1 CEREBELLUM INTRO Crucial for coordination, accuracy, efficiency of movement No direct access to motorneurons or to spinal cord Indirect role Gather and integrate signals from - relevant sensory modalities - pre-motor interneuron pools Modulate other descending pathways Air traffic control analogy (can fly without it, but uncoordinated, unsafe) Cerebellar lesions produce: No paralysis No sensory loss Ataxia (incoordination) GROSS STRUCTURE Surface features Hemisphere Vermis Flocculo-nodular lobe Folia Peduncles One per brainstem division (lesions in each division can produce ataxia by affecting these fibers) Input vs. output fibers (by peduncle) Middle mainly input; superior mainly output; inferior both Internal anatomy Cortex White matter Deep nuclei Dentate Interpositus (= globus + emboliform) Fastigial MICROSCOPIC ANATOMY Layers of cortex Molecular layer Granule-cell layer Purkinje cell layer Purkinje cells as sole cortical output Input fibers Mossy fibers granule cell parallel fibers (granule cell axons) Climbing fibers direct contacts onto Purkinje cells 2 Interneurons Several (basket, stellate, Golgi) Basket-cell example Lies in molecular layer Parallel fiber inputs Inhibit Purkinje soma and hillock Lateral inhibition (flanks a "beam" of Purkinje cells excited by their parallel fibers) Purkinje cells are inhibitory Sculpting of high resting activity of deep nuclei INPUT PATHWAYS Logic - what need for coordination Muscle / joint (from SPINAL CORD) Cutaneous (from CORD) Interneuron pool status (from CORD) Orientation with respect to gravity; movement (VESTIBULAR) Ongoing motor plans with respect to complex extrapersonal space (from NEOCORTEX) These classes of information target different cerebellar regions Vestibulo-cerebellum (flocculo-nodular lobe and vermis) Spino-cerebellum (vermis and para-vermal hemisphere) Cerebro-cerebellum (cerebellar hemisphere) Spinal inputs SPINOCEREBELLUM Vermis Paravermal Topography (axial structures represented at midline vermis) IPSILATERAL relationship of cerebellum to spinal cord and body DORSAL SPINOCEREBELLAR TRACT Afferents: Class I and II; muscle, joint, cutaneous Dorsal columns Clarke's column (thoracic) Lateral funiculus (relatively dorsal there, hence the tract's name) Inferior cerebellar peduncle Spinocerebellum CUNEO-CEREBELLAR TRACT Upper-limb equivalent of dorsal spinocerebellar tract Afferents same Relay: external cuneate nucleus (medulla) VENTRAL SPINOCEREBELLAR TRACT Origin: interneuron pools Does carry some crude sensory information But mainly "premotor" signals (status of interneurons) 3 Decussation Lateral funiculus Relatively ventral there, hence tract name Superior peduncle Cerebellum Crossed pathway? Doesn't fit rule, but - some recross - axial representation which typically gets bilateral input ROSTRAL SPINOCEREBELLAR TRACT Upper-limb equivalent of ventral spinocerebellar Vestibular inputs to cerebellum Secondary sensory mainly Some primary (from Scarpa's ganglion) Cerebellar targets Vestibulocerebellum Vermis Cerebral Cortical inputs to cerebellum Pathway indirect via pontine nuclei Cerebellar targets Hemisphere (cerebro-cerebellum) Paravermis (convergence with spinal input) Major component of cortico-brainstem fibers Of fibers in cerebral peduncle - fully 50% participate in corticopontine pathway - only 5% contribute to corticospinals Explains why medullary pyramid so much smaller than cerebral peduncle Decussation of ponto-cerebellar fibers Preserves cortical and cerebellar lateralities Transverse fibers visible in sections Inferior olive All above systems terminate as mossy fibers Olive as sole climbing-fiber source Inputs to olive Spinal Brainstem (midbrain) Cerebral cortex Olivo-cerebellar fibers are crossed (decussating) Reticular formation Diverse functions Partly, feedback from origin of one descending path used by cerebellum Other output relays also feed back (e.g., vestibular nuclei; cerebral cortex; red nuclei). 4 FUNCTIONAL ZONES OF CEREBELLUM (refer to table) Inputs distinguish (see above) Outputs to deep nuclei differ, which in turn relate them to different… - muscle groups (preferential targeting of axial; proximal limb/girdle; or distal muscle groups) - functional association (balance/posture; gait; fine manipulation) - descending pathways REGION ZONE DIVISION Flocculonodular Vermis Flocculonodular Median Vestibulocerebellum Paravermal Paramedian or Intermediate Hemisphere Lateral Spinocerebellum Cerebrocerebellum DEEP NUCLEUS Vestibular nuclei Fastigial Interpositus Dentate INPUTS MUSCLES FUNCTION Vestibular (1º and 2º) Spinal (trunk) and vestibular Spinal (limb) & corticopontine Axial Equilibrium Posture Equilibrium Posture Corticopontine All, especially distal limb Axial Limb Posture Gait Global, especially fine movement, planning CEREBELLUM Cerebral Cortex Lateral Paramedian Median Flocculo-nodular Dentate Interpositus Fastigial Vestibular nuclei Thalamus (VL) Red Nucleus Reticular Formation Spinal Cord 5