The oculomotor system Bijan Pesaran April 29, 2008 Classes of eye movements • Reflexive – gaze stabilization – VOR • Stabilize for head movements – Optokinetic • Stabilize for image motion • Voluntary – gaze shifting – Saccades • Acquire stationary target – Smooth pursuit • Acquire moving target – Vergence • Acquire target in depth Oculomotor muscles and nerves • Oculomotor nerve (III) – Medial rectus – Superior/Inferior recti – Inferior oblique • Trochlear nerve (IV) – Superior oblique • Abducens nerve (VI) – Lateral rectus • Medial longitudinal fasciculus Motor neurons command muscle forces • Linear increase for static forces • Pulse for dynamic forces Optokinetic reflex • Optokinetic nystagmus • Neural pathway convergent with VOR Saccadic system Brainstem saccadic control • Paramedian pontine reticular formation (PPRF) – – – – – Burst and omnipause neurons Aim to reduce horizontal motor error Project to directly to lateral rectus motor neurons Projects indirectly to contralateral medial rectus Medial longitudinal fasciculus • Mesencephalic reticular formation – Also influenced by omnipause neurons – Vertical motor error – Projects to superior and inferior rectus motor neurons Eye movements diagnose brainstem lesions • PPRF lesions impede horizontal eye movements • MRF lesions impede vertical eye movements • MLF lesions impede medial rectus contraction – Internuclear opthalmoplegia – No impact on vergence Superior colliculus • 7 layered structure. Mammalian optic tectum. • Superficial layers (3 layers) – Visual input from retina and striate cortex – Modulated by saccades but not attention • Intermediate (2) and deep (2) layers – Input from dorsal stream and FEF – Build-up and burst neurons • Topographic maps encode motor error • Fixation zone in rostral SC -> Dorsal raphe nucleus • Lesions disrupt saccades temporarily Population averaging scheme Sensory-motor transformations • Deep layers • Auditory-oculomotor – Auditory neurons – Bimodal neurons • Somatosensory-oculomotor – Body maps • Update in response to eye movements Parietal cortex • Area LIP – Early stage of movement planning – Visual responses modulated by attention • Lesions disrupt sensory-motor processes – Neglect – Optic ataxia – Balint’s syndrome Frontal cortex • Frontal eye fields – Visual, movement and visual-movement neurons – Project to PPRF and MRF – Lesions: Temporary paresis, long term memory deficit • Supplementary eye fields – Object-centered saccades • Dorsolateral prefrontal cortex – Working memory Smooth pursuit • Track movement on part of retina • Two theories – Motor (Robinson) • Retinal slip only provides velocity • Does not capture pursuit onset – Sensory (Lisberger and Krauzlis) • Position, velocity and acceleration Smooth pursuit system Smooth pursuit brainstem • Eye velocity for pursuit medial vestibular nucleus and nucleus prepositus hypoglossi – Project to abducens and oculomotor nuclei – Input from flocculus of cerebellum encodes velocity • PPRF also encodes velocity – Input from vermis of cerebellum encodes velocity • Dorsolateral pontine nucleus – Relays inputs from cortex to cerebellum and oculomotor brainstem Smooth pursuit cortex • Visual motion areas MT and MST – Active in visual processing for pursuit – Stimulation influences pursuit speed – Projects to DLPN and FEF – Does not initiate pursuit • Frontal eye fields – Stimulation initiates pursuit – Lesions diminish pursuit Vergence • Four sources – – – – Disparity Accomodation Tonic Proximal vergence • Brainstem – Burst and Burst-tonic neurons • Similar to saccadic system Coordinated vergence/version movements • Vergence starts sooner • Saccade finishes faster • Systems interact – Saccade omnipause inhibits vergence bursters 3-D eye movements • Donder’s Law – Relates torsion to eye position • Listing’s law – Torsion results from rotation of eye around perpendicular axis • Listing’s plane – Plane orthogonal to line of sight • Does not apply when head is free Clinical diagnosis from eye movements