DESCENDING TRACTS Fiber Types A Fibers: Somatic, myelinated. Alpha (α): Largest, also referred to as Type I. Beta (β): Also referred to as Type II. Gamma (γ): Delta (δ): Smallest, referred to as Type IV. Fiber Types B Fibers: Lightly myelinated. Preganglionic fibers of ANS. C Fibers: Unmyelinated. Found in somatic and autonomic systems. Also referred to as Type IV fibers. Fiber Types Sensory fibers are either: A-α or A-β fibers: Conduction rate = 30-120 m/sec. A-δ fibers: Conduction rate = 4-30 m/sec. C fibers: Conduction rate is less than 2.5m/sec. Fiber Types Nociceptors and thermoreceptors are related to C fibers or A-δ fibers. Generalizations: Motor Paths Typical descending pathway consists of a series of two motor neurons: Upper motor neurons (UMNs) Lower motor neurons (LMNs) Does not take into consideration the association neurons between UMNs and LMNs Upper Motor Neurons Are entirely within the CNS. Originate in: Cerebral cortex Cerebellum Brainstem Form descending tracts Lower Motor Neurons Begin in CNS. From anterior horns of spinal cord. From brainstem cranial nerve nuclei. Made up of alpha motor neurons (A-α). Make up spinal and cranial nerves. UMN Classification Classified according to where they synapse in the ventral horn: Medial activation system: Innervate postural and girdle muscles Lateral activation system: Associated with distally located muscles used for fine movements Nonspecific activating system: Facilitate local reflex arcs Pyramidal System Characteristics: Upper motor neurons: 75 – 85% Decussate in pyramids. Remainder decussate near synapse with lower motor neurons. Most synapse with association neurons in spinal cord central gray. Pyramidal System Components: Corticospinal Tract Corticobulbar Tract Corticospinal Tract Divisions Lateral corticospinal tract: Made up of corticospinal fibers that have crossed in medulla. Supply all levels of spinal cord. Anterior corticospinal tract: Made up of uncrossed corticospinal fibers that cross near level of synapse with LMNs. Supply neck and upper limbs. Corticospinal Tract Functions Add speed and agility to conscious movements: Especially movements of hand. Provide a high degree of motor control: (i.e., movement of individual fingers) Corticospinal Tract Lesions Reduced muscle tone Clumsiness Weakness Not complete paralysis Note: complete paralysis results if both pyramidal and extrapyramidal systems are involved (as is often the case). Corticobulbar Tract Innervates the head Most fibers terminate in reticular formation near cranial nerve nuclei. Association neurons: Leave reticular formation and synapse in cranial nerve nuclei. Synapse with lower motor neurons. Extrapyramidal System Includes descending motor tracts that do not pass through medullary pyramids or corticobulbar tracts. Includes: Rubrospinal tracts Vestibulospinal tracts Reticulospinal tracts Rubrospinal Tract Begins in red nucleus. Decussates in midbrain. Descends in lateral funiculus (column). Function closely related to cerebellar function. Lesions: Impairment of distal arm and hand movement. Intention tremors (similar to cerebellar lesions) Vestibulospinal Tract Originates in vestibular nuclei: Receives major input from vestibular nerve: (CN VIII) Descends in anterior funiculus (column). Synapses with LMNs to extensor muscles: Primarily involved in maintenance of upright posture. Reticulospinal Tract Originates in various regions of reticular formation. Descends in anterior portion of lateral funiculus (column). Thought to mediate larger movements of trunk and limbs that do not require balance or fine movements of upper limbs. BASAL NUCLEI Basal Ganglia Functions Compare proprioceptive information and movement commands. Sequence movements. Regulate muscle tone and muscle force. May be involved in selecting and inhibiting specific motor synergies. Basal Ganglia Functions Basal ganglia are vital for normal movement but they have no direct connections with lower motor neurons. Influence LMNs: Through planning areas of cerebral cortex. Pedunculopontine nucleus of midbrain. Basal Ganglia Functions Basal nuclei set organism’s level of responsiveness to stimuli. Extrapyramidal disorders are associated with basal nuclei pathology: Negative symptoms of underresponsiveness: Akinesias i.e. Parkinson disease Positive symptoms of over-responsiveness: Choreas, athetoses, ballisms i.e. Huntington’s chorea Basal Nuclei Components Corpus striatum Substantia nigra (within the midbrain) Subthalamic nuclei (diencephalon) Red nucleus (?) Claustrum (?) Nucleus accumbens (?) Corpus Striatum Composed of caudate nucleus + lentiform nucleus: Striatum = caudate nucleus + putamen. Pallidum = globus pallidus. Putamen + globus pallidus = lentiform nucleus. Controls large subconscious movements of the skeletal muscles. The globus pallidus regulates muscle tone. Corpus Striatum Corpus Striatum (Telencephalon) Striatum Caudate Nucleus Pallidum Putamen Globus Pallidus Substantia Nigra Subdivisions Dorsal pars compacta: Has melanin containing neurons and dopaminergic neurons. Ventral pars reticularis: Has iron-containing glial cells. Has serotonin and GABA (no melanin). Substantial Nigra Substantia Nigra Mesencephalon Dorsal Pars compacta Ventral Pars reticularis Melanin containing neurons Dopaminergic neurons Iron-containing glial cells Serotonin and GABA Input Nuclei Striatum Caudate nucleus Putamen Nucleus accumbens Receive widespread input from: Neocortex Intralaminar nuclei Substantia nigra Dorsal raphe nucleus Input Nuclei Striatum projects to: Globus pallidus Substantia nigra Pars reticularis Via gabaminergic fibers Motor and sensory cortices project to putamen. Association areas of all lobes project to caudate nucleus. Output Nuclei Globus pallidus (medial part) Substantia nigra: Pars reticularis Ventral pallidum Fibers project to: VA/VL nuclei Mostly inhibitory General Core Circuit Cerebral cortex to: Striatum to: Globus pallidus to: Thalamus to: Portions of motor cortex to: Upper motor neurons Thalamic Fasciculi Ansa lenticularis: Consists of fibers from dorsal portion of globus pallidus. Loops under internal capsule. To VA/VL complex. Thalamic Fasciculi Lenticular fasciculus: Consists of fibers from ventral portion of globus pallidus. Passes across the internal capsule. To VA/VL complex. Dopamine Neuronal System Consists of nigrostriatal fibers From pars compacta of substantia nigra To striatum Dopaminergic Direct Basal Ganglia Circuit Motor cortex projects to putamen: Excitatory (glutamate) Putamen projects to output nuclei (globus pallidus internus and substantia nigra reticularis): Inhibitory (GABA and substance P) Basal Ganglia Connections Red = excitatory; Black = Inhibitory Motor areas of cerebral cortex Ventrolateral thalamus Putamen Globus pallidus externus Output nuclei PedunculoPontine nuclei Reticulospinal and Vestibulospinal pathways Subthalamic nuclei Substantia nigra compacta Lateral Activation pathways Direct Basal Ganglia Circuit Output nuclei project to motor thalamus (VA-VL) and pedunculopontine nuclei: Inhibitory (GABA) Ventrolateral (VA-VL) thalamus projects to motor cortex: Excitatory Therefore: Increasing input to putamen increases activity in corticofugal fibers Direct Basal Ganglia Circuit Pedunculopontine nuclei project to reticulospinal and vestibulospinal pathways. Stimulation of pedunculopontine nuclei elicit rhythmical behaviors such as locomotor patterns. Indirect Basal Ganglia Circuit Motor cortex to putamen: Excitatory (glutamate) Putamen to globus pallidus externus: Inhibitory (GABA and enkephalins) Globus pallidus externus to subthalamic nuclei: Inhibition (GABA) Indirect Basal Ganglia Circuit Subthalamic nuclei to output nuclei (substantia nigra reticularis) Excitatory (glutamate) Output nuclei to VA-VL complex (motor thalamus) Inhibitory (GABA) Indirect Basal Ganglia Circuit VA-VL complex to motor cortex: Excitatory Therefore: decrease in corticofugal pathways. Input from Substantia Nigra Compacta Projects to putamen: Excitatory (dopamine) Two kinds of receptors in basal ganglia circuit: D1: facilitates activity in direct pathway D2: inhibits activity in indirect pathway