eye-neck" type (ENNs) were selected on the basis of qualitative correlations of their discharges with visually triggered eye saccades and electromyographic activity (EMG) of dorsal neck muscles. It was tested whether ENNs participate both in visually triggered gaze shifts requiring eye-head coordination and in gaze-stabilizing movements, such as vestibulo-ocular and vestibulo-collic reflexes (VOR, VCR). Firing patterns were studied during passive sinusoidal rotation (0.2-1.0 Hz; 2.0-21.5 deg peak-to-peak) in the horizontal plane. Responses to electrical stimulation of the superior colliculus and the vestibular nerve were recorded to assess the convergence of tectal and vestibular synaptic inputs. The same methods were applied to a control sample of neurons with discharges apparently "unrelated" to orienting movements. ENNs did not show any modulation of firing rate correlated to compensatory VOR or VCR during passive sinusoidal rotations. Among "unrelated" cells, the fraction of modulated units was close to that reported for reticular neurons projecting in the medial reticulospinal tract. Phasic and sustained components of ENN bursts were associated with anticompensatory movements induced by rotation, such as quick phases, ocular beating field shift, and the increase of EMG activity in neck muscles acting in the direction of passive rotation. Monosynaptic excitation from the contralateral superior colliculus was observed in 92.3% of ENNs, but only 2 out of 17 tested showed an excitatory response to vestibular nerve stimulation. In the control group of "unrelated" neurons the proportions of monosynaptic tectal and excitatory vestibular nerve inputs were, respectively, 75.6 and 71.4%. It is concluded that ENNs are specifically related to active gaze shifts, derived from either visual or from head velocity inputs. Rhombencephalic connections of vestibular nuclei to these neurons appear to be quite weak. Parallel inputs from the mid- or forebrain must be assumed to explain their firing patterns during rotation-induced anticompensatory gaze shifts. Within the studied range of frequencies and amplitudes of passive rotation, ENNs did not participate in the vestibulo-collic reflex. It is therefore unlikely that reticular neurons controlling orienting eye-neck synergies act also as a premotor pathway for gaze-stabilizing movements.

Post-spike facilitation of neck EMG by cat tectoreticulospinal neurones during orienting movements.

1. The activity of fourteen tectoreticulospinal neurones (TRSNs) was recorded intraaxonally in the caudal pons of alert cats during orienting movements towards visual stimuli. TRSN spikes were used to compute the spike-triggered average (STA) of rectified EMG of dorsal neck muscles. 2. Eight TRSNs for which 400-2532 spikes were available were analysed with the STA technique. When the STA was computed from all spikes, significant post-spike facilitation (PSF) was obtained for six of eighteen cell-muscle pairs investigated (5 TRSNs). The mean relative amplitude of PSFs was 7.4% (S.D. 3.7). The onset latencies ranged from 1.1 to 5.0 ms and mean duration was 11.4 +/- 3.1 ms (mean +/- S.D.). 3. Interspike interval distributions were unimodal, with modes between 2.7 and 12.7 ms. Spike trains of TRSNs that produced significant PSFs contained 5-13% of the interspike intervals < or = 5 ms and 22-37% of the intervals < or = 10 ms. To evaluate the contribution of short intervals to PSF, STAs were computed separately for spikes preceded by 'short' (< or = 5 or < or = 10 ms) and 'long' (> 5 or > 10 ms) intervals. 4. When computed from spikes preceded by 'long' intervals, PSF amplitudes were small (mean +/- S.D., 5.3 +/- 2.7%) and onset latencies measured by cusum ranged between 2.4 and 5.4 ms. This is longer than the estimated minimal latency of monosynaptic facilitatory effect on neck EMG (1.9-2.1 ms). 5. Relative amplitudes of PSF obtained with spikes preceded by 'short' intervals were much larger (mean +/- S.D., 14.8 +/- 7.4%), but cusums indicated negative latencies for four of six PSFs. The unrealistically short onset latencies could be accounted for by the summation of facilitation from the trigger spike with that of the preceding spikes. In four of five TRSNs a large increase of PSF amplitude (from 3.2 to 7.2 times the amplitude obtained from 'long' intervals) suggests the presence of frequency-dependent potentiation of synaptic transmission. 6. This study unequivocally demonstrates that some TRSNs produce significant post-spike facilitation of neck motoneurones. This facilitation could be mediated by monosynaptic tectomotoneuronal connections although a contribution by disynaptic connections cannot be definitively ruled out. The high instantaneous firing rates of TRSNs produce a potentiation of the otherwise weak facilitatory action of TRSNs that presumably contributes to a rapid recruitment of motoneurones during initiation of head orienting movements.

Anatomical evidence for ipsilateral collicular projections to the spinal cord in the cat.

Injections of WGA-HRP were made within the C1 segment of spinal cord in cats with a midsagittal section of the midbrain. A small number of labelled cells were found in the latero-caudal part of the deeper layers of the superior colliclus (SC) ipsilateral to the injection sites. Because of the complete section of the dorsal tegmental decussation, these results definitively demonstrate the existence of an ipsilateral tecto-spinal pathway projecting to upper cervical segments in the cat. Ipsilaterally projecting tecto-reticulo-spinal neurons represent about 5% of the total population of tecto-spinal neurons. They were exclusively located in the deeper collicular layers and most of them were found in the latero-caudal part of the SC. Comparison with our previous studies suggests that more ipsilateral tecto-spinal projections that found after the section of the dorsal tegmental decussation probably exist. They may arise from tecto-reticulo-spinal neurons recrossing the midline in the brainstem or in the rostral part of C1. By analogy with the cortico-spinal tract, we suggest that the existence of an ipsilateral tecto-spinal pathway can be regarded as evidence for a substantial development of the cat tecto-spinal system as compared with other mammals.

Tracing premotor brain stem networks of orienting movements.

Methods allowing a direct matching of movement-related firing patterns and connectivity of individual neurons have been used in the analysis of premotor networks controlling orienting movements. Advances have been made in the description of coding properties of orienting-related tectal output neurons, as well as in specifying their distributed connections in the brain stem and possible modes of coupling to saccadic pattern generators in the reticular formation. New data on the properties of signals and connectivity patterns have also been obtained for the tecto-recipient reticulo-spinal neurons. At least a small portion of the network performing the spatio-temporal transformations of orienting-related tectal efferent signals can now be described both in functional and in morphological terms.

Stimulation of the caudate nucleus induces contraversive saccadic eye movements as well as head turning in the cat.

The effects of stimulation of the caudate nucleus were investigated in alert cats, with special reference to the induction of eye and head movements. Stimulation of caudal portions of the caudate nucleus on one side with trains of current pulses induced gaze shifts towards the contralateral side. When the head of the animal was restrained, the majority of evoked eye movements were single conjugate saccades. The amplitude and direction of the evoked saccade varied depending on the initial eye position. The amplitude of the horizontal component tended to be larger for saccades initiated from more ipsilateral positions, and became gradually smaller as the initial eye position shifted to the contralateral side. If the eye was far into the contralateral positions, no saccades were induced. Furthermore, the saccades tended to have a downward component when the eye was initially focused upward, and an upward component when the eye was focused downward. When the head was made free to move, the same stimulation induced a sequence of contraversive staircase gaze shifts composed of coordinated eye and head movements. The eye movements in the orbit resembled nystagmus, consisting of contraversive saccades followed by reverse compensatory movements. The head turning, though smooth and continuous, was also suggested to consist of a series of movements coupled with saccadic eye movements. This study indicates a potential role of the caudate nucleus in the control of orienting reflexes.

Role of premotor vestibular nucleus neurons in vertical gaze.

The firing properties and projection patterns of secondary vestibular nucleus neurons involved in the vertical vestibulo-ocular pathways were investigated in alert cats. Recordings were made in the medial longitudinal fasciculus (MLF) from axons that were monosynaptically activated from the vestibular nerve. Many identified axons discharged in relation to vertical eye movements. The majority of these axons increased their firing rate for downward eye position (DPVs). During pitch rotation, the firing rate of DPVs was also related to upward head velocity, suggesting that they received monosynaptic input from the posterior canal. DPVs could be divided into two groups on the basis of their firing regularity. There was a tendency for regular DPVs to have a higher firing rate, a higher correlation for the rate-position relationship, and a larger phase lag and a smaller gain re head velocity than irregular DPVs. Spike-triggered average method and intraaxonal HRP techniques demonstrated that ipsilaterally projecting (i-) DPVs made inhibitory connections with up-on extraocular motoneurons, and contralaterally projecting (c-) DPVs made excitatory connections with down-on motoneurons. Virtually all i-DPVs were of regular type, while c-DPVs included both regular and irregular types. Stimulation of the caudal MLF at the level of the obex indicated that all the irregular c-DVPs and some of the regular c-DPVs had a collateral to the spinal cord, while none of the regular i-DPVs had such a collateral.

Vertical eye movement-related secondary vestibular neurons ascending in medial longitudinal fasciculus in cat I. Firing properties and projection pathways.

1. The firing characteristics and projection patterns of secondary vestibular nucleus neurons involved in the vertical vestibuloocular pathways were investigated in alert cats. Single-unit recordings were made in the medial longitudinal fasciculus (MLF) near the trochlear nucleus from axons that were monosynaptically activated after electrical stimulation of the vestibular nerve. In a total of 253 identified secondary neurons, 225 discharged in relation to vertical eye movements; 189 of these increased their firing rate for downward eye movements and 36 for upward movements. The activity of the remaining 28 axons was not related to eye movements when the head was still. 2. Virtually all of the secondary neurons with downward on-direction displayed tonic activity that was primarily related to steady eye position during fixation (DPV neurons). The slope of the relationship between firing rate and vertical eye position ranged from 1.2 to 9.1 (spikes/s)/deg with a mean of 3.2 (spikes/s)/deg. The regularity of firing was quantified by calculating the coefficient of variation (CV) of interspike intervals. A comparison of the CV in the population units indicated that DPV neurons could be classified as either regular or irregular neurons. There was a tendency for regular neurons to have higher firing rates and higher correlation coefficients for the rate-position relationships than irregular neurons. 3. During pitch rotation in the light, all the DPV neurons tested increased their firing rate with upward head rotation. Both the phase and the amplitude of the response indicated that DPV neurons discharged not only in relation to eye position but also in relation to head velocity, suggesting that they received monosynaptic input from the posterior semicircular canal. The gain and phase lag of the response relative to head velocity were measured at 0.5 Hz. The range of the gain was 1.1-5.1 (spikes/s)/(deg/s), and that of the phase lag was 18.3-62.4 degrees. There was a tendency for irregular DPV neurons to have a larger gain and smaller phase lag than regular DPV neurons. 4. Ascending and descending projection pathways were determined for 147 DPV axons. Of these, 69 ascended in the contralateral MLF with respect to their soma (crossed-DPV axons), and 78 in the ipsilateral MLF (uncrossed-DPV axons), as revealed by their monosynaptic activation from the contralateral or ipsilateral vestibular nerve. Stimulation of the caudal MLF at the level of the obex evoked direct responses caused by antidromic activation of descending collaterals in approximately 70% (49/69) of the crossed-DPV axons.(ABSTRACT TRUNCATED AT 400 WORDS)

Vertical eye movement-related secondary vestibular neurons ascending in medial longitudinal fasciculus in cat. II. Direct connections with extraocular motoneurons.

1. The preceding study in the alert cat has shown that many secondary vestibular axons that ascend in the medial longitudinal fasciculus (MLF) increase their firing rate in proportion to downward eye position. In the present study, projection and termination of these downward-position-vestibular (DPV) neurons within extraocular motoneuron pools were studied electrophysiologically by spike-triggered averaging techniques and morphologically be reconstructing their axonal trajectory after intra-axonal injection of horseradish peroxidase (HRP). 2. Extracellular field potentials recorded within the trochlear nucleus and/or the inferior rectus subdivision of the oculomotor nucleus were averaged by the use of spike potentials of single DPV neurons as triggers. All the crossed-DPV axons tested induced negative unitary field potentials in the trochlear nucleus (n = 9) and in the inferior rectus subdivision of the oculomotor nucleus (n = 5), suggesting that they made monosynaptic excitatory connection with motoneurons in these nuclei. The four crossed-DPV axons tested in the two motoneuron pools induced unitary field potentials in both. The majority of crossed-DPV axons terminated in these nuclei were directly activated from the caudal MLF, indicating that they had descending collaterals projecting to the spinal cord as well. The uncrossed-DPV axons did not induce such unitary field potentials either in the trochlear nucleus (n = 4) or in the inferior rectus subdivision (n = 3). 3. All the uncrossed-DPV axons examined (n = 14) induced positive unitary field potentials in the superior rectus subdivision of the oculomotor nucleus, suggesting that they made monosynaptic inhibitory connections with motoneurons innervating the superior rectus muscle. These uncrossed-DPV axons displayed regular firing patterns and were not activated from the caudal MLF. None of the crossed-DPV axons tested (n = 4) induced unitary field potentials in the superior rectus subdivision. 4. Five crossed-DPV axons were injected with HRP. All these axons ascended in the MLF contralateral to their soma, gave off many collaterals to the trochlear nucleus, and projected more rostrally. For three well-stained axons, numerous terminal branches were also found in the rostroventral part of the contralateral oculomotor nucleus, the area corresponding to the inferior rectus subdivision. Some collaterals in the oculomotor nucleus recrossed the midline to terminate in the medial part of the ipsilateral oculomotor nucleus. Other terminations were observed in the interstitial nucleus of Cajal and in the periaqueductal gray adjacent to the oculomotor nucleus. The crossed axons injected included both regular and irregular types, and three of the four examined were activated from the caudal MLF.(ABSTRACT TRUNCATED AT 400 WORDS)