Directional asymmetry in vertical smooth-pursuit and cancellation of the vertical vestibulo-ocular reflex in juvenile monkeys

Young primates exhibit asymmetric eye movements during vertical smooth-pursuit across a textured background such that upward pursuit has low velocity and requires many catch-up saccades. The asymmetric eye movements cannot be explained by the un-suppressed optokinetic reflex resulting from background visual motion across the retina during pursuit, suggesting that the asymmetry reflects most probably, a low gain in upward eye commands (Kasahara et al. in Exp Brain Res 171:306–321, 2006). In this study, we examined (1) whether there are intrinsic differences in the upward and downward pursuit capabilities and (2) how the difficulty in upward pursuit is correlated with the ability of vertical VOR cancellation. Three juvenile macaques that had initially been trained only for horizontal (but not vertical) pursuit were trained for sinusoidal pursuit in the absence of a textured background. In 2 of the 3 macaques, there was a clear asymmetry between upward and downward pursuit gains and in the time course of initial gain increase. In the third macaque, downward pursuit gain was also low. It did not show consistent asymmetry during the initial 2 weeks of training. However, it also exhibited a significant asymmetry after 4 months of training, similar to the other two monkeys. After 6 months of training, these two monkeys (but not the third) still exhibited asymmetry. As target frequency increased in these two monkeys, mean upward eye velocity saturated at ∼15°/s, whereas horizontal and downward eye velocity increased up to ∼40°/s. During cancellation of the VOR induced by upward whole body rotation, downward eye velocity of the residual VOR increased as the stimulus frequency increased. Gain of the residual VOR during upward rotation was significantly higher than that during horizontal and downward rotation. The time course of residual VOR induced by vertical whole body step-rotation during VOR cancellation was predicted by addition of eye velocity during pursuit and VOR x1. These results support our view that the directional asymmetry reflects the difference in the organization of the cerebellar floccular region for upward and downward directions and the preeminent role of pursuit in VOR cancellation.     

Further evidence for selective difficulty of upward eye pursuit in juvenile monkeys: effects of optokinetic stimulation, static roll tilt, and active head movements

The smooth-pursuit system moves the eyes in space accurately to track slowly moving objects of interest despite visual inputs from the moving background and/or vestibular inputs during head movements. Recently, our laboratory has shown that young primates exhibit asymmetric eye movements during vertical pursuit across a textured background; upward eye velocity gain is reduced. To further understand the nature of this asymmetry, we performed three series of experiments in young monkeys. In Experiment 1, we examined whether this asymmetry was due to an un-compensated downward optokinetic reflex induced by the textured background as it moves across the retina in the opposite direction of the pursuit eye movements. For this, we examined the monkeys’ ability to fixate a stationary spot in space during movement of the textured background and compared it with vertical pursuit across the stationary textured background. We also examined gains of optokinetic eye movements induced by downward motion of the textured background during upward pursuit. In both task conditions, gains of downward eye velocity induced by the textured background were too small to explain reduced upward eye velocity gains. In Experiment 2, we examined whether the frame of reference for low-velocity, upward pursuit was orbital or earth vertical. To test this, we first applied static tilt in the roll plane until the animals were nearly positioned on their side in order to dissociate vertical or horizontal eye movements in the orbit from those in space. Deficits were observed for upward pursuit in the orbit but not in space. In Experiment 3, we tested whether asymmetry was observed during head-free pursuit that requires coordination between eye and head movements. Asymmetry in vertical eye velocity gains was still observed during head-free pursuit although it was not observed in vertical head velocity. These results, taken together, suggest that the asymmetric eye movements during vertical pursuit are specific for upward, primarily eye pursuit in the orbit.     

Oculo-manual coordination control: respective role of visual and non-visual information in ocular tracking of self-moved targets

We evaluated the role of visual and non-visual information in the control of smooth pursuit movements during tracking of a self-moved target. Previous works have shown that self-moved target tracking is characterised by shorter smooth pursuit latency and higher maximal velocity than eye-alone tracking. In fact, when a subject tracks a visual target controlled by his own arm, eye movement and arm movement are closely synchronised. In the present study, we showed that, in a condition where the direction of motion of a self-moved visual target was opposite to that of the arm (same amplitude, same velocity, but opposite direction of movement), the resulting smooth pursuit eye movements occurred with low latency, and continued for about 140 ms in the direction of the arm movement rather than in the direction of the actual visual target movement. After 140 ms, the eye movement direction reversed through a combination of smooth pursuit and saccades. Subsequently, while arm and visual target still moved in opposite directions, smooth pursuit occurred in pace with the visual target motion. Subjects were also submitted to a series of 60 tracking trials, for which the arm-to-target motion relationship was systematically reversed. Under these conditions subjects were able to initiate early smooth pursuit in the actual direction of the visual target. Overall, these results confirm that non-visual information produced by the arm motor system can trigger and control smooth pursuit. They also demonstrate the plasticity of the neuronal network handling eye-arm coordination control.     

Directional organization of eye movement and visual signals in the floccular lobe of the monkey cerebellum

The floccular lobe of the monkey is critical for the generation of visually-guided smooth eye movements. The present experiments reveal physiological correlates of the directional organization in the primate floccular lobe by examining the selectivity for direction of eye motion and visual stimulation in the firing of individual Purkinje cells (PCs) and mossy fibers. During tracking of sinusoidal target motion along different axes in the frontoparallel plane, PCs fell into two classes based on the axis that caused the largest modulation of simple-spike firing rate. For horizontal PCs, the response was maximal during horizontal eye movements, with increases in firing rate during pursuit toward the side of recording (ipsiversive). For vertical PCs, the response was maximal during eye movement along an axis just off pure vertical, with increases in firing rate during pursuit directed downward and slightly contraversive. During pursuit of target motion at constant velocity, PCs again fell into horizontal and vertical classes that matched the results from sinusoidal tracking. In addition, the directional tuning of the sustained eye velocity and transient visual components of the neural responses obtained during constant velocity tracking were very similar. PCs displayed very broad tuning approximating a cosine tuning curve; the mean half-maximum bandwidth of their tuning curves was 170–180 °. Other cerebellar elements, related purely to eye movement and presumed to be mossy fibers, exhibited tuning approximately 40 ° narrower than PCs and had best directions that clustered around the four cardinal directions. Our data indicate that the motion signals encoded by PCs in the monkey floccular lobe are segregated into channels that are consistent with a coordinate system defined by the vestibular apparatus and eye muscles. The differences between the tuning properties exhibited by PCs compared with mossy fibers indicate that a spatial transformation occurs within the floccular lobe.     

Temporospatial properties of the effects of bottom-up attention on smooth pursuit initiation in humans

We examined the temporal and spatial properties of the effects of target saliency on the initiation of smooth pursuit eye movement in humans. Visual stimuli consisted of random dots projected on a large-field screen. During a fixation period, a cluster of dots (2×2 deg) was blinked (turned off for a short period) to make that region stand out from the remaining background and serve as a cue. After a delay (cue lead time), a cluster of dots (2×2 deg) started to move as a pursuit target. The target and cue were presented at either identical or different locations so that the subject could not predict the target location from the cue location. We examined the time course of the effect of the cue on pursuit initiation by changing the cue lead time. In half of the trials, the cue and target locations were identical, and in the other trials, they were not identical. There was a clear effect of the cue only for the trials where the cue and target locations were identical. The effect of the cue increased as cue lead time increased, peaked at ~160 ms, and then decreased. We also examined the spatial extent of the effect of the cue by varying the distance between the target and the cue. The largest effect was observed when the cue and the target were presented at the same location. Facilitating effects were observed when the target and the cue were presented in the same hemifield but not when presented in the opposite hemifield.     

Prediction in the oculomotor system: smooth pursuit during transient disappearance of a visual target

Summary Eye movements were recorded in human subjects who tracked a target spot which moved horizontally at constant speeds. At random times during its trajectory, the target disappeared for variable periods of time and the subjects attempted to continue tracking the invisible target. The smooth pursuit component of their eye movements was isolated and averaged. About 190 ms after the target disappeared, the smooth pursuit velocity began to decelerate rapidly. The time course of this deceleration was similar to that in response to a visible target whose velocity decreased suddenly. After a deceleration lasting about 280 ms, the velocity stabilized at a new, reduced level which we call the residual velocity. The residual velocity remained more or less constant or declined only slowly even when the target remained invisible for 4 s. When the same target velocity was used in all trials of an experiment, the subjects' residual velocity amounted to 60% of their normal pursuit velocity. When the velocity was varied randomly from trial to trial, the residual velocity was smaller; for target velocities of 5, 10, and 20 deg/s it reached 55, 47, and 39% respectively. The subjects needed to see targets of unforeseeable velocity for no more than 300 ms in order to develop a residual velocity that was characteristic of the given target velocity. When a target of unknown velocity disappeared at the very moment the subject expected it to start, a smooth movement developed nonetheless and reached within 300 ms a peak velocity of 5 deg/s which was independent of the actual target velocity and reflected a default value for the pursuit system. Thereafter the eyes decelerated briefly and then continued with a constant or slightly decreasing velocity of 2–4 deg/s until the target reappeared. Even when the subjects saw no moving target during an experiment, they could produce a smooth movement in the dark and could grade its velocity as a function of that of an imagined target. We suggest that the residual velocity reflects a first order prediction of target movement which is attenuated by a variable gain element. When subjects are pursuing a visible target, the gain of this element is close to unity. When the target disappears but continued tracking is attempted, the gain is reduced to a value between 0.4 and 0.6.Supported by grants DFG Be 783/1 and Be 783/2-1 (1), and NIH RR 00166 and EY 00745 (2)     

Combined smooth and saccadic ocular pursuit during the transient occlusion of a moving visual object

Accurate ocular pursuit during a transient occlusion interval would minimize retinal position and velocity error, and could provide an advantage when discriminating object characteristics at reappearance. This study was designed to examine how the smooth and saccadic response extrapolates the trajectory of a moving visual object during a transient occlusion. We confirmed that subjects could not maintain unity gain smooth pursuit during the transient occlusion. Eye velocity decayed significantly without visual feedback but then in the majority of subjects, there was a recovery that brought eye velocity back towards object velocity. However, eye velocity did not increase to a level that eliminated the developing position error. Subjects corrected for the resulting error in eye position by releasing saccades that generally placed the eye ahead of the occluded object’s extrapolated position. The majority of saccadic correction occurred between 220 and 600 ms of the occlusion interval, and when combined with the smooth response enabled accurate pursuit of a 10°/s object for up to 1,200 ms of occlusion. The lack of saccadic correction after 600 ms of occlusion combined with the reduced eye velocity resulted in significant undershoot of eye position at the moment of object reappearance when pursuing an 18°/s object. We suggest that extra-retinal information regarding eye velocity and smooth eye displacement could be available from a continually updating efference copy of eye motion in MST, whereas a veridical representation of extrapolated object velocity and displacement could be obtained from persistent activity in FEF.     

Oculo-manual coordination control: Ocular and manual tracking of visual targets with delayed visual feedback of the hand motion

Summary The aim of this study was to examine coordination control in eye and hand tracking of visual targets. We studied eye tracking of a self-moved target, and simultaneous eye and hand tracking of an external visual target moving horizontally on a screen. Predictive features of eye-hand coordination control were studied by introducing a delay (0 to 450 ms) between the Subject's (S's) hand motion and the motion of the hand-driven target on the screen. In self-moved target tracking with artificial delay, the eyes started to move in response to arm movement while the visual target was still motionless, that is before any retinal slip had been produced. The signal likely to trigger smooth pursuit in that condition must be derived from non-visual information. Candidates are efference copy and afferent signals from arm motion. When tracking an external target with the eyes and the hand, in a condition where a delay was introduced in the visual feedback loop of the hand, the Ss anticipated with the arm the movement of the target in order to compensate the delay. After a short tracking period, Ss were able to track with a low lag, or eventually to create a lead between the hand and the target. This was observed if the delay was less than 250–300 ms. For larger delays, the hand lagged the target by 250–300 ms. Ss did not completely compensate the delay and did not, on the average, correct for sudden changes in movement of the target (at the direction reversal of the trajectory). Conversely, in the whole range of studied delays (0–450 ms), the eyes were always in phase with the visual target (except during the first part of the first cycle of the movement, as seen previously). These findings are discussed in relation to a scheme in which both predictive (dynamic nature of the motion) and coordination (eye and hand movement system interactive signals) controls are included.     

Oculo-manual tracking of visual targets in monkey: role of the arm afferent information in the control of arm and eye movements

Summary The study was aimed at defining the role of hand (and arm) kinaesthetic information in coordination control of the visuo-oculo-manual tracking system. Baboons were trained to follow slow-moving and stepping visual targets either with the eyes alone or with the eyes and a lever moved by the forelimb about the vertical axis. A LED was attached to the lever extremity. Four oculo-manual tracking condidtions were tested and compared to eye-alone tracking: Eye and hand tracking of a visual target presented on a screen, eye tracking of the hand, and eye tracking of an imaginary target actively moved by the arm. The performance of the animals evaluated in terms of latency, and velocity and position precision for both eye and hand movements was seen to be equivalent to that of humans in similar situations. After dorsal root rhizotomy (C1-T2) the animals were unable to produce slow arm motion in response to slow-moving targets. Instead, they produced successions of ballistic-like motions whose amplitude decreased as retraining proceeded. In addition, the animals could not longer respond with smooth pursuit eye movements to an imaginary target actively displaced by the animal's forelimb. It was concluded that the absence of ocular smooth pursuit after lesion results from the disruption of a signal derived from arm kinaesthetic information and addresses to the oculomotor system. This signal is likely to be used in the control of coordination between arm and eye movements during visuo-oculo-manual tracking tasks. One cause of the animal's inability to achieve slow arm movement in response to slow target motion is thought to be due to a lesion-induced alteration of the spinal common pathway dynamics which normally integrate the velocity signal descending from the arm movement command system.     

Directional tuning of cells in area 18 of the feline visual cortex for visual noise,bar and spot stimuli: a comparison with area 17

Summary Directional tuning for visual noise, bar and single spot stimuli was compared over a wide range of velocities in cells from areas 17 and 18 of the visual cortex in lightly-anaesthetized cats. In each area, S-cells were predominantly insensitive to motion of a field of visual noise. C-cells were more sensitive to noise motion than B-cells, but showed heterogeneity in noise sensitivity, which was associated with other response properties: strongly noise-sensitive C-cells had relatively high spontaneous activity and broad directional tuning, and were predominantly direction-selective and binocularly-driven. Frequently, directional tuning for noise was unimodal at low velocity, but became progressively more bimodal as velocity was increased: a trough of depressed response corresponding to the peak in tuning for the bar separated two progressively more widely disparate preferred directions. In area 18, cells with velocity tuned (VT) functions for bar motion developed bimodal tuning for noise well below the optimum velocity for bar or for noise motion, while velocity high-pass (VHP) cells became progressively more bimodally tuned for noise over a wide range of velocities, in parallel with a steep increase in response to bar and noise motion. A high proportion of VT and VHP cells was bimodally tuned for noise at all velocities, one VHP cell showing two discrete lobes of tuning for noise below the threshold velocity for bar motion. Among cells which remained unimodally tuned for noise, VT and VHP cells in area 18 had radically dissimilar preferred directions for noise and bar motion at all velocities. With the exception of VHP cells, velocity bandpass was higher for noise than for bar motion. These results, together with other novel observations on the modality of tuning for noise in preferred and opposite directions of motion, demonstrate that bimodality of tuning for noise cannot simply be an effect of upper cut-off velocity for bar motion (Movshon et al. 1980; Orban 1984). It is argued that the trough between the lobes of tuning arises through laterally-directed inhibitory convergence from superficial- and deep-layer, large basket cells. In 40% of noise-sensitive cells, tuning for bar motion was broader on the flank closest to the preferred direction for noise and for a moving spot, while some 25% of cells showed variations in tuning for bar motion with velocity, which were associated with velocity-dependent changes in tuning for noise. Thus, the broad, asymmetrical tuning of these cell types for bar motion presumably reflects to some extent stimulation of the directional mechanism by the moving bar. Quantitative comparisons showed that C-cells in each area had similar tuning for bar motion, which was substantially broader than that of S- or B-cells. S-cells had the narrowest tuning, though those in area 18 were more broadly tuned than those in area 17.     

An fMRI study on smooth pursuit and fixation suppression of the optokinetic reflex using similar visual stimulation

This study compares brain activation patterns evoked by smooth pursuit and by fixation suppression of the optokinetic reflex (OKR) using similar retinal stimulation. Functional magnetic resonance imaging (fMRI) was performed during smooth pursuit stimulation in which a moving target was presented on a stationary pattern of stripes, and during fixation suppression of OKR in which a stationary target was presented on a moving pattern of stripes. All subjects could effectively ignore the background pattern and were able to keep the target continuously on the fovea with few saccades, in both experiments. Smooth pursuit evoked activation in the frontal eye fields (FEF), the supplementary eye fields (SEF), the parietal eye fields (PEF), the motion-sensitive area (MT/V5), and in lobules and vermis VI of the cerebellum (oculomotor areas). Fixation suppression of OKR induced activation in the FEF, PEF, and MT/V5. The direct comparison analysis revealed more activation in the right lobule VI of the cerebellum and in the right lingual and calcarine gyri during smooth pursuit than during fixation suppression of OKR. Using similar retinal stimulation, our results show that smooth pursuit and fixation suppression of the OKR appear to activate largely overlapping pathways. The increased activity in the oculomotor areas of the cerebellum during smooth pursuit is probably due to the presence of an active eye movement component.     

Modulatory influences of a moving visual noise background on bar-evoked responses of cells in area 18 of the feline visual cortex

Summary The modulatory influence of a synchronously moving visual noise background on responsiveness to an optimally-oriented moving bar stimulus was investigated in visual cortical area 18 of the lightly-anaesthetized cat. The bar and noise background were swept along the axis orthogonal to bar orientation, with the same phase, velocity and amplitude of motion. Cells which were insensitive to motion of visual noise per se or weakly responsive to individual grains in the noise sample showed suppression of bar-evoked responses by simultaneous motion of the noise background. Percent suppression declined with increase in bar length, over a range which could exceed the maximum estimate of receptive field length. The decline in percent suppression was non-linear, becoming progressively flatter in slope as bar length was increased until an asymptotic value was reached; observations on end-stopped cells and on end-free cells with restricted length summation verified that percent suppression was related specifically to the length of the comparison bar and not to the strength of response it evoked. Percent suppression and the extent over which it declined with increase in bar length were comparable for preferred and opposite directions of bar motion even in cells with radically different length-response functions in the two directions, including end-stopped cells with direction-selective end-zones. In contrast to end-inhibition, which was maximal at or near the preferred velocity for a bar of optimal length, percent suppression by motion of the noise background was essentially velocity-invariant; in velocity tuned and velocity high-pass cells, background motion reduced the slope(s) of the velocity-response function, implying that the suppressive action of moving noise backgrounds is divisive rather than subtractive. It is argued that the suppression derives predominantly from an axo-somatic noise-sensitive inhibitory input from superficial- and deep-layer, large basket cells in orientation columns at some distance from those of their target cells.     

Laminar,columnar and topographic aspects of ocular dominance in the primary visual cortex ofCebus monkeys

Summary The representation of the two eyes in striate cortex (V1) ofCebus monkeys was studied by electrophysiological single-unit recordings in normal animals and by morphometric analysis of the pattern of ocular dominance (OD) stripes, as revealed by cytochrome oxidase histochemistry in V1 flat-mounts of enucleated animals. Single-unit recordings revealed that the large majority of V1 neurons respond to the stimulation of either eye but are more strongly activated by one of them. As in other species of monkey, neurons with preference for the stimulation of the same eye are grouped in columns 300–400 µm wide, spanning all cortical layers. Monocular neurons are clustered in layer IVc, specially in its deeper half (IVc-beta), and constitute less than 10% of the population of other layers. Neurons with equal responses to each eye are more commonly found in layer V than elsewhere in V1. In the supragranular layers and in granular layer IVc-alpha neurons strongly dominated by one of the eyes tend to be broadly tuned for orientation, while binocularly balanced neurons tend to be sharply tuned for this parameter. No such correlation was detected in the infragranular layers, and most neurons in layer IVc-beta responded regardless of stimulus orientation. Ocular dominance stripes are present throughout most of V1 as long, parallel or bifurcating bands alternately dominated by the ipsi- or the contralateral eye. They are absent from the cortical representations of the blind spot and the monocular crescent. The domains of each eye occupy nearly equal portions of the surface of binocular V1, except for the representation of the periphery, where the contralateral eye has a larger domain, and a narrow strip along the border of V1 with V2, where either eye may predominate. The orderliness of the pattern of stripes and the relationship between stripe arrangement and the representation of the visual meridians vary with eccentricity and polar angle but follow the same rules in different animals. These results demonstrate that the laminar, columnar and topographic distribution of neurons with different degrees of OD in V1 is qualitatively similar in New- and Old World monkeys of similar sizes and suggest that common ancestry, rather than parallel evolution, may account for the OD phenotypes of contemporaneous simians.     

Independent roles for the dorsal paraflocculus and vermal lobule VII of the cerebellum in visuomotor coordination

Two distinct areas of cerebellar cortex, vermal lobule VII and the dorsal paraflocculus (DPFl) receive visual input. To help understand the visuomotor functions of these two regions, we compared their afferent and efferent connections using the tracers wheatgerm agglutinin horseradish peroxidase (WGA-HRP) and biotinilated dextran amine (BDA). The sources of both mossy fibre and climbing fibre input to the two areas are different. The main mossy fibre input to lobule VII is from the nucleus reticularis tegmenti pontis (NRTP), which relays visual information from the superior colliculus, while the main mossy fibre input to the DPFl is from the pontine nuclei, relaying information from cortical visual areas. The DPFl and lobule VII both also receive mossy fibre input from several common brainstem regions, but from different subsets of cells. These include visual input from the dorsolateral pons, and vestibular–oculomotor input from the medial vestibular nucleus (MVe) and the nucleus prepositus hypoglossi (Nph). The climbing fibre input to the two cerebellar regions is from different subdivisions of the inferior olivary nuclei. Climbing fibres from the caudal medial accessory olive (cMAO) project to lobule VII, while the rostral MAO (rMAO) and the principal olive (PO) project to the DPFl. The efferent projections from lobule VII and the DPF1 are to all of the recognised oculomotor and visual areas within the deep cerebellar nuclei, but to separate territories. Both regions play a role in eye movement control. The DPFl may also have a role in visually guided reaching.     

Immunocytochemical localization of calcineurin in the adult and developing primary visual cortex of cats

An immunocytochemical method was used to localize calcineurin, a calcium-dependent calmodulinstimulated protein phosphatase, in the primary visual cortex of developing and adult cats. In the adult calcineurin immunoreactivity exhibits a laminar distribution with dense labeling in the upper half of layers II/III and two lightly labeled bands in lower layer IV and in layer VI. Most of the immunoreactive neurons are pyramidal in shape and appear to form a subpopulation of cortical neurons, but non-pyramidal neurons were also labeled, especially during early stages of postnatal development. The distribution pattern of calcineurin immunoreactivity showed developmental changes until at least 3 months of age. The number of calcineurin-positive cells abruptly increased at 3 weeks, and heavily labeled neurons appeared in a well-delineated band in layer IV between 3 and 5 weeks of age. At 6 to 10 weeks, neurons in layers II/III also became strongly immunoreactive. At this developmental stage intensely stained cells were thus distributed throughout layers II to IV. Thereafter, there was a marked decrease in the number of immunoreactive cells in layer IV and beyond 12 weeks the distribution pattern of calcineurin immunoreactivity became similar to that of adult animals. These changes of calcineurin expression show some relation with the inside-out pattern of cortical maturation and with the time course and the laminar selectivity of use-dependent malleability. Therefore, we suggest that calcineurin may be involved in processes of neuronal differentiation and experience-dependent plasticity.     

Stimulus specificity of binocular cells in the cat's visual cortex: ocular dominance and the matching of left and right eyes

Summary Most cells in the striate cortex respond to visual stimulation through either eye. We have examined quantitatively the matching of response specificity for the two eyes. Our intention was to determine the degree to which this matching depends on ocular dominance. We used standard single cell recording techniques and studied responses to sinusoidal gratings of different spatial frequencies, orientations, and contrasts. For all tests, stimuli were randomly interleaved both with respect to the value of each parameter, and the eye which was stimulated. After estimating ocular dominance qualitatively and quantitatively, we measured: response modulation (to help identify whether a cell was simple or complex), orientation and spatial frequency tuning, and contrast response functions (to estimate contrast thresholds). Results show that: (1) Response modulation is well matched between the two eyes, but there is a slight tendency for the dominant eye to respond with less modulation. (2) Optimal orientation and spatial frequency and their respective tuning widths were similar for the two eyes. In general, tuning functions for the two eyes differed mainly in slope. However, in each case, there was a tendency for the dominant eye to have broader tuning widths. (3) In most cases, contrast response functions for the two eyes differed mainly in their slopes. Extrapolation to spontaneous levels suggests that estimated contrast thresholds are relatively independent of ocular dominance although, again, there was a tendency for the dominant eye to exhibit slightly lower estimated thresholds. These findings demonstrate that response characteristics between the two eyes are generally well matched regardless of relative response strength. There are, however, small but clear differences between the two eyes for all parameters we measured which are related to and demonstrate that ocular dominance influences the degree of matching between the two eyes.     

Purkinje cell activity in the flocculus of vestibular neurectomized and normal monkeys during optokinetic nystagmus (OKN) and smooth pursuit eye movements

Summary 1. Single unit activity was recorded in the primate flocculus after the vestibular nerves were cut (bilateral vestibular neurectomy) during optokinetic nystagmus (OKN), smooth pursuit eye movements (SP) and whole field visual stimulation with gaze fixed on a stationary target light (OKN-suppression). Following vestibular neurectomy monkeys had no vestibular responses and no optokinetic after-nystagmus (OKAN) in the horizontal plane. However, OKN slow phases still reached steady state velocities of up to 100 deg/s. 2. After neurectomy, simple spike (SS) activity of Purkinje cells (P-cells) was modulated in relation to eye velocity, regardless of whether eye velocity was induced by a small target light moving in darkness (SP) or by a moving visual surround (OKN). In over 90% of the P-cells firing rate increased with eye velocity to the ipsilateral side and decreased with velocities to the contralateral side. Modulation in firing rate increased monotonically with increasing eye velocity. The strength of modulation was similar during SP and OKN for the same eye velocity. 3. The change in firing rate of P-cells in response to a sudden change in optokinetic stimulus velocity contained a component related to eye velocity and a component related to eye acceleration. Only a few P-cells were also modulated with image slip velocity during OKN-suppression. 4. The modulation of P-cells during SP and OKN was compared in normal and vestibular neurectomized monkeys. The sensitivity of floccular P-cells to eye velocity during SP was 1.14 imp·s^–1/deg·s^–1 in normal monkey and 1.28 imp·s^–1/deg·s^–1 after neurectomy. The similarity of eye velocity sensitivities demonstrates that neurectomy does not change the characteristics of floccular P-cell modulation during SP. In contrast, during OKN modulation of P-cells is quite different in normal and neurectomized monkey. In normal monkey, P-cells are modulated during steady state OKN for eye velocities above 40–60 deg/s only. This threshold velocity corresponds approximately to the maximal initial OKAN velocity (i.e. OKAN saturation velocity). After neurectomy, the threshold velocity is 0 deg/s and P-cells are modulated during steady state OKN also over ranges of eye velocities that do not cause a response in normal monkey. Sensitivities of P-cells to eye velocity during OKN for eye velocities above the threshold velocity are 1.0 imp·s^–1/deg·s^–1 in neurectomized monkey and 1.43 imp·s^–1/deg·s^–1 in normal monkey. 5. The hypothesis has been put forward that OKN slow phase velocity in normal monkey has two dynamically different components, a fast and a slow component. The results strongly suggest that the two components depend on different neuronal populations. Firing rate of floccular P-cells is modulated in relation to the fast component only. The results furthermore support the idea that it is the smooth pursuit system which may generate the fast component in the OKN slow phase velocity response.Supported by Swiss National Foundation for Scientific Research (Nr. 3.718-0.80 and 3.593-0.84)     

Prediction and a stationary, structured visual background influence the dynamics of the smooth-pursuit offset in humans

 It is still not clear whether the transition from pursuit eye movements to fixation is mediated by the same system that initiates pursuit, or whether another system, a specialized fixation system, is responsible. To investigate this question we measured smooth-pursuit eye movements and smooth-pursuit termination in five normal subjects using both predictable and unpredictable step-ramp stimuli (velocities 10° and 20°/s) in front of a homogeneous and a structured visual background in order to compare the profile of eye velocity under these different conditions. With the predictable and/or structured visual background there was a gradual transition of eye velocity toward zero. In contrast, with the unpredictable stimulus in front of a homogeneous background, eye velocity during the offset was characterized by an overshoot (on the average, 2.2±1.0°/s for 10°/s ramps) before eye velocity settled at zero. Under this condition, steady-state velocity gain and the deceleration of the offset were significantly higher than during the other paradigm with the same target velocity. The latency of the pursuit offset was significantly shorter when a predictable stimulus was used. The duration of the offset did not depend on the experimental condition used. These findings imply that the pursuit onset and offset have some similarities and may be mediated by the same oculomotor system. Received: 6 February 1998 / Accepted: 8 July 1998     

Receptive fields of units in the visual cortex of the cat in the presence and absence of bodily tilt

Summary The receptive fields of units in the visual cortex of anaesthetised cats were studied using spots or slits of light. Some fields were found to be stable when they were repeatedly plotted with the cat maintained in the horizontal position: other fields were not stable and the sharpness of spatial tuning varied though the orientation of the axis did not shift. When the cat was tilted the field axis of the majority of cells followed the tilt. In 14 cells, however, changes occurred in the receptive field which were not observed when the animal remained in the horizontal plane. These changes included drifts of the field axis in a direction which, with one exception, was opposite to the tilt, and alterations in the spatial extent of the field. On returning the animal to horizontal the axis of 4 fields drifted past the original orientation. These effects were not eliminated by either bilateral destruction of the labyrinth or high cervical transection of the spinal cord. The time of onset of the tilt effects varied from cell to cell: some of this variability is probably an effect of anaesthesia.The findings are consistent with the view that the receptive field of certain cells in the visual cortex are capable of being modified, one of the modifying influences being the orientation of the body in space.This work was supported by grants from the Science Research Council to G. Horn and from the U.S. Public Health Service to G. Stechler (Grant MH 16215) and R.M. Hill (Grant NB 05653).     

Clear-cell adenocarcinoma of the female genital tract: presence of hyaline stroma and tigroid background in various types of cytologic specimens

Hyaline basement membrane-like stromal material and tigroid background are distinctive cytologic features observed in Diff-Quik (DQ)- or Giemsa-stained smears of clear-cell adenocarcinoma (CCA) of the female genital tract. However, it is uncertain how often these features are present in different types of cytologic specimens, and which type of preparation is optimal for this diagnosis. We therefore reviewed the cytologic features of CCA in three types of specimens, including 15 scrape cytology specimens, 7 fine-needle aspiration (FNA) specimens, and 15 peritoneal cytology specimens, with emphasis on the features observed in DQ-stained smears. The cell morphology in scrape cytology specimens and FNA specimens was comparable, whereas in peritoneal cytology specimens, the cytoplasm was better preserved. Most tumor cells had fragile cytoplasm containing variable amounts of fine vacuoles, and round nuclei with distinct or prominent nucleoli. Hyaline stroma was present in 93% of scrape cytology specimens, 71% of FNA specimens, and 80% of peritoneal cytology specimens. Tigroid background was observed in 47% of scrape cytology specimens, 43% of FNA specimens, but in none of the peritoneal cytology specimens. Formation of a tigroid background may be prevented by the abundant fluid content in peritoneal cytology specimens. Hyaline stroma and tigroid background were uncommonly seen in scrape smears from other types of primary ovarian tumors, mainly juvenile granulosa cell tumor and yolk sac tumor. However, the additional presence of papillary structures allows CCA to be readily distinguished from these other tumors. We propose that scrape cytology offers the best approach for the intraoperative cytologic diagnosis of CCA.