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L. Itti: CS564 - Brain Theory and Artificial Intelligence University of Southern California Lecture 16. Saccades 2 Reading Assignments: The NSL Book The Modular Design of the Oculomotor System in Monkeys Peter Dominey, Michael Arbib, and Amanda Alexander Supplementary Reading: Crowley-Arbib Saccade Model M. Crowley, E. Oztop, and S. Marmol Laurent Itti: CS564 - Brain Theory and Artificial Intelligence. Saccades 2 1 delay FEF PP FOn PPct r ms Peter Dominey & Michael Arbib: Cerebral Cortex, 2:153-175 switch Filling in the Schemas: Neural Network Models Based on Monkey Neurophysiology qv sm vm FEF vs PP MD VisCx sm CAUDATE Vis Cx SC CD TH LGN vm SNr SNR vs SG sm delay Develop hypotheses on Neural Networks that yield an equivalent functionality: mapping schemas (functions) to the cooperative cooperation of sets of brain regions (structures) FEFvs FEFms SC vs ms qv FOn wta eye movement FEFvs FEFms Brainstem Saccade Generator Retina VisInput Last time, we saw that… - Double-saccade experiments suggest direction/amplitude coding rather than absolute target location - Lesion/stimulation studies suggest that the overall system still works when either SC or FEF is missing (but not both!) - FEF stimulation just after presentation of a visual target (SC lesioned) elicits a saccade towards the “fake” FEF target first Laurent Itti: CS564 - Brain Theory and Artificial Intelligence. Saccades 2 3 Experimental Findings FEF PP - FEF, PPC SC - SC saccade generator (SG) - FEF BG (CD and SNr) SC - connection (role in disinhibition of SC for saccades) - Simple saccade: study topographic relations between sensory and motor areas - Memory saccade: study cortical and subcortical activity that sustains spatial memory - Double saccade: study dynamic remapping of target location with intervening eye movements Laurent Itti: CS564 - Brain Theory and Artificial Intelligence. Saccades 2 4 Basic Model Element: Layer 2D array of neurons delay FEF PPct r ms topographic correspondence from layer to layer switch PP FOn qv sm vm vs VisCx sm CD external world: 27x27 array TH LGN vm SNR vs sm model retina: 9x9 layer; so, each model neuron represents a small population of biological neurons delay FEFvs FEFms SC vs ms qv FOn Laurent Itti: CS564 - Brain Theory and Artificial Intelligence. Saccades 2 wta eye movement FEFvs FEFms Brains tem Saccade Generator Retina VisInput 5 Visual Input At every iteration, eye position determines position of 9x9 retinal window within 27x27 outside world if eye velocity over 200deg/sec, retinal input is reduced (saccadic suppression) Laurent Itti: CS564 - Brain Theory and Artificial Intelligence. Saccades 2 6 Direct connection retinaSC To superficial layer of SC (vs) responsible for reflex saccades = short-latency saccades to target which has not been recently fixated Laurent Itti: CS564 - Brain Theory and Artificial Intelligence. Saccades 2 7 visual pre-processing LGN, V1, V2, V4 and MT areas abstracted by a single layer possible only because we have a very coarse (9x9) retinal input with no image noise! Laurent Itti: CS564 - Brain Theory and Artificial Intelligence. Saccades 2 8 quasi-visual cells in PP Andersen et al. (1988) found in PP cells that code for future eye movements. Quasi-visual because in double-saccade task found cells which fire at location of second target respective to first target, while there never was a retinal stimulus there! right movement field but wrong receptive field Laurent Itti: CS564 - Brain Theory and Artificial Intelligence. Saccades 2 9 Double Saccade Experiment + time Laurent Itti: CS564 - Brain Theory and Artificial Intelligence. Saccades 2 10 remapping Hypothesis: occurs primarily in PP (in reality, may occur in many regions at once, with connections between regions serving for fine-tuning). problem: eye velocity signals have not been found in PP. but eye position signals have Dominey and Arbib’s computational hypothesis: remapping is done such as to compensate for difference between current eye position, and a damped/delayed eye position signal Laurent Itti: CS564 - Brain Theory and Artificial Intelligence. Saccades 2 11 frontal eye fields Bruce & Goldberg (1984): FEF contains: - visual cells (vm) (receive input from PP) - movement cells (ms) (fire just before saccade) - visuomovement cells (sm) (memory: fire during delay in memory saccade task) - postsaccadic cells PPctr: active as long as fixation cross present (inhibits eye movements) Laurent Itti: CS564 - Brain Theory and Artificial Intelligence. Saccades 2 FOn = fixation is on 12 superior colliculus -Input from retina (reflex saccades) -Input from PPqv SC qv layer (yield saccades when FEF lesioned) -Inputs from FEF How can we choose? WTA array: saccade to currently strongest target Laurent Itti: CS564 - Brain Theory and Artificial Intelligence. Saccades 2 13 basal ganglia SNr provides tonic inhibition of SC and thalamus, unless prevented to do so by FEF (directly or via CD) Goals: - prevent saccades while a target is being fixated - memorise location of future target in memory saccade task FEF can selectively control the targets for saccades, overriding collicular attempts to initiate saccades to distracting peripheral targets Laurent Itti: CS564 - Brain Theory and Artificial Intelligence. Saccades 2 14 The Full Dominey Model DCEP trig SC PN RI dimension Retina, VisCx Delay (d1) Mechanism A FEF velocity EBN LLBN, MLBN TN MN 7a/LIP/PP Brain Stem – Saccade Burst Generator DCEP-Damped Change in Eye Position 7a/LIP-Oculomotor Region of Posterior Parietal Cortex Laurent Itti: CS564 - Brain Theory and Artificial Intelligence. Saccades 2 15 Laurent Itti: CS564 - Brain Theory and Artificial Intelligence. Saccades 2 16 Laurent Itti: CS564 - Brain Theory and Artificial Intelligence. Saccades 2 17 Laurent Itti: CS564 - Brain Theory and Artificial Intelligence. Saccades 2 18