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Spinal cord: ascending & descending pathways 2012/3/26 Josh Wu Spinal cord • White matter (pathways) abundant in upper cord contain sensory & motor pathways serving all 4 limbs o cuneate fasciculus (CF) carries information from UE only in C segments ogracile fasciculus (GF) carry information from LE present in both C & L levels • Gray matter (neuron cell body) cervical & lumbosacral enlargements innervation of limbs Types of spinal neuron • In thick sections of spinal cord nerve cells exhibit a laminar (layered) arrangement. • True lamination is confined to post. horn • 10 laminae of Rexed defined in gray matter correlate findings from animal research in different labs Types of spinal neuron • Smallest neurons soma diameters 5-20μm propriospinal entirely contained within cord Some are confined within a single segment; others span 2 or more segments by way of neighboring propriospinal tract Functions 1. participate spinal reflexes 2. intermediate cell stations motor projection: interposed between fiber tracts descending from brain to locomotor apparatus () sensory transmission ( to CNS) Types of spinal neuron • Medium-sized neurons soma diameters 20-50μm in all parts of gray matter except substantia gelatinosa Function relay (projection) cells: receiving inputs from post root afferents and projecting to brain () forming tracts (a functionally homogeneous group of fibers) • Largest neurons soma 50-100μm alpha motor neurons (αMN): supply skeletal muscles gamma motor neurons (γMN) (smaller): supply muscle spindles Renshaw cells (med part of ant horn): tonic inhibition on αMN Other structures • individual ganglion surround with modified Schwann cells called satellite cells • stem axon: bifurcates • ganglion cells enter post. n. roots Exception: visceral afferents enter cord by ventral roots failure post. rhizotomy (surgical section of post roots) to relieve pain for intra-abdominal cancer Spinal ganglia • located in intervertebral foramina ant. + post. roots = spinal n. thoracic ganglia: 50,000 unipolar neurons ganglia serving limbs: 100,000 cells • cord shows no evidence of segmentation internally • Reality: nuclear groups in cell columns spanning several segments Central terminations of post root afferents • medial stream medium + large fibers divide within post funiculus for ascending/descending branches swing into post gray horn synapse in laminae II, III, & IV largest ascending fibers run all way to post column nuclei (gracilis-cuneatus) in medulla oblongata these long fibers form bulk of FG & FC • lateral stream small (Aδ + C) fibers divide into short ascending/descending branches within posterolateral tract of Lissauer synapse on neurons in lamina I (marginal zone), lamina II (substantia gelatinosa), and some dendrite to laminae III-V ASCENDING SENSORY PATHWAYS • Conscious sensations (perceived at cerebral cortex) 1. Exteroceptive sensations From external world somatic receptors on body surface or telereceptors serving vision/hearing include touch, pressure, heat, cold, and pain. 2. Proprioceptive sensations arise within body locomotor system (muscles, joints, bones) and vestibular labyrinth pathways to cerebral cortex for position (stationary) & kinesthetic (movement) sense • Non-conscious sensations (refer to cerebellum) 1. Non-conscious proprioception to cerebellum through spinocerebellar pathways essential for smooth motor coordination 2. Interoception (Enteroception) unconscious afferent signals involved in visceral reflexes Sensory testing • Routine assessment of somatic exteroceptive sensation includes: 1. Touch: by finger tip or a cotton swab 2. Pain: by applying point of a pin 3. Thermal sense: by warm or cold test tubes In alert & cooperative patients, active & passive tests of conscious proprioception can be performed • Active proprioception tests: execute activities with eyes closed 1. toe the line without swaying 2. finger-to-nose test 3. heel-to-knee test • Passive proprioception tests: 1. Joint sense (mainly by passive stretching of neuromuscular spindles) clinician grasps thumb or great toe by sides and moves it while asking patient to name direction of movement ('up' or 'down') 2. Vibration sense. vibrations of a tuning fork applied to radial styloid process or to shaft of tibia Question 1 heel-to-knee test 1 2 3 4 Question 2 1 2 3 4 Question 3 1 2 3 4 Ascending Pathways Somatic Sensory Perception posterior column-medial lemniscal pathway (PCML) spinothalamic pathway SOMATIC SENSORY PATHWAYS Common features: • Both comprise 1st, 2nd, 3rd-order sets of sensory neurons • 1st-order somas (primary afferents) occupy post. root ganglia • 2nd-order somas occupy CNS gray matter on same side as 1st-order neurons • 2nd-order axons cross midline and then terminate in thalamus • 3rd-order neurons project: thalamus somatic sensory cortex • Both pathways are somatotopic: orderly map of body parts can be identified in gray matter • Synaptic transmission from 1 2 & 23 neuron can be modulated (inhibited or enhanced) by other neurons 3rd 1st 2nd spinothalamic pathway Posterior Column-Medial Lemniscal pathway • 1st-order afferents include largest somas in post. root ganglia receive info from largest sensory receptors: 1.Meissner's and Pacinian corpuscles 2.Ruffini endings 3.Merkel cell-neurite complexes 4.neuromuscular spindles 5.Golgi tendon organs LE: FG (fasciculus gracilis) UE: FC (fasciculus cuneatus) • 2nd-order afferent LE: FG NG (nucleus gracilis) UE: FC NC (nucleus cuneatus) • Crossed midline in great sensory decussation of medulla oblongata medial lemniscus pons & midbrain terminates in VPL (ventral posterolateral nucleus) of thalamus • trigeminal lemniscus terminating in VPM (ventral posteromedial nucleus) • 3rd -order afferents thalamus somatic sensory cortex Function of PCML pathway • Chief functions conscious proprioception + discriminative touch provide parietal lobe with an instantaneous body images both at rest & during movement Without background information impair execution of movements • Clinical correlation disturb PCML in demyelinating diseases, e.g. multiple sclerosis sensory ataxia: movement disorder resulting from sensory impairment P’t can stand unsupported only with feet well apart and with gaze directed downward broad-based gait, Max stamping action for remains conscious proprioceptive function severe swaying as patient stands feet together with eyes closed tandem Romberg's sign: Inability to 'toe the line' with eyes closed loss kinesthetic sense: finger-to-nose and/or heel-to-knee tests two-point discrimination test impair of tactile discrimination joint sense and vibration sense may also be impaired Tactile, painful, and thermal sensations are preserved Ascending Pathways • Uncrossed • Proprioception • Touch (2 point discrimination) Spinothalamic pathway 2nd-order sensory neurons • projecting from laminae I-II, IV-V of post gray horn to contralateral thalamus • receive excitatory & inhibitory synapses from neurons of substantia gelatinosa 'gating' (modulatory) effects on sensory transmission • cross midline in ant. commissure at all segmental levels • anterolateral pathway is divisible into ASTT (anterior spinothalamic tract) ant. funiculus LSTT (lateral spinothalamic tract) lat. funiculus merge in brainstem as spinal lemniscus joined by trigeminal afferents VP (ventral post) nucleus of thalamus 3rd-order sensory neurons • project from thalamus to somatic sensory cortex Functions of Spinothalamic pathway • modality segregation: postoperative sensory testing ASTT touch LSTT thermal & noxious sensations • Both LSTT & ASTT are somatotopically arranged front back neck leg Percutaneous cordotomy • interrupt spinothalamic pathway on one or both sides for relief of intractable pain 1. passed needle between atlas/axis into subarachnoid space 2. advanced into anterolateral region of cord under radiologic guidance 3. pass mild current elicit paresthesia (tingling) on opposite side of body destroy anterolateral pathway • P’t is insensitive to pinprick, heat, or cold on opposite side • touch sensitivity • performed for terminal cancer patients not benign wears off after about a year Ascending Pathways thermal noxious crossed Touch Spinocerebellar pathways • 4 fiber tracts run from spinal cord to cerebellum 1. PSCT (posterior spinocerebellar) non-conscious proprioception 2. cuneocerebellar report continuously state of internuncial 3. ASCT (anterior spinocerebellar) neurons in spinal cord 4. RSCT (rostral spinocerebellar) Non-conscious proprioception (1) • both uncross control by each cerebellar (2) hemisphere (3) PSCT (post. spinocerebellar tract) (4) from LE primary afferents enter FG nucleus dorsalis originates in post thoracic nucleus in lamina VII at base of post gray horn nucleus extends T1-L1 segmental levels very fast conduction largest fibers in CNS (ext diameter: 20μm) receives primary afferents from m. & joints also receives collaterals from cutaneous sensory neurons tract ascends close to surface of cord & enters inferior cerebellar peduncle cuneocerebellar tract from accessory NC UE & upper trunk through FC enters inferior cerebellar peduncle Stretch reflex Ia internuncial serving reciprocal inhibition non-conscious proprioception kinesthesia Information from reflex arcs • 2 tracts originate in intermediate gray matter of cord • their main function is to monitor activity state of spinal reflex arcs ASCT • lower half of cord component fibers cross initially run close to surface midbrain superior cerebellar peduncle & recross within cerebellar white matter RSCT • upper half of cord ascends without crossing & enters inferior cerebellar peduncle SRT (Spinoreticular tracts) • • • • oldest somatosensory pathways half of SR fibers may be uncrossed without somatotopically arranged SR fibers originate in laminae V-VII accompany spinothalamic pathway brainstem • 2 interrelated functions 1. induce cerebral cortex waking state 2. report to limbic cortex of ant cingulate gyrus about nature sti pleasurable or aversive • phylogenetically old 'paleospinothalamic' pathways reticular formation emotional somatic sensory sti. ant. SRT post. OTHER ASCENDING PATHWAYS • ST (spinotectal tract): runs alongside spinothalamic pathway resembles in its origin & functional composition ends in superior colliculus (joins crossed visual inputs) involved in visuospinal reflex • SOT (spinoolivary tract): sends tactile information to inf. olivary nucleus in medulla oblongata important function in motor learning modify cerebellar activity in response to environmental change motor adaptation • spinocervical tract well developed in cat (small spinothalamic pathways) vestigial or absent in humans Summary of Ascending Pathways PCML (motor) Spinocerebellar emotional Spinothalamic (Pain, Touch) Question 4 (2 point) • pathway for this functional performance 1. PSCT 2. RSCT 3. Spinothalamic 4. PCML Question 5 (2 point) • Location of the soma for this 2nd-order afferent 1. Left nucleus gracilis 2. Right nucleus gracilis 3. Left nucleus cuneatus Move “up” & “down” 4. Right nucleus cuneatus 16 Spinal cord: descending pathways Cell types α-motor neurons (αMN) large, supply extrafusal fibers of skeletal m. Tonic & phasic αMN • Tonic αMN innervate slow, oxidative-glycolytic m. fibers depolarized & slowly conducting axons with small spike amplitudes • Phasic αMN innervate squads of fast, oxidative & fast, oxidative-glycolytic m. fibers larger, higher thresholds, rapidly conducting axons with large spike amplitudes from propriospinal sources usually 1st recruits when voluntary movements are initiated γ -motor neurons (γ MN) small, supply intrafusal fibers of neuromuscular spindles Renshaw cells • in med. part of ant. horn • form inhibitory, glycinergic synapses on αMN • negative feedback, or recurrent inhibition Cell columns • Each motor neurons columns supplies a group of m. having similar functions • individual m. are supplied from cell groups (nuclei) within columns 6 somatomotor cell columns (C3-5), Diaphragm lat. limb m. (C5-8, L2-S2), Arm, thigh (C6-8, L3-S3), Forearm, leg (C8, T1, S1-2) intrinsic m. med. trunk m. (T1-L2), Intercostals, abdominals (all segments), Erector spinae DESCENDING MOTOR PATHWAYS • Important pathways descending to spinal cord are 1. corticospinal (pyramidal) (CST) 2. reticulospinal (extrapyramidal) (RST) 3. vestibulospinal 4. tectospinal 5. raphespinal 6. aminergic 7. Autonomic c c c Corticospinal tract (CST) • Main voluntary motor pathway • sources 1. about 50% from primary motor cortex in precentral gyrus 2. supplementary motor area on med. side of hemisphere 3. premotor cortex on lat. side 4. somatic sensory cortex 5. parietal lobe 6. cingulate gyrus (contributions from 2 sensory areas mentioned terminate in sensory nuclei of brainstem & spinal cord modulate sensory transmission) Corticospinal tract • CST pathway corona radiata & internal capsule brainstem crus of midbrain & basilar pons medulla oblongata forms pyramid • Corticonuclear: gives off fibers to activate motor cranial nerve nuclei for face, jaw, and tongue m. Corticospinal tract (CST) Just above spinomedullary junction 1.About 80% of fibers cross midline in pyramidal decussation 2.descend on contralateral side of spinal cord as LCST (lat corticospinal tract) 3.About 10% enter ACST (ant corticospinal tract) occupies ant funiculus at cervical & upper thoracic levels cross in white commissure & supply MN for deep m. in neck 4.About 10% of pyramidal fibers enter LCST on same side • CST contains about 1 million nerve fibers • All corticospinal fibers are excitatory & use glutamate as transmitter Targets of LCST Distal limb motor neurons • A unique property of corticomotoneuronal fibers of LCST is fractionation selectively activate small groups of neurons • most obvious in index finger flex/ext quite independently • essential for skilled movements (buttoning a coat or tying shoe laces) • when lost, seldom recover completely • α & γ MN are coactivated by LCST during a given movement spindles in prime movers are signaling active stretch while those in antagonists are signaling passive stretch Renshaw cells • cocontraction of prime movers & antagonists fix one or more joints • inactivation of Ia inhibitory internucials by Renshaw cells Excitatory internuncials • MN supplying axial & proximal limb m. are recruited indirectly by LCST using excitatory internuncials in intermediate gray matter & ant. horn base Targets of LCST Ia inhibitory internuncials • Also located in intermediate gray matter • first neurons to be activated by LCST during voluntary movements causes antagonist m. to relax before prime movers contract Sequence of voluntary movement (knee flexion) 1) Activation of Ia internuncials to inhibit antagonist αMN 2) activation of agonist α & γ MN 3) activation of extrafusal & intrafusal m. fibers 4) feedback from actively stretched spindles excitation of agonist αMN & antagonist αMN 5) Ia fibers from passively stretched antagonist spindles find respective αMN refractory DESCENDING MOTOR PATHWAYS • Important pathways descending to spinal cord are 1. corticospinal (pyramidal) (CST) 2. reticulospinal (extrapyramidal) (RST) 3. vestibulospinal 4. tectospinal 5. raphespinal 6. aminergic 7. Autonomic c c c Reticulospinal tracts (RST) • originate in reticular formation of pons & medulla oblongata • partially crossed • PRST (pontine reticulospinal tract) descends in ant funiculus acts on extensor MN • MRST (medullary reticulospinal tract ) Descends (partly crossed) in lat funiculus flexor MN • Both tracts act on MN supplying trunk & proximal limb muscles • Both pathways exert reciprocal inhibition • RS system is involved in 2 different kinds of motor behavior: locomotion & postural control Reticulospinal tracts (RST) Locomotion • Walking & running are rhythmic events involving all 4 limbs • 2 side movements are reciprocal to flexor/extensor contractions & relaxations • Locomotion: initiated from locomotor center in lower midbrain for humans (in pons for lab animal) • pattern generators: intermediate gray matter at upper end of spinal cord initiate rhythmic movements • human locomotion is less 'spinal' than quadrupeds • In human, removal of entire cerebral hemisphere during childhood or adolescence bilaterally organized motor system controlling proximal & axial m. exist for return near-perfect locomotor function, but never recover manual skill on contralateral side 2 distinct pathways: pyramidal vs. extrapyramidal (reticulospinal) Posture • position held between movements, e.g. standing, sitting • postural fixation: immobilization of proximal limb joints by cocontraction of surrounding muscles, leaving distal limb parts free to do voluntary business Resident Evil DESCENDING MOTOR PATHWAYS • Important pathways descending to spinal cord are 1. corticospinal (pyramidal) (CST) 2. reticulospinal (extrapyramidal) (RST) 3. vestibulospinal 4. tectospinal 5. raphespinal 6. aminergic 7. Autonomic c c c Vestibulospinal tract • originates in vestibular nucleus in medulla oblongata descends in ant funiculus • tone of antigravity m. is automatically increased as head is tilted • keep center of gravity between feet Tectospinal tract • crossed pathway: from tectum of midbrain med. part of ant gray horn at cervical & upper thoracic levels • access to axial MN • important in reptilian brain responsible for orienting head-trunk toward visual or auditory sources • similar to automatic functions in humans Raphespinal tract • originates in/beside raphe nucleus in medulla oblongata descend on both sides within Lissauer • modulate sensory transmission between 1st & 2nd-order neurons in post gray horn (particular pain) Aminergic pathways • from specialized cell groups in pons & medulla oblongata descend in outer parts of ant & lat funiculi wide distributed in spinal gray matter • neurotransmitters: norepinephrine & serotonin (classed as biogenic amines) • inhibitory effects on sensory neurons & facilitatory effects on motor neurons Central autonomic pathways • Originate part from autonomic control centers in hypothalamus & part from several nuclear groups in brainstem descend beside intermediate gray matter terminate in intermediolateral cell columns to preganglionic sympathetic & parasympathetic fibers of peripheral autonomic system • required for normal baroreceptor reflex activity SCI patient blood pressure in carotid sinus as sitting up compensatory sympathetic activity to maintain blood flow to brain • originate in reticular formation tonic inhibitory on sacral parasympathetic system required for normal bladder/rectal function severe injury to spinal cord or cauda equina results in reflex voiding when is only half full BLOOD SUPPLY OF SPINAL CORD Arteries • Close to foramen magnum, 2 vertebral a. give off ant/post spinal branches • ant branches fuse to form a single ant spinal a. • spinal a. are boosted by several radiculospinal branches from vertebral & intercostal a. • small radicular a. enter every intervertebral foramen to nourish nerve roots • rare vascular disorders in spinal cord • artery of Adamkiewicz : largest radiculospinal a. from lower intercostal or upper lumbar a. supplies lumbar enlargement & conus medullaris be careful when abdominal aortic aneurysm clamp is placed above a. postoperative paraplegia with incontinence Veins • Drainage by ant/post spinal v. outward along nerve roots • obstruction edema of cord progressive loss of function Clinical Panel 16.3 Spinal cord injury automobile accidents are commonest cause of SCI (16 ~30 y/o with cervical injury) • • T & L level results in paraplegia (paralysis of lower limbs) C level causes tetraplegia (quadriplegia) Spinal shock • Below injury level in first few days following a complete cord transection 1. 2. 3. 4. Paralysis movement limbs flaccid & absent tendon reflexes Anesthesia (loss of all forms of sensation) Paralysis of bladder & rectum • hyperpolarization of spinal neurons below lesion level (release inhibitory transmitter glycine) • patient develops postural hypotension interruption of baroreceptor reflex wearing an abdominal binder for compensation Return of spinal function • Several days or weeks later 1. 2. 3. 4. 5. Reflex progressively restored & upper motor neuron signs appear Muscle tone becomes excessive (spastic) Tendon reflexes become abnormally brisk Babinski sign can be elicited on both sides Ankle clonus is commonly seen • bladder condition is important for 2 dangers on infection & formation of bladder stones initial, atonic bladder, insert a sterile catheter to ensure unobstructed drainage later, automatic bladder, emptying itself every 4-6 h through a reflex arc • Considerable interest has been aroused by observations in several spinal rehabilitation centers, patients with complete cord transections can be trained to activate spinal locomotor generators Question 7 (2 point) • Which of the following symptoms can be observed in chronic complete SCI patients B A C Ans: (1) A only; (2) A+B; (3) A+B+C; (4) B+C sensory motor PCML (motor) Spinocerebellar c Spinothalamic (Pain, Touch) c c Thanks for your attention!!