Double Stimulation in a Spiking Neural Network Model of the Midbrain Superior Colliculus.

The midbrain superior colliculus (SC) is a crucial sensorimotor interface in the generation of rapid saccadic gaze shifts. For every saccade it recruits a large population of cells in its vectorial motor map. Supra-threshold electrical microstimulation in the SC reveals that the stimulated site produces the saccade vector specified by the motor map. Electrically evoked saccades (E-saccades) have kinematic properties that strongly resemble natural, visual-evoked saccades (V-saccades), with little influence of the stimulation parameters. Moreover, synchronous stimulation at two sites yields eye movements that resemble a weighted vector average of the individual stimulation effects. Single-unit recordings have indicated that the SC population acts as a vectorial pulse generator by specifying the instantaneous gaze-kinematics through dynamic summation of the movement effects of all SC spike trains. But how to reconcile the a-specific stimulation pulses with these intricate saccade properties? We recently developed a spiking neural network model of the SC, in which microstimulation initially activates a relatively small set of (~50) neurons around the electrode tip, which subsequently sets up a large population response (~5,000 neurons) through lateral synaptic interactions. Single-site microstimulation in this network thus produces the saccade properties and firing rate profiles as seen in single-unit recording experiments. We here show that this mechanism also accounts for many results of simultaneous double stimulation at different SC sites. The resulting E-saccade trajectories resemble a weighted average of the single-site effects, in which stimulus current strength of the electrode pulses serve as weighting factors. We discuss under which conditions the network produces effects that deviate from experimental results.