Medical Neurosciences Spinal Cord Spinal Cord DAVID GRIESEMER, MD Professor of Neurosciences and Pediatrics Key Concepts: 1. In adults the spinal cord ends at vertebral level L2, and the nerve roots continue as the cauda equina to exit at the appropriate vertebral level. The level of spinal cord termination allows a lumbar puncture to be performed with greater safety at spinal level L4-5. 2. Cell bodies for sensory neurons carrying information to the spinal cord are located in dorsal root ganglia, which are located outside the spinal canal. 3. Sensory information enters the spinal cord on the dorsal side. 4. The gray matter in the spinal cord contains motor neurons, neurons ascending to the brain, interneurons, and glial cells. 5. Alpha motor neurons from segments C3, C4 and C5 travel via the phrenic nerve to innervate the diaphragm. 6. Alpha motor neurons originate from lamina IX, with those innervating extensor muscles more ventral than those innervating flexor muscles and those innervating distal muscles more lateral than those innervating proximal muscles. 7. Gamma motor neurons, which innervate intrafusal muscle fibers necessary in the muscle stretch reflex arc, also originate from lamina IX. 8. Autonomic fibers originate from the intermediolateral region of the gray matter. In the thoracic and lumbar spinal cord these are preganglionic sympathetic neurons and in the sacral regions they are preganglionic parasympathetic neurons. No autonomic neurons arise from the spinal cord at the cervical level. 9. Axons that ascend in the posterior column of white matter have their cell bodies in the dorsal root ganglia. These neurons mediate vibration, position sense, two-point discrimination, and feeling for shape, pattern, or direction of stimulus on the skin. 10. The dorsal nucleus of Clarke, located in lamina V between T1 and L2, receives unconscious sensory information about leg position. It has a vital role in coordination of walking. 11. Bladder control and micturition require coordination of signaling from somatic innervation and from parasympathetic both derived from S2 – S4 and from sympathetic innervation derived from T11-L2. 12. Supplemental alpha motor neurons from segments S1 – S4 innervate the anal and urethral sphincters, as well as muscles necessary for male sexual function. Medical Neurosciences Spinal Cord INTRODUCTION The spinal cord contains the first synapse in all sensory pathways from the body and back of the head. It also contains all the motor neurons that innervate skeletal muscles of the body. (However, it does not have a role in sensory perception from the face or in motor control of facial muscles.) It also plays a vital role in the control of urination and the bladder. EXTERNAL VIEW The spinal cord is located in the vertebral canal. At birth the cord extends the full length of the spinal column, but in the adult it extends from the foramen magnum to the L1 or L2 vertebral level. Most of the cells in the spinal cord are present at birth, and it does not grow substantially. However, since the bony vertebral column continues to grow for another 20 years, the adult spinal cord ends at the level of the L1 or L2 vertebra. As a result the spinal nerves associated with progressively lowers levels of the cord must travel further downward within the canal to exit at the appropriate intervertebral foramen. This collection of long spinal nerves extending beyond the conus medullaris, or tapering end of the spinal cord, is called the cauda equina, or “tail of the horse.” By way of review, the spinal cord is divided into 31 segments: 8 cervical 12 thoracic 5 lumbar 5 sacral 1 coccygeal Medical Neurosciences Spinal Cord Dorsal and ventral roots exit the spinal cord at each spinal level. Dorsal roots carry sensory information to the spinal cord; their cell bodies are located in the dorsal root ganglia. The ventral roots carry motor and autonomic inform from the spinal cord; their cell bodies are located in the gray matter of the spinal cord. Both dorsal (somatosensory) and ventral (motor) roots fan into tiny rootlets that attach to the cord along a vertical line at the dorsolateral and ventrolateral surfaces of the cord. Before exiting the spinal canal, the dorsal and ventral roots join a short distance to form a spinal nerve. CROSS SECTIONAL VIEW Appearance. The spinal cord consists of gray matter that contains neuronal cell bodies (below, on left) and white matter that consists of myelinated fiber tracts (below, on right). These are both ascending and descending tracts. The gray matter is centrally located in the spinal cord. It is arranged in a “butterfly” of H-shaped pattern that varies depending upon the level of the cord. The white matter surrounds the gray matter and is therefore in the peripheral regions of the cord. The spinal cord is not perfectly symmetric, the right side being slightly larger in most people. This occurs because of more descending fibers originating from the left side of brain, regardless of handedness. The two halves of the spinal cord (where not connected around the central canal) are separated by the ventral (anterior) fissure and the dorsal (posterior) median septum. Medical Neurosciences Spinal Cord In some of the cervical and lumbar regions of the spinal cord there are additional neurons that innervate muscles and extra skin receptors for the arms and legs. The addition of these neurons expands the gray matter at the cervical enlargement (C5 to T2) and at the lumbar enlargement (L2 to S1). Ascending from lower sacral segments to higher cervical segments of the cord, there is an increasing volume of white matter. This is because: at progressively higher levels there is an accumulation of axons bringing sensory input from all parts of the body; at progressively lower levels there is a decrease in the number of descending axons from the brain, as they distribute into the gray matter along the length of the cord; some white matter tracts are not present below certain levels. Organization of Gray Matter. The gray matter of the cord contains primarily the cell bodies of neurons and glia. The neurons are of two basic types: primary neurons and interneurons. Primary neurons are further divided into two broad categories: motor neurons that have axons leaving via the ventral nerve root, and ascending neurons that carry information to the brain. The traditional terminology divides the gray matter into four regions: dorsal horn (or, posterior column) – this region receives axons of dorsal root ganglia. It is concerned with sensory function. intermediolateral horn (part of the lateral column) -- this horn is limited to the thoracic and upper lumbar regions of the cord and is concerned with autonomic nervous system function. It consists of preganglionic sympathetic neurons which exit the spinal cord through the ventral roots. In this equivalent region of the sacral region of the cord the S2, S3 and S4 segments contain preganglionic parasympathetic neurons. ventral horn (or, anterior column) -- this region contains motor neurons, axons from which exit through the ventral nerve root. A modern view of the spinal cord is based upon layers of axon termination, based on cytological criteria. A series of ten layers, labeled I through X, was proposed by Bror Rexed. Details of the traditional and modern classification system, together with a functional overview are outlined in the chart below (for your reference only). TRADITIONAL TERMINOLOGY HORN REGIONS REXED LAMINA Posteromarginal nucleus Substantia gelatinosa I II Dorsal Nucleus proprius III-IV Intermedio lateral Neck of Posterior horn Base of Posterior horn Intermediate zone Commissural nucleus Ventral horn Grisea centralis V VI VII VIII IX X Ventral FUNCTION Exteroceptive sensation Proprioceptive sensation Pain and temperature Position sense, vibration touch, pressure, Position information from the legs (includes dorsal nucleus of Clarke) Relay between midbrain and cerebellum Modulation of motor activity via γ motor neurons Main motor nuclei of both α and γ motor neurons Surrounds the central canal, and contains neuroglia Medical Neurosciences Spinal Cord Looking at a cross section of the cervical cord as an example, the figure on the left highlights cell groups using the traditional nomenclature, while the figure on the right highlights synaptic layers identified by Rexed laminae. Regions of particular note include: 1 Lamina V. Of particular note is a region in this lamina between the T1 and L2 spinal segments. A noticeable “bump” on the mediobasal margin of the dorsal horn in this region corresponds to the dorsal nucleus of Clarke. This nucleus receives sensory data concerning position of the legs and therefore has a role in coordination of walking. Lamina IX. This is the primary motor area of the spinal cord. It contains large α motor neurons that supply extrafusal muscle fibers and smaller γ motor neurons that supply intrafusal muscle fibers. o α motor neurons are arranged by function neurons innervating muscles causing flexion of joints are located more dorsally neurons innervating muscles causing extension of joints are located ventrally neurons innervating distal muscles (e.g. hand) are located more laterally neurons innervating proximal muscles (e.g. trunk) are located medially o α motor neurons from segments C3, C4 and C5 travel via the phrenic nerve. Axons of this nerve innervate the diaphragm and are essential for breathing. o Onuf’s nucleus. Supplemental α motor neurons from segments S1 – S4 innervate the anal and urethral sphincters. They are therefore essential for bowel and bladder continence. In addition these motor neurons supply muscles essential for sexual function in males.1 o Interneurons. In addition to α and γ motor neurons, lamina IX contains interneurons, including the Renshaw cell. This interneuron is part of a “negative feedback” loop for the α motor neuron. It is stimulated by a collateral axon coming from the motor neuron, but its output is an inhibitory signal to the dendrite going to the originating motor neuron and neighboring motor neurons. Renshaw cell inhibition can therefore allow motor neurons to influence their own activity and damp output of selected neurons. Only these supplemental motor neurons are spared in motor neuron diseases like amyotrophic lateral sclerosis (Lou Gehrig’s disease). Medical Neurosciences Spinal Cord Motor neurons that innervate muscles are arranged into vertical columns in the anterior horn of the spinal cord. These columnar collections are considered nuclei, and they are analogous to nuclei for cranial nerves that are located in the brainstem. There is a more complex arrangement in the cervical and lumbar regions. During embryonic development, primitive muscles carry their original innervation with them, generating a motor column that sends its axons through multiple nerve roots from multiple spinal cord levels. Note that, while a muscle may be innervated by axons from multiple spinal nerve roots, those axons arise from a single motor column. Organization of White Matter. The white matter of the spinal cord contains primarily axons gathered into ascending and descending fiber tracts. The H-shaped appearance of gray matter on cross-sectional view allows division of the white matter into three symmetric, paired columns (or funiculi) which extend the length of the spinal cord. Each column contains bundles of axons that channel specific sensory information towards the brain or motor command signals from the brain. These are illustrated above. Medical Neurosciences Spinal Cord dorsal (posterior) column – lies medial to the dorsal horn of gray matter. It contains ascending axons that carry somatic sensory signals to the brain. Sensory axons from inferior regions of the body are located more medially in the posterior column than axons from higher regions. Dorsal nerve roots that enter the spinal cord below the T7 segment are located medially in the posterior column and form the gracile tract. Nerve roots that enter the spinal cord above the T6 segment are located laterally in the posterior column and form the cuneate tract. Lesions in this region manifest as a loss or diminution of the following sensations: o Position sense o Vibration sense o Two point discrimination o Touch o Feel for shapes or patterns o Discerning direction of stimulus on skin lateral column – lies laterally to the gray matter between the dorsal horn and the ventral horn. It contains ascending somatic sensory axons and descending motor control axons. ventral (anterior) column – lies medial to the ventral horn of gray matter. descending motor control axons. It contains TRACTS OF THE SPINAL CORD While the posterior column contains only one ascending tract which conveys conscious perception to the cerebral cortex, the lateral and ventral columns contain both ascending and descending tracts. Ascending tracts. All of the clinically significant ascending tracts have cell bodies of origin in the dorsal root ganglia. Not all, however, convey information of which a person is conscious. There are four main ascending pathways: o dorsal (posterior) spinocerebellar tract – conveys to the cerebellum information about strength, rate and phase of muscle contraction o ventral (anterior) spinocerebellar tract – conveys information about the effect of descending signals and interneuronal activity. Dorsal and ventral spinocerebellar tracts together provide unconscious information about position. o lateral spinothalamic tract – conveys pain and temperature information o anterior spinothalamic tract – conveys sensation of light touch and some pain. Functionally the anterior and lateral spinothalamic tracts are one. Medical Neurosciences Spinal Cord Descending tracts. While neurons of all ascending tracts are located in dorsal root ganglia, axons in descending tracts come from neurons in several locations. Key pathways include: o corticospinal tract – conveys information essential for speed and agility of movement. It does not by itself initiate movement. o rubrospinal tract – conveys information for correcting errors in movement. o lateral vestibulospinal tract – conveys information to activate extensor motor neurons that maintain upright posture. o medial vestibulospinal tract – conveys information to control head position and activate flexor motor neurons. o reticulospinal tract – axons arising from the pons facilitate extensor motor neurons, and axons arising from the medulla facilitate flexor motor neurons o tectospinal tract – facilitates turning of the head in response to light Medical Neurosciences Spinal Cord CLINICAL APPLICATION: MICTURITION AND BLADDER CONTROL From a clinical perspective, one of the most important functions of the spinal cord it to mediate control of the bladder and the function of urination. The bladder had three sources of efferent and afferent signal transmission: somatic innervation from Onuf’s nucleus at S2, S3 and S4 via the pudendal nerve sympathetic innervation from the intermediolateral cell column at T11 – L2 via the hypogastric nerve parasympathetic innervation from the intermediolateral-like cell column at S2, S3 and S4 via the pelvic nerve Bladder filling is dependent upon tonic activity of somatic neurons and sympathetic neurons. Somatic stimulation causes contraction of the external urethral sphincter Sympathetic stimulation causes contraction of the internal urethral sphincter and relaxation of the detrusor muscle that expels urine from the bladder Bladder emptying is dependent upon all three innervations: Inhibition of somatic innervation to relax the external urethral sphincter Inhibition of sympathetic stimulation to relax the internal urethral sphincter Parasympathetic stimulation to stimulate detrusor contraction Central nervous system control is also an essential element: Descending pathways originating in the pons coordinate starting and stopping of micturition Afferent information about bladder filling travels to the periaqueductal region of the midbrain. At the point when it is filled enough to empty, the midbrain activates neurons in the micturition center of the pons The medial preoptic area of the hypothalamus is involved in micturition The right inferior frontal gyrus and the right anterior cingulate gyrus of the cerebral cortex are also involved in micturition.