Adonis K. Moschovakis

Neural network simulations of the primate oculomotor system

The performance of a neural network that simulates the vertical saccade-generating portion of the primate brain is evaluated. Consistent with presently available anatomical evidence, the model makes use of an eye displacement signal for its feedback. Its major features include a simple mechanism for resetting its integrator at the end of each saccade, the ability to generate staircases of saccades in response to stimulation of the superior colliculus, and the ability to account for the monotonic relation between motor error and the instantaneous discharge of presaccadic neurons of the superior colliculus without placing the latter within the local feedback loop. Several experimentally testable predictions about the effects of stimulation or lesion of saccaderelated areas of the primate brain are made on the basis of model output in response to “stimulation” or “lesion” of model elements.

The superior colliculus and eye movement control

Saccade metrics are largely determined by command signals generated in the superior colliculus. Recent work has improved our knowledge of the highly complex axonal trees that disseminate these signals. It has also demonstrated that electrical stimulation of the superior colliculus evokes slow (drifts) as well as fast (saccades) eye movements and has provided detailed quantitative descriptions of their metrical properties. New biologically influenced models of the superior colliculus provide a detailed and realistic account of the discharge of tectal pre-saccadic neurons, the properties of evoked saccades and the ability of subjects to execute correct saccades in a variety of circumstances.

The neural integrators of the mammalian saccadic system.

The neural velocity to position integrators transform the saccade related signal of the burst generators into an eye position related tonic signal they convey to motoneurons. They are largely confined to three heavily interconnected midbrain structures: 1) The interstitial nucleus of Cajal (NIC), 2) The nucleus prepositus hypoglossi (NPH), 3) The vestibular nuclei (VN). Integration in the horizontal and vertical planes is accomplished largely independently by the NPH-VN and the NIC-VN complexes, respectively. Cells in these regions carry a more or less intense phasic signal related to saccades and a tonic signal related to eye position. Depending on the relationship between the rate of their discharge and the position of the eyes, these cells have been further subdivided into regular or irregular, more or less sensitive, and bi-directionally or uni-directionally modulated. The present review provides a brief description of their discharge pattern and that of burst neurons and extraocular motoneurons. Then, evidence concerning the input-output connections of relevant cell classes is summarized. Finally, several modelling attempts to simulate the neural velocity-to-position integrators are presented and their verisimilitude is evaluated in the light of psychophysical, anatomical, physiological and neurological evidence.

Optimal Control of Gaze Shifts

To explore the visible world, human beings and other primates often rely on gaze shifts. These are coordinated movements of the eyes and head characterized by stereotypical metrics and kinematics. It is possible to determine the rules that the effectors must obey to execute them rapidly and accurately and the neural commands needed to implement these rules with the help of optimal control theory. In this study, we demonstrate that head-fixed saccades and head-free gaze shifts obey a simple physical principle, “the minimum effort rule.” By direct comparison with existing models of the neural control of gaze shifts, we conclude that the neural circuitry that implements the minimum effort rule is one that uses inhibitory cross talk between independent eye and head controllers.

The place code of saccade metrics in the lateral bank of the intraparietal sulcus.

The lateral intraparietal area (LIP) of monkeys is known to participate in the guidance of rapid eye movements (saccades), but the means it uses to specify movement variables are poorly understood. To determine whether area LIP devotes neural space to encode saccade metrics spatially, we used the quantitative [14C]deoxyglucose method to obtain images of the distribution of metabolic activity in the intraparietal sulcus (IPs) of rhesus monkeys trained to repeatedly execute saccades of the same amplitude and direction for the duration of the experiment. Different monkeys were trained to perform saccades of different sizes and in different directions. A clear topography of saccade metrics was found in the cytoarchitectonically identified area LIP ventral (LIPv) contralateral to the direction of the eye movements. We demonstrate that the representation of the vertical meridian runs parallel to the fundus of the IPs and that it is not orthogonal to the representation of the horizontal meridian. Instead, the latter runs through the middle of LIPv parallel to its border with area LIP dorsal (LIPd). The upper part of oculomotor space is represented rostrally and dorsally relative to the horizontal meridian toward the LIPv–LIPd border, whereas the lower part of oculomotor space is represented caudally and ventrally toward the caudal edge of the IPs. Saccade amplitude is also represented in an orderly manner.

Saccade-Related Information in the Superior Temporal Motion Complex: Quantitative Functional Mapping in the Monkey

Although the role of the motion complex [cortical areas middle temporal (V5/MT), medial superior temporal (MST), and fundus of the superior temporal (FST)] in visual motion and smooth-pursuit eye movements is well understood, little is known about its involvement in rapid eye movements (saccades). To address this issue, we used the quantitative 14C-deoxyglucose method to obtain functional maps of the cerebral cortex lying in the superior temporal sulcus of rhesus monkeys executing saccades to visual targets and saccades to memorized targets in complete darkness. Fixational effects were observed in MT-foveal, FST, the anterior part of V4-transitional (V4t), and temporal-occipital areas. Saccades to memorized targets activated areas V5/MT, MST, and V4t, which were also activated for saccades to visual targets. Regions activated in the light and in the dark overlapped extensively. In addition, saccades to visual targets activated areas FST and the intermediate part of the polysensory temporal-parietal-occipital area. Cortical activity related to visually guided saccades could be explained, at least in part, by visual motion. Because only oculomotor signals can account for the equally robust activations induced by memory saccades in complete darkness, we suggest that areas V5/MT, MST, and V4t receive and/or process saccade-related oculomotor information.

Topography of Visuomotor Parameters in the Frontal and Premotor Eye Fields

To determine whether the periarcuate frontal cortex spatially encodes visual and oculomotor parameters, we trained monkeys to repeatedly execute saccades of the same amplitude and direction toward visual targets and we obtained quantitative images of the distribution of metabolic activity in 2D flattened reconstructions of the arcuate sulcus (As) and prearcuate convexity. We found two topographic maps of contraversive saccades to visual targets, separated by a region representing the vertical meridian: the first region straddled the fundus of the As and occupied areas 44 and 6-ventral, whereas the second one occupied areas 8A and 45 in the anterior bank of the As and the prearcuate convexity. The representation of the vertical meridian runs along the posterior borders of areas 8A and 45 (deep in the As). In both maps, the upper part of visuo-oculomotor space is represented ventrally and laterally and the lower part dorsally and medially whereas dorsal and ventral regions are separated by the representation of the horizontal meridian.

Functional imaging of the intraparietal cortex during saccades to visual and memorized targets.

The representation of perceived space and intended actions in the primate parietal cortex has been the subject of considerable debate. To address this issue, we used the quantitative 14C-deoxyglucose method to obtain maps of the activity pattern in the intraparietal cortex of rhesus monkeys executing saccades to visual and memorized targets. The principal effect induced by memory-guided saccades was found more caudally in the deepest part of the middle third of the lateral bank (within area LIPv) whereas that induced by visually guided saccades extended more rostrally and superficially in the anterior third of the bank (within area LIPd). The memory-saccade-related and the visual-saccade-related regions of activation overlapped only within area LIPv. Besides saccade execution, maximal activity in area LIPd required a visual stimulus. The region activated by visual fixation was located at the border of LIPv and LIPd, extending mainly within area LIPd, and occupying about one third of the neural space of the region activated for visual-saccades. We suggest that the lateral intraparietal cortex represents visual and motor space in segregated, albeit partially overlapping, regions.

The pharmacological effect of fractions obtained by smoking cannabis through a water-pipe. II. A second fractionation step.

The catatonic activity, prolongation of phenobarbital sleeping-time, convulsant action and disruption of nestbuilding activity were assessed in mice subjected to 4 cannabis pyrolysis products and their tobacco analogues. All but one of the cannabis fractions prolonged the pentobarbital sleeping-time and disrupted the nest-building activity of mice in a way not related to their content in the main cannabinoids. Nest-building activity seems to be the most valid assay we have used so far.

Hashish smoke interfers with Sidman avoidance in mice.

Hashish smoke has been proved to be active in the Sidman avoidance. Its activity is similar to that of hallucinogens.