Role of basal ganglia and ectostriatum in the context-dependent properties of the optocollic reflex (OCR) in the pigeon (Columba livia): a lesion study

The possible participation of basal ganglia and associated structures [dorsal striato-pallidum, nucleus spiriformis lateralis (SpL), ectostriatum] in the elaboration of the optocollic reflex (OCR) was investigated by making bilateral chemical lesions (ibotenic acid). Previous data have shown that both the slow and fast phases of the OCR are dependent on the behavioural context. The slow phase velocity (SPV) and the peak velocity of fast phases obtained in non-flying pigeons ('resting condition') were enhanced in pigeons in which a flying posture was experimentally provoked ('flying condition'). Therefore, the effect of lesions was analysed in pigeons standing in the 'resting' or 'flying' condition. In the 'resting' as in the 'flying' condition, all the lesions provoked a decrease in SPV, which augmented with the stimulation velocity. Velocity step stimuli revealed greater OCR deficits than velocity ramp stimuli. Extensive lesions (including the striato-pallidum, ectostriatum and a part of the neostriatum), as well as SpL lesions, provoked a greater SPV decrease over a longer time than lesions restricted to the striato-pallidum or the ectostriatum. The peak velocity of fast phases was only reduced by the 'extensive lesion' in the 'flying condition'. The present data show that the basal ganglia system is involved in the elaboration of optokinetic responses and suggest that, to work in an optimal range, the optokinetic centres need to receive integrated information from basal ganglia in addition to direct visual input.     

Vestibulo-ocular reflex (VOR), cervico-ocular reflex (COR) and its interaction in active head movements

Summary In normal adults the vestibulo-ocular reflex (VOR) and the cervico-ocular reflex (COR) were investigated during passive and active head or body movements, respectively. Sinusoidal rotations around the vertical axis of the body at frequencies of 0.05, 0.1, and 0.2 s^–1 and total amplitudes of 20°, 40°, 60°, or 80° were employed.The average eye deviations (Schlagfeld) during VOR were directed opposite to the direction of the head turning. During COR, however, slow eye deviations of higher amplitude were anticompensatory relative to the fixed head. During active head turnings the average eye deviations showed the same anticompensatory direction as in COR, but were still larger. They increased with stimulus amplitudes up to 60°.At least a weak cervical nystagmus was elicited in all subjects, with its fast phases beating in the direction of the relative head movement. Its gain reached marked values up to 0.5, but only for peak stimulus velocities below 25°/s. The nystagmus gain during active head turnings was only slightly higher than during VOR.With higher stimulus velocities, large anticompensatory saccades appeared just before the change of stimulus direction; these are typical for active head movements, but were also found during COR.Supported by Sonderforschungsbereich Hirnforschung und Sinnesphysiologie (SFB 70) der Deutschen Forschungsgemeinschaft (DFG)     

Vestibulo-ocular reflex (VOR), cervico-ocular reflex (COR) and its interaction in active head movements

In normal adults the vestibulo-ocular reflex (VOR) and the cervico-ocular reflex (COR) were investigated during passive and active head or body movements, respectively. Sinusoidal rotations around the vertical axis of the body at frequencies of 0.05, 0.1, and 0.2s-1 and total amplitudes of 20 degrees, 40 degrees, 60 degrees, or 80 degrees were employed. The average eye deviations (Schlagfeld) during VOR were directed opposite to the direction of the head turning. During COR, however, slow eye deviations of higher amplitude were anticompensatory relative to the fixed head. During active head turnings the average eye deviations showed the same anticompensatory direction as in COR, but were still larger. The increased with stimulus amplitudes up to 60 degrees. At least a weak cervical nystagmus was elicited in all subjects, with its fast phases beating in the direction of the relative head movement. Its gain reached marked values up to 0.5, but only for peak stimulus velocities below 25%. The nystagmus gain during active head turnings was only slightly higher than during VOR. With higher stimulus velocities, large anticompensatory saccades appeared just before the change of stimulus direction; these are typical for active head movements, but were also found during COR.     

Optokinetic eye and head movements in the unrestrained cat

Eye and head movements of the unrestrained and alert cat were measured with the search coil technique. Optokinetic responses were elicited by a moving random dot pattern. With the head free, slow phase gaze velocity (SPGV) during optokinetic nystagmus (OKN) was not faster than slow phase eye velocity (SPEV) with the head fixed neither with binocular nor with monocular stimulation. Cats with the head free also showed a directional asymmetry of SPGV during monocular stimulation with higher gain values for slow phases from temporal to nasal as compared to slow phases from nasal to temporal. Eye, head and body movements contributed to SPGV to a varying degree. Although SPGV remained rather constant (comparable to the head-fixed condition), the relative contributions of head velocity and eye velocity could fluctuate strongly. During the resetting saccades, the head ended its fast phase later than the eye (102 ± 46 ms). During this period, the eye compensated the remaining part of the fast phase of the head and simultaneously executed an already new slow phase of gaze. It is discussed whether the cerebellar flocculus might provide a system to achieve constant gaze during OKN.     

The effect of cervical and vestibular reflexes on eye movements in Huntington's chorea

In 8 patients with manifest Huntington's Chorea vestibulo-ocular (VOR) and cervico-ocular (COR) reflexes were compared with eye movements during active head turnings. Seated patients were stimulated with their eyes closed by sinusoidal swings around the vertical axis at frequencies of 0.05, 0.1 and 0.2s-1 with amplitudes of 20, 40 and 60 degrees. 1) With all stimuli and in all patients a weak nystagmus was elicited in the direction of head movements, superimposed on larger slow eye deviations. 2) The averaged total saccadic amplitudes were smaller than in normals, increased with stimulus amplitudes and were smallest for COR, followed by VOR and active head movements. 3) The gain (peak velocity of slow phase of nystagmus to peak stimulus velocity) was only slightly below norm values and decreased with increasing stimulus frequency and amplitude. 4) The peak amplitudes of average slow eye deviations increased with stimulus amplitudes. In VOR they were comparable to norm values but were below them during COR and active head movements. 5) In normal subjects these slow eye deviations were compensatory to head movements in VOR but anticompensatory in COR and during active head movements. In choreic patients during COR and more often during active head movements these slow eye movements were compensatory for the head turning.     

Vestibulo-ocular and optokinetic impairments in left unilateral neglect

Right brain damaged patients affected by left unilateral neglect (N+) typically fail to explore the contralesional space. For the first time, this study investigates the dynamic and spatial features of the horizontal vestibular-ocular response (VOR), the optokinetic response (OKR) and the VOR-OKR interaction in six N+ and in five right brain damaged patients without neglect (N-). No lateral asymmetry of the gain (i.e. eye velocity to head velocity ratio) of VOR slow phases was found in either group. In the VOR, N+ had higher frequency of slow-rightward/fast-leftward phases and higher contralesional shift of the beating field (i.e. orbital position of fast phases). In the VOR-OKR, there was an increase of gain in both lateral directions and in both groups even though in N-, there was a lower phase shift between eye and head velocity. In contrast to the VOR, in the VOR-OKR, N+ had higher frequency of slow-leftward/fast-rightward phases. The VOR-OKR interaction also introduced an ipsilesional shift of the beating field in both N+ and N-. In the OKR, N+ showed a drop in the velocity, amplitude and frequency of slow-rightward/fast-leftward phases. These findings potentially suggest that each hemisphere modulates VOR with contralaterally directed slow phases and OKR with ipsilaterally directed slow phases. This organisation could facilitate maintenance or fast recovery of combined VOR + OKR after unilateral brain damage. The same findings suggest that by inducing slow-leftward phases, vestibular and optokinetic stimulation improve left side neglect through the activation of different hemispheric pathways. No ipsilesional deviation of the subjective "straight ahead" was found in N+. These results show that chronic unilateral neglect can be dissociated both from deficits of ipsilesionally directed VOR and from ipsilesional deviation of the subjective midsagittal plane of the body.     

Autistic traits modulate cortical responses to affective but not discriminative touch

The sense of touch is primarily considered a discriminative and exteroceptive sense, facilitating the detection, manipulation and exploration of objects, via an array of low‐threshold mechanoreceptors and fast conducting A‐beta (Aβ) afferents. However, a class of unmyelinated, low‐threshold mechanoreceptors identified in the hairy skin of mammals have been proposed to constitute a second, anatomically distinct system coding the affective qualities of touch. Unlike Aβs, which increase their firing rate linearly with the velocity of a stimulus moving across their receptive field, the response of these C‐tactile afferents (CTs) is described by an inverted ‘U’ curve fit, responding optimally to a skin temperature stimulus moving at between 1 and 10 cm/s. Given the distinct velocity tuning of these fast and slow touch fibres, here we used event‐related potentials to compare the time course of neural responses to 1st (fast) and 2nd (slow) touch systems. We identified a higher amplitude P300 in response to fast, Aβ‐targeted, versus slow CT‐targeted, stroking touch. In contrast, we identified a previously described, C‐fibre specific, ultra‐late potential (ULP) associated with CT‐targeted input. Of special note as regards the function of CTs is that the amplitude of the ULP was negatively correlated with self‐reported levels of autistic traits, which is consistent with the hypothesized affective and social significance of this response. Taken together, these findings provide further support for distinct discriminative and affective touch systems and suggests the temporal resolution of EEG provides an as yet underutilized tool for exploring individual differences in response sensitivity to CT‐targeted touch.     

Eye instability induced by vestibular stimulation in rabbits.

The slow compensatory phases of the vestibulo-ocular reflex (VOR) in the rabbit tend to drift and the drift reverses the direction. This periodic alternating drift (PAD) has two peculiar characteristics: (1) it is induced by sinusoidal vestibular stimulation in naive animals, being evoked immediately after stimulus onset and persisting after the end of stimulation; (2) the peak velocity and period of the drift are dependent on stimulus amplitude. PAD of the rabbit has strong similarities with PAN, a periodic alternating nystagmus observed in humans with cerbellar disorders and in monkeys after nodulo-uvulectomy, although its peak velocity is smaller. It is hypothesized that PAD is due to a slight instability, caused by vestibular stimulation in darkness, of the cerebellar adaptive loop, which exerts a variable gain control on the time constant of the velocity storage integrator.     

Orientational asymmetries in small-field optokinetic nystagmus in human infants

Optokinetic nystagmus (OKN) is a pattern of reflexive eye movements which occurs when portions of the visual field are in continuous motion. Gratings moving at 7 deg/s either horizontally (left or right) or vertically (up or down) were presented on a viewing screen subtending 30° by 22°. Horizontal (HOKN) and vertical (VOKN) OKN were recorded under binocular viewing conditions from infants and adults. Eye movements were recorded by means of an infrared corneal reflection eye movement recorder. OKN to horizontal and vertical stimuli was different in pattern for infants. Infants' HOKN was of significantly higher frequency and lower amplitude than their VOKN. Infants below 4 months of age also showed an asymmetry within VOKN between upward and downward stimulus motion, with markedly lower gains and more variable slow phase following movements for downward moving stimuli. No differences were found in HOKN to the right and left. There was also no evidence of a build-up of slow phase velocity over time. Infants' fast phases showed peak velocity/amplitude relationships like those of adults, and like those of infant saccades recorded in a previous study of infants' fast eye movements. Across all directions of stimulus movement, infants had lower slow phase gains and OKN frequencies, and larger slow phase amplitudes than adults. The characteristics of infants' OKN are discussed in relation to those observed in other species and in adult clinical patients with eye movement disorders.     

The role of compensatory eye and head movements for gaze stabilization in the unrestrained frog

Compensatory eye, head and gaze movements of unrestrained frogs were recorded simultaneously in response to table movements in the light. Passive displacement was compensated with a gain between 0.55 and 0.85, depending on stimulus amplitude. At small stimulus amplitudes gaze was stabilized exclusively by compensatory eye movements. At larger stimulus amplitudes compensatory head movements contributed up to 80% gaze stabilization. The contribution of compensatory eye movements became increasingly more restricted to those brief transient periods, at which head velocity changed only slowly in response to a change in stimulus direction or velocity. The wave forms of both eye and head movements exhibited characteristic and complementary distortions. Their combination, the gaze wave form compensated almost exactly in phase for the imposed passive displacement in space. Head saccades of small amplitude were rather well compensated by fast eye movements in the opposite direction, with the result that the combined gaze movement was smooth. The occurrence of these compensatory fast eye movements depended neither upon the function of the labyrinthine organs nor upon retinal image slip.     

Participation of different groups of afferent fibers of mesenteric nerves in vasomotor reflexes]

The dependence of the magnitude and character of vasomotor reflexes on the amplitude of mesenteric nerve tetanic stimulation was studied in experiments with cats anesthetized with urethane and chloralose. A comparison of the stimulus amplitude reflex magnitude curves with the previously obtained data on electrical excitability of various groups of mesenteric nerves afferent fibres revealed that there are 3 groups of "vasomotor" afferents in mesenteric nerves: "fast" Adelta-fibres (conduction velocity above 8 m/s) the impulses of which evoke depressor or small pressor reflexes; "slow" Adelta-fibres (conduction velocity below 8 m/s) the impulses of which evoke pressor reflexes or interacting with impulses of lower-threshold "fast" Adelta-fibres, either decrease depressor reflexes evoked by impulses of these fibres or increase pressor reflexes evoked by these fibres impulses; C-fibres the impulses of which increase pressor reflexes evoked by impulses of "slow" Adelta-fibres.     

Vestibulo-ocular reflex (VOR) induced by passive head rotation and goal-directed saccadic eye movements do not simply add in man

Additivity between vestibulo-ocular reflex (VOR) and saccadic eye movements was quantified in man by passively rotating the subject's head as he tracked a stepping target. A systematically increased gaze (i.e. eye + head) saccadic velocity was observed when the head was rotated toward the target, as compared to a head-fixed condition, indicating that VOR and saccades do not fully add. Moreover, although VOR was assumed to be inhibited during ocular saccades, mean gaze saccadic amplitude remained unchanged. This would suggest an on-line computation of gaze position to be fed back to the saccadic system in order to stop the saccade once gaze reaches its goal.     

Action-potential initiation and maintained activity of the isolated frog muscle spindle

The initiation of afferent action potentials represents the basic signalling process integrating and coding information of an external stimulus. There is also evidence in sensory receptor neurons of spontaneously generating action potentials that interfere with and modify the stimulus evoked activity. The present study investigates the generation of spontaneous action potentials in the isolated muscle spindle of the frog by recording receptor potentials, small subthreshold depolarizations, propagated impulses and interspike transients from the first Ranvier-node of the afferent stem fibre. The temporal pattern of spontaneous discharges at resting length depended on several variables interacting at the encoding site. In the absence of mechanical stimulation, a large resting receptor potential steadily depolarized the encoding node and provoked action potentials at irregular intervals. After each action potential, the hyperpolarizing afterpotential provided a slowly increasing depolarizing interspike transient of decreasing slope (time constant 128 ms), which evoked small subthreshold depolarizations (decay time > 5 ms; multimodal amplitude distribution) before the following action potential discharged. The probability of the small subthreshold depolarizations increased the longer the resting receptor potential stayed constant at its maximum amplitude. When increasing the static prestretch level encoding depended also on the stretch-evoked receptor potential as an additional parameter. The resulting depolarizing interspike transients were then larger and also more steeply rising, so that the afferent discharges increased in both rate and regularity. The experiments show dynamic threshold patterns that control action-potential initiation by the assessment of the actual amplitude of depolarization and its rate of rise.     

Dissociations in Hippocampal 5-Hydroxytryptamine Release in the Rat Following Pavlovian Aversive Conditioning to Discrete and Contextual Stimuli

The experiments examined the release of 5-hydroxytryptamine using in vivo microdialysis methods in the hippocampus of freely moving rats following Pavlovian aversive conditioning to discrete and contextual stimuli. Differential conditioning was achieved by manipulating the interval between the offset of a discrete auditory ‘clicker’stimulus and the onset of a mild foot-shock reinforcer (0.5 mA, 0.5 s). Foot-shock occurred either simultaneously with the last second of the discrete auditory stimulus (in short-trace subjects) or 60 s later (long-trace subjects). In this way, subjects were preferentially conditioned to the discrete stimulus and background ‘contextual’stimuli respectively. During conditioning subjects also received two identical unpaired visual stimuli. At test, dialysates were collected and behavioural measures taken as all animals experienced (i) the aversive and two other ‘neutral’environments, and (ii) the discrete unconditioned and conditioned stimuli presented in both aversive and neutral environments. Exposure to the aversive environment, but not to either of the two neutral environments, was associated with significantly increased hippocampal 5-hydroxytryptamine release in long-trace subjects. There was also a small but non-significant increase in 5-hydroxytryptamine release in short-trace animals. In contrast, hippocampal 5-hydroxytryptamine release was unaffected by presentation of either of the discrete stimuli under all conditions. The last result was obtained despite robust behavioural responses (freezing) to the discrete conditioned stimulus. These data do not agree with the hypothesis that aversive cues generally activate 5-hydroxytryptamine function in the hippocampus. Rather, they suggest a degree of specificity whereby 5-hydroxytryptamine release in the hippocampus was determined primarily by other qualitative properties of the conditioned aversive stimulus, namely whether the aversive cue was discrete or contextual, as well as by the magnitude of conditioning.     

The mismatch negativity to frequency deviant stimuli during natural sleep

Eight subjects spent a single night in the sleep laboratory. Event-related potentials (ERPs) were recorded during the presentation of two auditory ‘oddball’ stimulus conditions in which tonal frequency was manipulated. In the first condition, 1000 Hz ‘standard’ and 2000 Hz ‘deviant’ tones were presented. In the second condition, the deviant tone was reduced to 1050 Hz. In both conditions, deviant probability was 0.2. Stimuli were presented every 600 ms during wakefulness and stages 2, 4, and REM of sleep. A distinctive N1 wave was visible in both stimulus conditions when the subject was awake. The deviant stimuli elicited a ‘mismatch negativity’ (MMN) that inverted in polarity at the mastoid. In REM sleep, an N1 and a MMN were also elicited in both conditions. In the large deviance condition, the MMN had a slightly attenuated amplitude and was shorter in duration while in the small deviant condition, its peak latency was unusually early. Neither the N1 nor the MMN could be recorded in non-REM sleep.     

Averaged and single-trial brain responses in the assessment of human sound detection

We investigated sound detection in humans with magnetoencephalography and behavioural measurements. Sounds with intensity increasing smoothly over 125-1000 ms elicited a transient response in auditory cortex with a peak latency in the 200-600 ms range. Importantly, peak latency accurately predicted behavioural reaction time and was unaffected by attentional engagement. Peak amplitude was augmented when the study participants attended to the stimuli and when stimulus duration was decreased. For investigating the cause of these amplitude variations in the averaged response we designed a wavelet-based method for analysing single-trial responses. We found that attention affects the amplitude of the single-trial responses whereas the intensity slope of the stimulus modifies their latency distribution. The transient response reported here holds promise for rapid, objective hearing assessment not requiring a behavioural task.     

Influence of ventral tegmental area (A10 region) on flight behaviour elicited by hypothalamic stimulation in the cat

The influence of the ventral tegmental area (VTA) (dopaminergic A10 group neurons) on flight behaviour, induced by hypothalamic stimulation, was studied in the cat. Co-stimulation of hypothalamus and VTA (ipsi- or contralateral) induced an increase of the flight latency. Slow-motion analysis of flight behaviour showed that this increase was due to the augmentation of the fixation latency (FL) whereas the upright latency (UL) was not modified. Sulpiride injection (50 mg/kg i.p.) provoked the disappearance of VTA effect without affecting the basal behavioural sequence. The results show that DA A10 group neurons increase the attentive component of the flight reaction, suggesting a possible influence of the DA A10 system on the mechanisms modulating focused attention in the animal.     

Eye and head movements during vestibular stimulation in the alert rabbit

Rabbits passively oscillated in the horizontal plane with a free head tended to stabilize their head in space (re: earth-fixed surroundings) by moving the head on the trunk (neck angular deviation, NAD) opposite the passively imposed body rotation. The gain (NAD/body rotation) of head stabilization varied from 0.0 to 0.95 (nearly perfect stability) and was most commonly above 0.5. Horizontal eye movement (HEM) was inversely proportional to head-in-space stability, i.e. the gaze (sum of HEM, NAD, and body rotation) was stable in space (regardless of the gain of head stabilization). When the head was fixed to the rotating platform, attempted head movements (head torque) mimicked eye movements in both the slow and fast phases of vestibular nystagmus; tonic eye position was also accompanied by conjugate shifts in tonic head torque. Thus, while eye and head movements may at times be linked, that the slow eye and head movements vary inversely during vestibular stimulation with a free head indicates that the linkage is not rigid.Absence of a textured stationary visual field consistently produced a response termed ‘visual inattentiveness,’ which was characterized by, among other things, a reduction of head and gaze stability in space. This behavioral response could also be reproduced in a subject allowed vision during prolonged vestibular stimulation in the absence of other environmental stimuli. It is suggested that rabbits optimize gaze stability (re: stationary surroundings), with the head contributing variably, as long as the animal is attending to its surroundings.     

Effects of stimulus repetition rate on slow and fast components of auditory brain-stem responses

Effects of stimulus repetition rate on the slow and fast components of the auditory brain-stem response (ABR) were investigated in 10 adult subjects with normal hearing. The ABRs were recorded with click stimuli at repetition rates of 8, 13.3, 23.8, 40 and 90.9/sec and at an intensity level of 55 dB nHL. Power spectral analysis of the averaged responses was performed. Then the responses were divided into a slow component (0-400 Hz) and a fast component (400-1500 Hz) by using digital filtering technique. The magnitude of the slow component was little affected with increasing stimulus rate from 8/sec to 90.9/sec, while successive waves of the fast component, including wave V, decreased in amplitude as stimulus rate was increased. The latency of the slow component and each wave of the fast component was prolonged with increasing click rates. The shift of latency became longer in the later waves than in the earlier waves.