Modelling 3D saccade generation by feedforward optimal control

An interesting problem for the human saccadic eye-movement system is how to deal with the degrees-of-freedom problem: the six extra-ocular muscles provide three rotational degrees of freedom, while only two are needed to point gaze at any direction. Measurements show that 3D eye orientations during head-fixed saccades in far-viewing conditions lie in Listing’s plane (LP), in which the eye’s cyclotorsion is zero (Listing’s law, LL). Moreover, while saccades are executed as single-axis rotations around a stable eye-angular velocity axis, they follow straight trajectories in LP. Another distinctive saccade property is their nonlinear main-sequence dynamics: the affine relationship between saccade size and movement duration, and the saturation of peak velocity with amplitude. To explain all these properties, we developed a computational model, based on a simplified and upscaled robotic prototype of an eye with 3 degrees of freedom, driven by three independent motor commands, coupled to three antagonistic elastic muscle pairs. As the robotic prototype was not intended to faithfully mimic the detailed biomechanics of the human eye, we did not impose specific prior mechanical constraints on the ocular plant that could, by themselves, generate Listing’s law and the main-sequence. Instead, our goal was to study how these properties can emerge from the application of optimal control principles to simplified eye models. We performed a numerical linearization of the nonlinear system dynamics around the origin using system identification techniques, and developed open-loop controllers for 3D saccade generation. Applying optimal control to the simulated model, could reproduce both Listing’s law and and the main-sequence. We verified the contribution of different terms in the cost optimization functional to realistic 3D saccade behavior, and identified four essential terms: total energy expenditure by the motors, movement duration, gaze accuracy, and the total static force exerted by the muscles during fixation. Our findings suggest that Listing’s law, as well as the saccade dynamics and their trajectories, may all emerge from the same common mechanism that aims to optimize speed-accuracy trade-off for saccades, while minimizing the total muscle force during eccentric fixation.

Microstimulation in the primary visual cortex: activity patterns and their relation to visual responses and evoked saccades

Intra cortical microstimulation (ICMS) in the primary visual cortex (V1) can generate the visual perception of phosphenes and evoke saccades directed to the stimulated location in the retinotopic map. Although ICMS is widely used, little is known about the evoked spatio-temporal patterns of neural activity and their relation to neural responses evoked by visual stimuli or saccade generation. To investigate this, we combined ICMS with Voltage Sensitive Dye Imaging in V1 of behaving monkeys and measured neural activity at high spatial (meso-scale) and temporal resolution. Small visual stimuli and ICMS evoked population activity spreading over few mm that propagated to extrastriate areas. The population responses evoked by ICMS showed faster dynamics and different spatial propagation patterns. Neural activity was higher in trials w/saccades compared with trials w/o saccades. In conclusion, our results uncover the spatio-temporal patterns evoked by ICMS and their relation to visual processing and saccade generation.

Impairment but not abolishment of express saccades after unilateral or bilateral inactivation of the frontal eye fields

Express saccades are the shortest-latency saccade. The frontal eye fields (FEF) are thought to promote express saccades by presetting the superior colliculus. Here, by reversibly inactivating the FEF either unilaterally or bilaterally via cortical cooling, we support this by showing that the FEF plays a facilitative but not critical role in express saccade generation. We also found that FEF inactivation lowered express saccade peak velocity, emphasizing a contribution of the FEF to express saccade kinematics.