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Eye Movements
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1. The Plant
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The Oculomotor Plant Consists
Of only 6 muscles in 3 pairs
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This Yields 3 degrees of Mechanical Freedom
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Donder’s Law/ Listing’s Law
Neural Constraints Reduce this to 2 degrees of freedom
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3-D eye movements Donder’s Law Listing’s law Listing’s plane
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
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Kinematics vs Dynamics In the Oculomotor System
Rotations about the Center of Gravity No Loads No Inertia Force = Position
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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
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2. The Behaviors Gaze Holding: VOR OKN Gaze Shifting: Saccades
Vergence Smooth Pursuit
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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
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Gaze During Nystagmus
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Saccades
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3-D Gaze Trajectory Vergence
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2. The Motor Neurons
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Force Patterns Robinson’s Lollipop Experiments Statics Dynamics
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Oculomotor Neurons During Static Gaze
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Dynamics and Statics
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3. VOR
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Cupula and otoliths move sensory receptors
Cristae Maculae
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Angular Acceleration Angular Velocity Angular Position Cupula Deflection
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Canal afferents code velocity
Spontaneous activity allows for bidirectional signaling S-curve is common Different cells have different ranges and different dynamics Population code
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Canal Output During Slow Sinusoidal Rotation
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VOR With and Without Vision
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rVOR gain varies with frequency
Almost perfect > 1Hz Low gain for low frequencies (0.1Hz) Sensory mechanisms can compensate (optokinetic reflex)
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Oculomotor muscles and nerves
Oculomotor nerve (III) Medial rectus Superior/Inferior recti Inferior oblique Trochlear nerve (IV) Superior oblique Abducens nerve (VI) Lateral rectus
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The 3-Neuron Arc Primary Effects of Canals on Eye Muscles
Canal Excites Inhibits Horizontal Ipsi MR, Contra LR Ipsi LR, Contra MR Anterior Ipsi SR, Contra IO Ipsi IR, Contra SO Posterior Ipsi SO, Contra IR Ipsi IO, Contra SR
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Robinson’s Model of the VOR
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Robinson
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4. OKN
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Type I Vestib Neuron
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Bode Plot of OKN
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Bode Plot of VOR
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Bode Plot of OKN
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5. Saccades
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Saccadic system
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OPN Stimulation
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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
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Robinson’s Model of the VOR
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Lee, Rohrer and Sparks
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Jay and Sparks
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5. Pursuit
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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
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Smooth pursuit system
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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
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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
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Jergens
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Scudder
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