Motor System Hierarchy

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Motor Pathways

Lectures Objectives Define the terms upper and lower motor neurons with examples. Describe the corticospinal (pyramidal) tract and the direct motor pathways from the cortex to the trunk and limbs. Briefly describe the indirect motor pathways from the cortex to the trunk and limbs through extrapyramidal tracts such as rubrospinal and reticulospinal tracts. Describe motor pathways to the face muscles. Compare the signs and symptoms of the upper and lower motor neuron lesions. Identify the centers that make the basal ganglia. Identify the different parts, regions and nuclei of the cerebellum. Summarize the motor system circuitry.

Motor System Hierarchy Control Systems Initiator Executers Lower motor neuron Muscle Basal Ganglia Cerebellum Cortex Upper motor neurons Lower motor neuron Muscle

Upper and Lower Motor Neurons

Lower Motor Neurons Neurons innervates muscles Alpha motoneurons Innervates normal fibers Gamma motor neurons Innervates fibers in muscle spindle Present in: Spinal cord Brainstem (in the cranial nerves nuclei) Muscle tone

Muscle Tone Determined by the level of activity in the lower motor neurons Tone refers to the resistance of a muscle to passive stretch Primary determinant of muscle tone is the level of activity in the stretch reflex

Upper Motor Neurons Project to LMN Innervate α and γ motor neurons and inhibitory interneurons Location of UMN Reticular formation: reticulospinal tract Vestibular nuclei: vestibulospinal tract Superior colliculus: tectospinal tract Red nucleus: rubrospinal tract Motor cortex: corticospinal & corticobulbar tracts (+ corticorubral & corticoreticular)

Descending Tracts Classification Classical classification: Pyramidal system Axons traverse the pyramids in medulla Corticospinal axons Extrapyramidal system Other descending tracts Basal ganglia and their connections with motor region Functional classification: Medial system Innervate medial motor nucleus Lateral system Innervate lateral motor nucleus

Descending Tracts Lateral pathways Terminate laterally in the ventral horn Crossed Involved in movements of the distal limbs (initiation & fine movement) Damage weakness or paralysis Contains Lateral corticospinal tract Ruprospinal tract

Ruprospinal tract From red nucleus Crossed immediately

The Pyramidal (Corticospinal & corticobulbar) Tract Origin cerebral cortex ⅓ from primary motor cortex ⅓ from premotor areas ⅓ from primary somatosensory cortex Terminate in dorsal horn and brainstem Modify processing in the somatosensory pathways

Corticospinal Tract Axons pass through: Corona radiata Internal capsule (posterior limb) Basis pedunculi midbrain Medullary pyramids Decussation Crossed Lateral corticospinal 90 95% Uncrossed Anterior corticospinal tract

Lateral Corticospinal Tract Somatotopic Organization

Corticobulbar Tract Same origin & course as corticospinals Leave tract in brainstem Terminate in the cranial nerve nuclei Bilateral in general Except to facial nucleus

Descending Tracts Medial pathways Terminate Medially in the ventral horn Trunk & proximal limb movements Important in maintaining balance & position Mostly happened unconsciously Damage Deficits in maintaining balance & posture Changes in muscle tone Contains: Reticulospinal tract Vestibuspinal tract Tectospinal tract Anterior corticospinal tract

Reticulospinal Tract From the reticular formation Important influence on muscle activity and reflexes Controlled by cortex (corticoreticular) Contains descending autonomic fibers controlled by thalamus Crossed and uncrossed Two tracts Lateral reticulospinal tract from Medulla Medial reticulospinal tract from Pons

Vestibulospinal Tract Importance in maintaining balance Influence axial muscles Uncrossed Medial & lateral vestibulospinal tracts

Tectospinal Tract Superior colliculus Terminate in the cervical regions Head movements in response to visual stimuli Mostly crossed

Anterior Corticospinal Tract Similar to other medial pathways Terminate in the medial motor n. Except it is voluntary Do not cross in pyramidal decussation May cross before termination

Lower Motor Neuron Lesion Flaccid paralysis or paresis (weakness) Hypo or areflexia Decreased muscle tone Atrophy muscle wasting Develops over time (weeks) Fasciculations small twitches that are visible to the eye

Upper Motor Neuron Lesion Paralysis or paresis Spasticity Hypertonia Hyperreflexia And maybe: Babinski sign Clonus Decreased superficial reflexes Abdominal reflex & Cremasteric reflex

Upper Motor Neuron Lesion Babinski Sign Abnormal response to stroking the lateral planter surface of the foot Not useful in babies Normal response: toes planter flex Abnormal: dorsiflexion of big toe

Upper Motor Neuron Lesion Clonus Repetitive flexion extension of a joint in response to single flexion or extension https://youtu.be/ux75k8s5que

Superficial Reflexes Decrease With UMN lesions Abdominal reflex; abdominal muscles contract on stroking the abdomen Cremasteric reflex (useful in babies); testes elevation with stroking inside of the thigh

Lower-versus Upper-Motor- Neuron Lesions Variable Lower-Motor-Neuron Lesion Upper-Motor-Neuron Lesion Weakness Flaccid paralysis Spastic paralysis Deep tendon reflexes Decreased or absent Increased Babinski's reflex Absent Present Atrophy May be marked Absent or resulting from disuse Fasciculations and fibrillations May be present Absent

Spinal Shock Follows severe acute injury to the spinal cord For short period (days or weeks) Loss of all functions (motor & sensory) bellow level of injury Loss of reflexes Due to sudden loss of supraspinal inputs

Upper Motor Neurons Cerebral Cortex Lateral system Medial system Motor cortex Red nu. Spinal cord & Brainstem Retic. Form. Sup. Colliculus Vestibular Nu. γ α Intrafusal Muscle Extrafusal

Basal Nuclei (Ganglia)

Basal Ganglia The basal ganglia include the caudate, putamen, and globus pallidus and number of closely related nuclei They influence motor system primarily through projections to upper motor neurons Motor deficits depend on the specific nucleus damaged Understanding the neurochemistry of basal ganglia drives the development of clinical treatment

Basal Ganglia The basal ganglia act as Brake against involuntary movement Switch to turn on a fixed action pattern Their major output is to the VA of the thalamus Projects primarily to area 6 (premotor & supplementary motor areas)

Basal Ganglia Terminology Striatum (neostriatum) = caudate + putamen Lentiform nucleus = putamen + globus pallidus Corpus striatum = caudate + lentiform Basal ganglia = corpus striatum + amygdala Globus pallidus = pallidum = paleostriatum Claustrum is some times included with the basal ganglia Basal ganglia is included by the extrapyramidal system

Caudate nucleus Parts Location Relations Lateral ventricle Amygdaloid nucleus Basal Ganglia: Gross Anatomy

Lentiform nucleus Parts Putamen Globus pallidus Internal (GPi) External (GPe) Shape Location Relations External & internal capsules Claustrum Basal Ganglia: Gross Anatomy

Basal Ganglia: Gross Anatomy Amygdaloid nucleus Subthalamic nucleus Substantia nigra Pars reticulata (SNr) Pars compacta (SNc) Claustrum

Inputs Most inputs enter the striatum From cerebral cortex & thalamus These inputs are excitatory Outputs Most leave from Gpi & SNr Most go to VA nucleus of the thalamus, which projects to motor cortex The outputs are GABAergic and inhibitory VA excites motor cortex, leading to movements Increase basal ganglia output will inhibit the VA and reduce overall movements Basal Ganglia Circuitry

Basal Ganglia Circuitry Intrinsic Circuits Large number of connections between components of the basal ganglia Can be grouped into Direct pathway Indirect pathway These pathways affect the VA activity and thus the motor cortex activity

From striatum to Gpi Uses GABA, which inhibits another GABAergic projection (Gpi to VA) Disinhibition Cortical activity direct pathway Gpi activity VA activity Activity in the direct pathway leads to increased motor cortex activity and increased movements The Direct Pathway

Goes from striatum to GPe (GABA) to the subthalamic nucleus (GABA) Subthalamic nucleus to Gpi (Glu) activity in the cortex activity of subthalamic nu. GPi VA activity motor cortex activity The Indirect Pathway

Basal Ganglia Circuitry The direct pathway increase movements The indirect pathway decrease movements Normal behavior requires a balance between the direct and indirect pathways All pathways are uncrossed Right basal ganglia modulate right cortex and affect movements on the left side of the body Acetylcholine is used by the interneurons in the striatum It affect the output of the direct and indirect pathways It s a target for drug therapy

In the striatum different cell types give rise to the direct and indirect pathways Both cell types receive dopaminergic input from SN pars compacta These cells have different receptors for DA For direct pathway, DA excites the striatal cells For indirect pathway, DA inhibits the striatal cells Thus the nigrostriatal pathway the activity of the VA and motor cortex PD leads to direct pathway activity indirect pathway activity activity of VA and motor cortex Nigrostriatal Pathway

Cerebellum

Cerebellum The cerebellum is essential for normal movements It affects motor behavior by affecting UMNs The cerebellum acts as a comparator Compares intended movements (data from cerebral cortex) to the actual movements (sensory data) Sends corrective signals into the descending motor pathways

Cerebellar Function It affects all movements, it is important in: Balance Locomotion Simple & complex movements Eye movements, etc. Site of motor learning Important for learning new motor skills and adjusting movements to changing sensory inputs

Location. Relations.. The cerebellum consists of two hemispheres The hemispheres are connected by vermis Cerebellar Anatomy Gross Anatomy

Cerebellum: Gross Anatomy Three main lobes Anterior lobe Primary fissure Posterior lobe (middle lobe) Cerebellar tonsils Posterolateral fissure (uvulonodular fissure) Flocculonodular lobe

Cerebellum: Gross Anatomy

Cerebellum: Internal Structure Content Cerebellar cortex (folia) & central nuclei are grey matter Arbor vitae = tree of life = white matter

Cerebellum includes a cortex & deep nuclei The deep nuclei are the major source of output from the cerebellum Four nuclei from medial to lateral Fastigial Globose Emboliform Dentate Cerebellar Anatomy Interposed nuclei

Cerebellar cortex includes 5 cell types in 3 layers Five cell types Inhibitory cells Purkinje, basket, Golgi, and stellate cells Excitatory cells Granule cells Three layers Molecular layer Purkinje cell layer Granule cell layer Cerebellar Cortex

Inputs to the cerebellum Climbing fibers From inferior olivary complex (olivocerebellar fibers) Decussate Inferior cerebellar peduncle Mossy fibers All remaining inputs: spinal cord, vestibular n. & nuclei, & pontine nuclei Each type of input fibers branches Branch to deep nuclei Branch to cerebellar cortex Cerebellar Inputs

Cerebellar Circuit The basic cerebellar circuit includes Main excitatory loop Inhibitory cortical side loop

The Main Excitatory Loop Includes the input and the deep cerebellar nuclei Both the inputs & the cells of the deep nuclei are excitatory

The Inhibitory Cortical Side Loop Serves to modulate the activity in the deep cerebellar nuclei Mossy & climbing fibers are inputs to cerebellar cortex Climbing fibers contact Purkinje cells directly Mossy fibers contact granule cells Granule cells contact Purkinje cells Output of cerebellar cortex (Purkinje fibers) depend on the mossy & climbing fibers

The Inhibitory Cortical Side Loop Remaining cells (Golgi, basket & stellate) are inhibitory interneurons Alter granule & Purkinje cells Purkinje cells (cerebellar cortex output) are inhibitory Purkinje cells targets deep cerebellar nuclei & vestibular nuclei Thus cerebellar output is driven by the main excitatory loop and limited by the inhibitory cortical side loop

Cerebellar Functional Divisions 1. Vestibulocerebellum Flocculonodular lobe & fastigial nu. Balance, eye movements 2. Spinocerebellum Vermis & paravermal parts of hemispheres & interposed nuclei (emboliform & globose) Motor execution 3. Cerebrocerebellum Lateral hemispheres & dentate nu. Motor planning

Vestibulocerebellum Function Balance & eye movements Inputs Vestibular n. fibers Vestibular nuclei Inferior olive Deep nucleus Fastigial nucleus Outputs (From fastigial nu. & Purkinje cells) Vestibular nuclei Reticular formation VL of thalamus Part of motor system targeted UMNs of medial pathway Major signs of damage Staggering or falling, nystagmus

Function Spinocerebellum Execution of movement Compensates for changes in load, regulates muscle tone, guides limb movement, helps maintain posture Organized somatotopically Head & trunk vermis Limbs paravermal areas Inputs Spinal & trigeminal inputs Inferior olive Deep nucleus Fastigial & interposed nuclei Outputs Vermis Reticular formation & Vestibular nu. Paravermal Red nucleus, VL of thalamus & Inferior olive Part of motor system targeted UMNs of medial & lateral pathways Major signs of damage Staggering gait, intention tremor

Cerebrocerebellum Function Coordination, planning of voluntary movements Inputs Pontine nuclei (relaying information from sensory & motor cerebral cortex) Inferior olive Deep nucleus Dentate nucleus Outputs Red nucleus (to inferior olive, back to cerebellum) VL of thalamus Part of motor system targeted Motor cortex (via VL) Major signs of damage Decomposition of movements

Cerebellar Peduncles Peduncle Inferior Restiform body Juxtarestiform body Middle (brachium pontis) Major inputs to cerebellum Fibers from: Inferior olive (climbing fibers) Dorsal spinocerebellar tract Cuneocerebellar tract Vestibular nerve Vestibular nuclei Pontine nuclei (relay inputs from cerebral cortex) Major outputs from cerebellum Fibers to: Vestibular nuclei None Superior (brachium conjunctivum) Ventral spinocerebellar Rostral spinocerebellar Red nucleus VL thalamus Reticular formation Inferior olive

Cerebellar Circuitry Cerebra l Cortex Motor cortex SCP V A V L Thalamus Red nu. Inferior olive Retic. Form. Sup. Colliculus Vestibular Nu. UMNs Lateral system Medial system Spinal cord & Brainstem γ α Pontine nuclei MCP Deep nuclei Cortex + + Cerebellum ICP Vestibular nerve Intrafusal Extrafusal

Blood Supply of Cerebellum SCA Superior cerebellar hemispheres Superior vermis Dentate nucleus Most of white matter Superior cerebellar peduncle AICA Middle cerebellar peduncle Flocculus Anteroinferior surface of the cerebellum PICA Posteroinferior cerebellar hemispheres Inferior portion of the vermis Inferior cerebellar peduncle

Motor System Intralaminar nu. Of thalamus SN compacta Basal Ganglia Globus pallidus Cerebral Cortex Glu + Striatum GABA GABA GPe Gpi SNr GABA Glu Subthalamic nucleus Motor cortex Glu VA VL Dorsal thalamus Red nu. GABA To other UMNs: Retic. Form. & Sup. Coll. Upper Motor Neurons Spinal cord & Brainstem SCP Inferior olive Retic. Form. Sup. Colliculus Vestibular Nu. γ α Pontine nuclei MCP Deep nuclei Cortex + + Cerebellum ICP Vestibular nerve Intrafusal Extrafusal