Difference between revisions of "Spinal cord" - New World Encyclopedia

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{{Infobox anatomy
[[Image:Spinal cord direv.svg|thumb|200px|right|The Spinal cord nested in the [[spine (anatomy)|vertebral column]].]]
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| Name        = {{PAGENAME}}
[[Image:Spinalcord trirev.svg|thumb|200px|right|A closer look.]]
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| Latin      = medulla spinalis
[[Image:Medulla spinalis - Section - English.svg|thumb|200px|right||Cross-section through cervical spinal cord.]]
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| GraySubject =
[[Image:Spinalcord trirev rexedlamina.svg|thumb|200px|right|Gray Matter's Rexed Lamina.]]
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| GrayPage    =
[[Image:somatosensory.jpg|thumb|200px|right|Somatosensory Tracts.]]
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| Image      = Nervous system diagram.png
[[Image:alarplate.jpg|thumb|200px|right|Spinal Cord Development of the Alar and Basal Plates]]
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| Caption    = The spinal cord (in red) connects the brain to the nerves throughout the body
[[Image:Medulla spinalis - tracts - English.svg|thumb|200px|right|Spinal Cord Tracts]]
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| Width      = 240
 
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| Image2      =
The '''spinal cord''' is a thin, tubular bundle of [[nerve]]s that is an extension of the [[central nervous system]] from the brain and is enclosed in and protected by the bony [[spine (anatomy)|vertebral column]]. The main function of the spinal cord is transmission of neural inputs between the [[periphery]] and the [[brain]].  
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{{Vertebral column}}
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The '''spinal cord''' is a long, thin, tubular bundle of [[nervous tissue]] and [[glia|support cells]] that extends from the [[brain]] (the [[medulla oblongata]] specifically).  The brain and spinal cord together make up the [[central nervous system]] (CNS). The spinal cord begins at the [[occipital bone]] and extends down to the space between the first and second [[lumbar vertebrae]]; it does not extend the entire length of the [[vertebral column]]. It is around {{convert|45|cm|in|abbr=on}} in men and around {{convert|43|cm|in|abbr=on}} long in women.  Also, the spinal cord has a varying width, ranging from 1/2 inch thick in the cervical and lumbar regions to 1/4 inch thick in the thoracic area. The enclosing bony [[spine (anatomy)|vertebral column]] protects the relatively shorter spinal cord. The spinal cord functions primarily in the transmission of neural signals between the [[brain]] and the rest of the body but also contains [[neural circuit]]s that can independently control numerous [[reflex]]es and [[central pattern generator]]s.
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The spinal cord has three major functions:
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as a conduit for motor information, which travels down the spinal cord, as a conduit for sensory information in the reverse direction, and finally as a center for coordinating certain reflexes.
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<ref>{{cite book
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  | last = Maton
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  | first = Anthea
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  | authorlink =
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  | coauthors = Jean Hopkins, Charles William McLaughlin, Susan Johnson, Maryanna Quon Warner, David LaHart, Jill D. Wright
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  | title = Human Biology and Health
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  | publisher = Prentice Hall
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  | year = 1993
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  | location = Englewood Cliffs, New Jersey, USA
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  | pages = 132–144
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  | url =
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  | doi =
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  | id =
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  | isbn = 0-13-981176-1}}</ref>
  
 
==Structure==
 
==Structure==
The human spinal cord extends from the [[medulla oblongata]] in the brain and continues to the [[conus medullaris]] near the lumbar level at L1-2, terminating in a fibrous extension known as the [[filum terminale]].
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The spinal cord is the main pathway for information connecting the brain and peripheral nervous system. The length of the spinal cord is much shorter than the length of the bony spinal column. The human spinal cord extends from the [[foramen magnum]] and continues through to the [[conus medullaris]] near the second [[lumbar vertebra]], terminating in a fibrous extension known as the [[filum terminale]].
  
It is about 45 cm long in men and 42 cm long in women, ovoid-shaped, and is enlarged in the cervical and lumbar regions. The peripheral regions of the cord contains neuronal white matter tracts containing sensory and motor neurons. The central region is a four-leaf clover shape that surrounds the central canal (an anatomic extension of the [[fourth ventricle]]) and contains nerve cell bodies.  
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It is about {{convert|45|cm|in|abbr=on}} long in men and around {{convert|43|cm|in|abbr=on}} in women, ovoid-shaped, and is enlarged in the cervical and lumbar regions. The cervical enlargement, located from C3 to T2 spinal segments, is where sensory input comes from and motor output goes to the arms. The lumbar enlargement, located between L1 and S3 spinal segments, handles sensory input and motor output coming from and going to the legs.
  
The three [[meninges]] that cover the spinal cord—the outer [[dura mater]], the [[arachnoid membrane]], and the innermost [[pia mater]]—are continuous with that in the brainstem and cerebral hemispheres, with [[cerebrospinal fluid]] found in the [[subarachnoid space]]. The cord within the pia mater is stabilized within the dura mater by the connecting [[denticulate ligaments]] which extends from the pia mater laterally between the dorsal and ventral roots. The dural sac ends at the vertebral level of S2.
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The spinal cord is protected by three layers of tissue, called spinal [[meninges]], that surround the canal. The [[dura mater]] is the outermost layer, and it forms a tough protective coating. Between the dura mater and the surrounding bone of the vertebrae is a space called the [[epidural space]]. The epidural space is filled with [[adipose tissue]], and it contains a network of [[blood vessel]]s. The [[arachnoid mater]] is the middle protective layer. Its name comes from the fact that the tissue has a spiderweb-like appearance. The space between the arachnoid and the underlying [[pia mater]] is called the [[subarachnoid space]]. The subarachnoid space contains [[cerebrospinal fluid]] (CSF). The medical procedure known as a [[lumbar puncture]] (or "spinal tap") involves use of a needle to withdraw cerebrospinal fluid from the subarachnoid space, usually from the [[lumbar]] region of the spine. The pia mater is the innermost protective layer. It is very delicate and it is tightly associated with the surface of the spinal cord. The cord is stabilized within the dura mater by the connecting [[denticulate ligaments]], which extend from the enveloping pia mater laterally between the dorsal and ventral roots. The '''dural sac''' ends at the vertebral level of the second [[sacrum|sacral]] vertebra.
  
== Sensory Organization ==
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In cross-section, the peripheral region of the cord contains neuronal [[white matter]] tracts containing [[sensory neuron|sensory]] and [[motor neuron]]s. Internal to this peripheral region is the gray, butterfly-shaped central region made up of [[neuron|nerve cell bodies]]. This central region surrounds the [[central canal]], which is an anatomic extension of the spaces in the brain known as the [[Ventricular system|ventricles]] and, like the ventricles, contains cerebrospinal fluid.  
Somatosensory organization is divided into a touch/[[proprioception]]/vibration sensory pathway and a pain/temperature sensory pathway, which are more formally known as the [[dorsal column-medial lemniscus tract]] and the [[spinothalamic tract]], respectively.
 
 
Each of these sensory pathways utilizes three different neurons to get from the sensory receptors to the [[cerebral cortex]]. These neurons are designated primary, secondary and tertiary sensory neurons. The primary neuron has its cell body in the [[dorsal root ganglia]] and  its [[axon]] projects into the spinal cord.
 
  
In the case of the touch/proprioception/vibration sensory pathway, the primary neuron enters the spinal cord and travels in the [[dorsal column]]. Below level T6, the neuron  travels in the [[fasciculus gracilis]] - the most medial part of the column. Above level T6, the neuron enters the [[fasciculus cuneatus]] - lateral to the fasiculus gracilis.
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The spinal cord has a shape that is compressed dorso-ventrally, giving it an [[elliptical]] shape. The cord has grooves in the dorsal and ventral sides. The [[posterior median sulcus of spinal cord|posterior median sulcus]] is the groove in the dorsal side, and the [[anterior median fissure of spinal cord|anterior median fissure]] is the groove in the ventral side.
  
As the primary axons reach the caudal [[medulla]], they leave their respective fasiculi and  enter and synapse on secondary neurons within the [[nucleus gracilis]] and the [[nucleus cuneatus]], respectively. At this point, the seconday neuronal axons  decussate and  continue to ascend as the [[medial leminiscus]]. They run up to the [[Ventral posterolateral nucleus|VPL]] nucleus of the [[thalamus]],and  synapse there on the tertiary neurons. From there, the tertiary neurons  ascend via the posterior limb of the [[internal capsule]] to the post central gyrus, or [[Brodmann]]'s Area 3,1,2.   
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===Spinal cord segments===
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[[File:Spinal readjustment 3.jpg|thumb|right|alt=Model of a section of a spine.|A model of segments of the human vertebral column, nerve roots can be seen extending laterally from the (not visible) spinal cord.]]
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The human spinal cord is divided into 31 different segments. At every segment, right and left pairs of spinal nerves (mixed; sensory and motor) form. Six to eight motor nerve rootlets branch out of right and left ventro lateral sulci in a very orderly manner. Nerve rootlets combine to form nerve roots. Likewise, sensory nerve rootlets form off right and left dorsal lateral sulci and form sensory nerve roots. The ventral (motor) and dorsal (sensory) roots combine to form [[spinal nerve]]s (mixed; motor and sensory), one on each side of the spinal cord. Spinal nerves, with the exception of C1 and C2, form inside [[intervertebral foramen]] (IVF).  Note that at each spinal segment, the border between the central and peripheral nervous system can be observedRootlets are a part of the peripheral nervous system.
  
The pain/temperature sensory pathway differs from that of the touch/proprioception/vibration pathway. The pain neurons enter as primary neurons and  ascend 1-2 levels before synapsing in the [[substantia gelatinosa]]. The tract that ascends those 1-2 levels before synapsing is known as [[Lissauer's tract]]. After synapsing, the secondary neurons  cross decussate and ascend as the [[spinothalamic tract]] in the anterior lateral portion of the spinal cord. Hence, the spinothalamic tract is also known as the [[anterior lateral system]] (ALS). The tract ascends all the way to the VPL of the thalamus where it  synapses on the tertiary neurons. The tertiary neuronal axons then project via the posterior limb of the internal capsule to the post-central gyrus or Broadmann's Area 3,1,2.  
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In the upper part of the vertebral column, spinal nerves exit directly from the spinal cord, whereas in the lower part of the vertebral column nerves pass further down the column before exiting. The terminal portion of the spinal cord is called the [[conus medullaris]]. The [[pia mater]] continues as an extension called the [[filum terminale]], which anchors the spinal cord to the [[coccyx]]. The [[cauda equina]] (“horse’s tail”) is the name for the collection of nerves in the vertebral column that continue to travel through the vertebral column below the [[conus medullaris]]. The cauda equina forms as a result of the fact that the spinal cord stops growing in length at about age four, even though the vertebral column continues to lengthen until adulthood. This results in the fact that sacral spinal nerves actually originate in the upper lumbar region. The spinal cord can be anatomically divided into 31 spinal segments based on the origins of the spinal nerves.
  
It should be noted that the pain fibers in the ALS can also deviate in their pathway towards the VPL. In one pathway, the axons  project towards the [[reticular formation]] in the midbrain. The reticular formation then project to a number of places including the [[hippocampus]] (to create memories about the pain), to the [[centromedian nucleus]] (to cause diffuse, non-specific pain) and the various places on the cortex. The third place that the neurons project to is the [[periaqueductal gray]] in the pons. The neurons form the periaqueductal gray  then project to the [[nucleus raphe magnus]] which  projects back down to where the pain signal is coming in from and inhibits it. This  reduces the pain sensation to some degree.
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Each segment of the spinal cord is associated with a pair of ganglia, called [[dorsal root ganglia]], which are situated just outside of the spinal cord. These ganglia contain cell bodies of sensory neurons. Axons of these sensory neurons travel into the spinal cord via the dorsal roots.
  
== Motor Organization ==
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Ventral roots consist of axons from motor neurons, which bring information to the periphery from cell
Upper motor neuronal input comes from two places- first  from the cerebral cortex and second  from more primitive brainstem nuclei. Cortical upper motor neurons originate in Brodmann Areas 4, 6, 3, 1 and 2. They then descend through the genu and the posterior limb of the [[internal capsule]]. This pathway is known as the [[corticospinal tract]]. After passing through the internal capsule, the tract descends through the cerebral peduncles, down through the pons and to the [[medullary pyramids]]. At this point, ~85% of these upper motor neuronal axons decussate. These fibers then descend as the lateral corticospinal tract. The remaining ~15% descend as the anterior corticospinal tract. <br /><br />
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bodies within the CNS. Dorsal roots and ventral roots come together and exit the intervertebral
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foramina as they become spinal nerves.
  
The midbrain nuclei include four motor tracts that send upper motor neuronal axons down the spinal cord to lower motor neurons. These are the [[rubrospinal tract]], the [[vestibulospinal tract]], the [[tectospinal tract]] and the [[reticulospinal tract]]. The rubrospinal tract descends with the lateral corticospinal tract and the remaining three descend with the anterior corticospinal tract.<br /><br />
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The gray matter, in the center of the cord, is shaped like a butterfly and consists of cell bodies of
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[[interneurons]] and motor neurons. It also consists of [[neuroglia]] cells and [[unmyelinated]] axons.
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Projections of the gray matter (the “wings”) are called horns. Together, the gray horns and the gray
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[[commissure]] form the “gray H.
  
The function of lower motor neurons can be divided into two different groups—first,the lateral corticospinal tract and second, the anterior cortical spinal tract. The lateral tract contains upper motor neuronal [[axons]] which synapse on dorsal lateral (DL) lower motor neurons. The DL neurons are involved in [[distal]] limb control. Therefore, these DL neurons are found specifically only in the cervical and lumbosaccral enlargements within the spinal cord. There is no decussation in the lateral corticospinal tract after the decussation at the medullary pyramids.<br />  <br />
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The white matter is located outside of the gray matter and consists almost totally of [[myelinated]] motor
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and sensory axons. “Columns” of white matter carry information either up or down the
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spinal cord.
  
The anterior corticospinal tract descends [[ipsilateral]]ly in the anterior column where the axons  emerge and either synapse on lower motor neurons, known as ventromedial (VM) lower motor neurons, in the ventral horn ipsilaterally, or descussate at the [[anterior white commissure]] where they synapse on VM lower motor neurons [[contralateral]]ly . The tectospinal, vestibulospinal and reticulospinal descend ipsilaterally in the anterior column, but do not synapse across the anterior white commissure. Rather, they only synapse on VM lower motor neurons ipsilaterally. The VM lower motor neurons control [[axial]] motor function—the large, postural muscles. These lower motor neurons, unlike those of the DL, are located in the ventral horn all the way throughout the spinal cord.
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Within the CNS, nerve cell bodies are generally organized into functional clusters, called nuclei. Axons
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within the CNS are grouped into tracts.
  
== Spinocerebellar Tracts ==
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There are 33 spinal cord nerve segments in a human spinal cord:
[[Proprioceptive]]information in the body travels up the spinal cord via three tracts. Below L2 the proprioceptive information travels up the spinal cord in the [[ventral spinocerebellar tract]]. Also known as the anterior spinocerebellar tract, sensory receptors take in the information and travel into the spinal cord. The cell bodies of these primary neurons are located in the [[dorsal root ganglia]]. In the spinal cord, the axons synapse and the secondary neuronal axons decussate and then travel up to the [[superior cerebellar peduncle]] where they decussate again. From here, the information is brought to deep nuclei of the cerebellum including the [[fastigial]] and [[interposed]] nuclei.  <br />From the levels of L2 to T1, the proprioceptive information enters the spinal cord and ascends ipsilaterally where it synapses in the [[Dorsal Nucleus of Clark]]. The secondary neuronal axons continue to ascend ispilaterally and enter the pass into the cerebellum via the [[inferior cerebellar peduncle]]. This tract is known as the [[dorsal spinocerebellar tract]] and also as the [[posterior spinocerebellar tract]].<br />  From above T1, proprioceptive primary axons enter the spinal cord and ascend ipsilaterally until reaching the [[accessory cuneate nucleus]], where they synapse. The secondary axons pass into the cerebellum via the inferior cerebellar peduncle where again, these axons synapse on cerebellar deep nuclei. This tract is known as the [[cuneocerebellar tract]].
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* 8 cervical segments forming 8 pairs of [[cervical nerves]] (C1 spinal nerves exit spinal column between occiput and C1 vertebra; C2 nerves exit between posterior arch of C1 vertebra and lamina of C2 vertebra; C3-C8 spinal nerves through IVF above corresponding cervica vertebra, with the exception of C8 pair which exit via IVF between C7 and T1 vertebra)
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* 12 thoracic segments forming 12 pairs of [[thoracic nerves]] (exit spinal column through IVF below corresponding vertebra T1-T12)
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* 5 lumbar segments forming 5 pairs of [[lumbar nerves]] (exit spinal column through IVF, below corresponding vertebra L1-L5)
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* 5 sacral segments forming 5 pairs of [[sacral nerves]] (exit spinal column through IVF, below corresponding vertebra S1-S5)
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* 3 coccygeal segments joined up becoming a single segment forming 1 pair of [[coccygeal nerves]] (exit spinal column through the sacral hiatus).
  
===Spinal cord segments=== <!--other articles link here—>
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In the fetus, vertebral segments correspond with spinal cord segments.  However, because the [[vertebra|vertebral column]] grows longer than the spinal cord, spinal cord segments do not correspond to vertebral segments in the adult, particularly in the lower spinal cord. For example, lumbar and sacral spinal cord segments are found between vertebral levels T9 and L2, and the spinal cord ends around the L1/L2 vertebral level, forming a structure known as the conus medullaris.
  
The human spinal cord is divided into 31 different segments, with motor nerve roots exiting in the [[ventral]] aspects and sensory nerve roots entering in the [[dorsal]] aspects. The ventral and dorsal roots later join to form paired [[spinal nerve]]s, one on each side of the spinal cord.  
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Although the spinal cord cell bodies end around the L1/L2 vertebral level, the spinal nerves for each segment exit at the level of the corresponding vertebra. For the nerves of the lower spinal cord, this means that they exit the vertebral column much lower (more caudally) than their roots. As these nerves travel from their respective roots to their point of exit from the vertebral column, the nerves of the lower spinal segments form a bundle called the cauda equina.
  
There are 31 spinal cord nerve segments in a human spinal cord:
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[[File:Dissection of spinal cord.jpg|thumb|right|300px|Spinal cord]]
* 8 cervical segments ([[cervical nerves]] exit spinal column above C1 and below C1-C7)
 
* 12 thoracic segments (nerves exit spinal column below T1-T12)
 
* 5 lumbar segments (nerves exit spinal column below L1-L5)
 
* 5 sacral segments (nerves exit spinal column below S1-S5)
 
* 1 coccygeal segment (nerves exit spinal column at coccyx)
 
 
 
Because the vertebral column grows longer than the spinal cord, spinal cord segments become higher than the corresponding [[vertebra]], especially in the lower spinal cord segments in adults. In a fetus, the vertebral levels originally correspond with the spinal cord segments. In the adult, the cord ends around the L1/L2 vertebral level at the [[conus medullaris]], with all of the spinal cord segments located superiorly to this. For example, the segments for the lumbar and sacral regions are found between the vertebral levels of T9 and L2. The S4 spinal nerve roots arise from the cord around the upper lumbar/lower thoracic vertebral region, and descend downward in the vertebral canal. After they pass the end of the spinal cord, they are considered to be part of the [[cauda equina]]. The roots for S4 finally leave the vertebral canal in the [[sacrum]].
 
  
 
There are two regions where the spinal cord enlarges:
 
There are two regions where the spinal cord enlarges:
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* [[Cervical enlargement]] - corresponds roughly to the [[brachial plexus]] nerves, which innervate the [[upper limb]]. It includes spinal cord segments from about C4 to T1. The vertebral levels of the enlargement are roughly the same (C4 to T1).
 
* [[Cervical enlargement]] - corresponds roughly to the [[brachial plexus]] nerves, which innervate the [[upper limb]]. It includes spinal cord segments from about C4 to T1. The vertebral levels of the enlargement are roughly the same (C4 to T1).
  
* [[Lumbosacral enlargement]] - corresponds to the [[lumbosacral plexus]] nerves, which innervate the [[lower limb]]. It comprises the spinal cord segments from L2 to S3, and is found about the vertebral levels of T9 to T12.
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* [[Lumbosacral enlargement]] - corresponds to the [[lumbosacral plexus]] nerves, which innervate the [[lower limb]]. It comprises the spinal cord segments from L2 to S3 and is found about the vertebral levels of T9 to T12.
  
 
===Embryology===
 
===Embryology===
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The spinal cord is made from part of the [[neural tube]] during development. As the neural tube begins to develop, the [[notochord]] begins to secrete a factor known as [[Sonic hedgehog]] or SHH. As a result, the [[floor plate]] then also begins to secrete SHH, and this will induce the basal plate to develop [[motor neurons]]. Meanwhile, the overlying [[ectoderm]] secretes [[bone morphogenetic protein]] (BMP). This induces the [[roof plate]] to begin to secrete BMP, which will induce the [[alar plate]] to develop [[sensory neurons]]. The alar plate and the basal plate are separated by the [[sulcus limitans]]. 
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[[File:Human embryo 8 weeks 6.JPG|thumb|300px|Medulla spinalis of 8-week-old human embryo]]
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Additionally, the floor plate also secretes [[netrin]]s. The netrins act as chemoattractants to [[decussation]] of pain and temperature sensory neurons in the alar plate across the anterior white commissure, where they then ascend towards the [[thalamus]].
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Lastly, it is important to note that the past studies of Viktor Hamburger and Rita Levi-Montalcini in the chick embryo have been further proven by more recent studies which demonstrated that the elimination of neuronal cells by [[programmed cell death]] (PCD) is necessary for the correct assembly of the nervous system.
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Overall, spontaneous embryonic activity has been shown to play a role in neuron and muscle development but is probably not involved in the initial formation of connections between spinal neurons.
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== Blood supply ==
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The spinal cord is supplied with blood by three arteries that run along its length starting in the brain, and many arteries that approach it through the sides of the spinal column. The three longitudinal arteries are called the [[anterior spinal artery]], and the right and left [[posterior spinal artery|posterior spinal arteries]].<ref name=Moore298>{{cite book|last=Moore|first=Keith|title=Essential Clinical Anatomy, Third Edition|year=2007|publisher=Lippincott Williams & Wilkins|isbn=0-7817-6274-X|pages=298|coauthors=Anne Agur}}</ref> These travel in the subarachnoid space and send branches into the spinal cord. They form anastamoses (connections) via the anterior and posterior [[segmental medullary artery|segmental medullary arteries]], which enter the spinal cord at various points along its length.<ref name=Moore298 /> The actual blood flow caudally through these arteries, derived from the posterior cerebral circulation, is inadequate to maintain the spinal cord beyond the cervical segments.
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The major contribution to the arterial blood supply of the spinal cord below the cervical region comes from the radially arranged posterior and anterior [[radicular artery|radicular arteries]], which run into the spinal cord alongside the dorsal and ventral nerve roots, but with one exception do not connect directly with any of the three longitudinal arteries.<ref name=Moore298 />  These intercostal and lumbar radicular arteries arise from the aorta, provide major anastomoses and supplement the blood flow to the spinal cord. In humans the largest of the anterior radicular arteries is known as the artery of Adamkiewicz, or anterior radicularis magna (ARM) artery, which usually arises between L1 and L2, but can arise anywhere from T9 to L5.  <ref>{{cite journal|last=Biglioli|first=Paolo|coauthors=et alia|title=Upper and lower spinal cord blood supply: the continuity of the anterior spinal artery and the relevance of the lumbar arteries|journal=Journal of Thoracic and Cardiovascular Surgery|year=2004|month=April|volume=127|issue=4|pages=1188–1192|doi=10.1016/j.jtcvs.2003.11.038|pmid=15052221}}</ref> Impaired blood flow through these critical radicular arteries, especially during surgical procedures that involve abrupt disruption of blood flow through the aorta for example during aortic aneursym repair, can result in spinal cord infarction and paraplegia.
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== Somatosensory organization ==
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{{anchor|tracts}}
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[[Image:Spinal cord tracts - English.svg|thumb|360px|right|Spinal cord tracts.]]
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Somatosensory organization is divided into the [[dorsal column-medial lemniscus tract]] (the touch/[[proprioception]]/vibration sensory pathway) and the [[anterolateral system]], or ALS (the pain/temperature sensory pathway). Both sensory pathways use three different neurons to get information from sensory receptors at the periphery to the [[cerebral cortex]]. These neurons are designated primary, secondary and tertiary sensory neurons. In both pathways, primary sensory neuron cell bodies are found in the [[dorsal root ganglia]], and their central [[axons]] project into the spinal cord.
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In the dorsal column-medial leminiscus tract, a primary neuron's axon enters the spinal cord and then enters the dorsal column. If the primary axon enters below spinal level T6, the axon travels in the [[fasciculus gracilis]], the medial part of the column. If the axon enters above level T6, then it travels in the [[fasciculus cuneatus]], which is lateral to the fasiculus gracilis. Either way, the primary axon ascends to the lower [[medulla oblongata|medulla]], where it leaves its fasiculus and synapses with a secondary neuron in one of the dorsal column nuclei: either the [[nucleus gracilis]] or the [[nucleus cuneatus]], depending on the pathway it took. At this point, the secondary axon leaves its nucleus and passes anteriorly and medially. The collection of secondary axons that do this are known as [[internal arcuate fibers]]. The internal arcuate fibers [[decussate]] and continue ascending as the contralateral [[medial lemniscus]]. Secondary axons from the medial lemniscus finally terminate in the [[ventral posterolateral nucleus]] (VPL) of the [[thalamus]], where they synapse with tertiary neurons. From there, tertiary neurons ascend via the posterior limb of the [[internal capsule]] and end in the [[primary sensory cortex]].
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The proprioception of the lower limbs differs from the upper limbs and upper trunk. There is a four-neuron pathway for lower limb proprioception. This pathway initially follows the dorsal spino-cerebellar pathway. It is arranged as follows: proprioceptive receptors of lower limb -> peripheral process -> dorsal root ganglion -> central process -> [[Clarke's column]] -> 2nd order neuron -> medulla oblogata ([[Caudate nucleus]]) -> 3rd order neuron -> VPL of thalamus -> 4th order neuron -> posterior limb of internal capsule -> corona radiata -> sensory area of cerebrum.
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The anterolateral system works somewhat differently. Its primary neurons axons enter the spinal cord and then ascend one to two levels before synapsing in the [[substantia gelatinosa of Rolando|substantia gelatinosa]]. The tract that ascends before synapsing is known as [[Lissauer's tract]]. After synapsing, secondary axons decussate and ascend in the anterior lateral portion of the spinal cord as the [[spinothalamic tract]]. This tract ascends all the way to the VPL, where it synapses on tertiary neurons. Tertiary neuronal axons then travel to the primary sensory cortex via the posterior limb of the internal capsule.
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It should be noted that some of the "pain fibers" in the ALS deviate from their pathway towards the VPL. In one such deviation, axons travel towards the [[reticular formation]] in the midbrain. The reticular formation then projects to a number of places including the [[hippocampus]] (to create memories about the pain), the [[centromedian nucleus]] (to cause diffuse, non-specific pain) and various parts of the cortex. Additionally, some ALS axons project to the [[periaqueductal gray]] in the pons, and the axons forming the periaqueductal gray then project to the [[nucleus raphes magnus]], which projects back down to where the pain signal is coming from and inhibits it. This helps control the sensation of pain to some degree.
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== Motor organization ==
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The [[corticospinal tract]] serves as the motor pathway for upper motor neuronal signals coming from the cerebral cortex and from primitive brainstem motor nuclei.
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Cortical upper motor neurons originate from [[Brodmann area]]s 1, 2, 3, 4, and 6 and then descend in the posterior limb of the [[internal capsule]], through the [[crus cerebri]], down through the pons, and to the [[medullary pyramids]], where about 90% of the axons cross to the contralateral side at the decussation of the pyramids. They then descend as the lateral corticospinal tract. These axons synapse with lower motor neurons in the ventral [[spinal cord horn|horns]] of all levels of the spinal cord. The remaining 10% of axons descend on the ipsilateral side as the ventral corticospinal tract. These axons also synapse with lower motor neurons in the ventral horns. Most of them will cross to the contralateral side of the cord (via the [[anterior white commissure]]) right before synapsing.
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The midbrain nuclei include four motor tracts that send upper motor neuronal axons down the spinal cord to lower motor neurons. These are the [[rubrospinal tract]], the [[vestibulospinal tract]], the [[tectospinal tract]] and the [[reticulospinal tract]]. The rubrospinal tract descends with the lateral corticospinal tract, and the remaining three descend with the anterior corticospinal tract.
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The function of lower motor neurons can be divided into two different groups: the lateral corticospinal tract and the anterior cortical spinal tract. The lateral tract contains upper motor neuronal [[axons]] which synapse on dorsal lateral (DL) lower motor neurons. The DL neurons are involved in [[distal]] limb control. Therefore, these DL neurons are found specifically only in the cervical and lumbosacral enlargements within the spinal cord. There is no decussation in the lateral corticospinal tract after the decussation at the medullary pyramids.
  
The spinal cord is made from part of the [[neural tube]] during development. As the neural tube begins to develop, the notochord begins to secrete a factor known as [[Sonic hedgehog]] or SHH. As a result, the [[floor plate]] then also begins to secrete SHH and this will induce the basal plate to develop [[motor neurons]]. Meanwhile, the overlying [[ectoderm]] secretes [[bone morphogenetic protein]] (BMP). This will induce the [[roof plate]] to begin to also secrete BMP which will induce the [[alar plate]] to develop [[sensory neurons]]. The alar plate and the basal plate are separated by the [[sulcus limitans]].
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The anterior corticospinal tract descends [[ipsilateral]]ly in the anterior column, where the axons emerge and either synapse on lower ventromedial (VM) motor neurons in the ventral horn ipsilaterally or descussate at the [[anterior white commissure]] where they synapse on VM lower motor neurons [[contralateral]]ly . The tectospinal, vestibulospinal and reticulospinal descend ipsilaterally in the anterior column but do not synapse across the anterior white commissure. Rather, they only synapse on VM lower motor neurons ipsilaterally. The VM lower motor neurons control the large, postural muscles of the [[axial skeleton]]. These lower motor neurons, unlike those of the DL, are located in the ventral horn all the way throughout the spinal cord.
  
Additionally, the floor plate will also secrete [[netrin]]s. The netrins act as chemoattractants to [[decussation]] of pain and temperature sensory neurons in the alar plate across the anterior white commissure where they will then ascend towards the [[thalamus]].
+
== Spinocerebellar tracts ==
 +
[[Proprioceptive]] information in the body travels up the spinal cord via three tracts. Below L2, the proprioceptive information travels up the spinal cord in the [[ventral spinocerebellar tract]]. Also known as the anterior spinocerebellar tract, sensory receptors take in the information and travel into the spinal cord. The cell bodies of these primary neurons are located in the [[dorsal root ganglia]]. In the spinal cord, the axons synapse and the secondary neuronal axons decussates and then travel up to the [[superior cerebellar peduncle]] where they decussate again. From here, the information is brought to deep nuclei of the cerebellum including the [[fastigial]] and [[Interposed nucleus|interposed nuclei]].
  
Lastly it is important to note that the past studies of Viktor Hamburger and Rita Levi-Montalcini in the chick embryo have been further proven by more recent studies which demonstrated that the elimination of neuronal cells by programmed cell death (PCD) is necessary for the correct assembly of the nervous system.
+
From the levels of L2 to T1, proprioceptive information enters the spinal cord and ascends ipsilaterally, where it synapses in [[Clarke's nucleus]]. The secondary neuronal axons continue to ascend ipsilaterally and then pass into the cerebellum via the [[inferior cerebellar peduncle]]. This tract is known as the dorsal spinocerebellar tract.
  
Overall, spontaneous embryonic activity has been shown to play a role in neuron and muscle development, but is probably not involved in the initial formation of connections between spinal neurons.
+
From above T1, proprioceptive primary axons enter the spinal cord and ascend ipsilaterally until reaching the [[accessory cuneate nucleus]], where they synapse. The secondary axons pass into the cerebellum via the inferior cerebellar peduncle where again, these axons synapse on cerebellar deep nuclei. This tract is known as the [[cuneocerebellar tract]].
 +
 
 +
Motor information travels from the brain down the spinal cord via descending spinal cord tracts.  Descending tracts involve two neurons: the upper motor neuron (UMN) and lower motor neuron (LMN).<ref name="SaladinAnatomy">Saladin. Anatomy and Physiology, 5th Ed.</ref>  A nerve signal travels down the upper motor neuron until it synapses with the lower motor neuron in the spinal cord.  Then, the lower motor neuron conducts the nerve signal to the spinal root where efferent nerve fibers carry the motor signal toward the target muscle.  The descending tracts are composed of white matter.  There are several descending tracts serving different functions.  The corticospinal tracts (lateral and anterior) are responsible for coordinated limb movements.<ref name="SaladinAnatomy" />
  
 
==Injury==
 
==Injury==
 +
{{main|Spinal cord injuries}}
  
{{main article|Spinal cord injury}}
+
Spinal cord injuries can be caused by trauma to the spinal column (stretching, bruising, applying pressure, severing, laceration, etc.).  The vertebral bones or [[intervertebral disk]]s can shatter, causing the spinal cord to be punctured by a sharp fragment of [[bone]]. Usually, victims of spinal cord injuries will suffer loss of feeling in certain parts of their body. In milder cases, a victim might only suffer loss of [[hand]] or foot function. More severe injuries may result in [[paraplegia]], [[tetraplegia]] (also known as quadriplegia), or full body [[paralysis]] below the site of injury to the spinal cord.
  
Spinal cord injuries can be caused by falling on the neck or back, or having the spinal cord moved or disrupted in another way. The vertebral bones or [[intervertebral disk]]s can shatter, causing the spinal cord to be punctured by a sharp fragment of [[bone]]. Usually victims of spinal cord injuries will suffer loss of feeling in certain parts of their body. In milder cases a victim might only suffer loss of [[hand]] or [[foot]] function. More severe injury may result in [[paraplegia]], [[tetraplegia]], or full body [[paralysis]] below the site of injury to the spinal cord.
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Damage to upper motor neuron axons in the spinal cord results in a characteristic pattern of ipsilateral deficits. These include [[hyperreflexia]], [[hypertonia]] and muscle weakness. Lower motor neuronal damage results in its own characteristic pattern of deficits. Rather than an entire side of deficits, there is a pattern relating to the [[myotome]] affected by the damage. Additionally, lower motor neurons are characterized by muscle weakness, [[hypotonia]], [[hyporeflexia]] and [[muscle atrophy]].
  
Damage to upper motor neurons axons in the spinal cord results in a characteristic pattern of ipsilateral deficits. These include [[hyperreflexia]], [[hypertonia]] and muscle weakness. Lower motor neuronal damage results in its own characteristic pattern of deficits. Rather than an entire side of deficits, there is a pattern relating to the [[myotome]] affected by the damage. Additionally, lower motor neurons are characterized by muscle weakness, [[hypotonia]], [[hyporeflexia]] and [[muscle atrophy]].
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Spinal shock and neurogenic shock can occur from a spinal injury. Spinal shock is usually temporary, lasting only for 24–48 hours, and is a temporary absence of sensory and motor functions. Neurogenic shock lasts for weeks and can lead to a loss of muscle tone due to disuse of the muscles below the injured site.
  
 
The two areas of the spinal cord most commonly injured are the [[cervical spine]] (C1-C7) and the [[lumbar spine]] (L1-L5). (The notation C1, C7, L1, L5 refer to the location of a specific [[vertebra]] in either the cervical, thoracic, or lumbar region of the spine.)
 
The two areas of the spinal cord most commonly injured are the [[cervical spine]] (C1-C7) and the [[lumbar spine]] (L1-L5). (The notation C1, C7, L1, L5 refer to the location of a specific [[vertebra]] in either the cervical, thoracic, or lumbar region of the spine.)
 +
 +
Spinal cord injury can also be non traumatic and caused by disease ([[transverse myelitis]], [[polio]], [[spina bifida]], [[Friedreich's ataxia]], [[spinal cord tumor]], [[spinal stenosis]] etc.){{cn|date=June 2013}}
  
 
==Additional images==
 
==Additional images==
Line 86: Line 162:
 
  Image:Gray664.png|Cross-section through the spinal cord at the mid-thoracic level.
 
  Image:Gray664.png|Cross-section through the spinal cord at the mid-thoracic level.
 
  Image:Gray666.png|Cross-sections of the spinal cord at varying levels.
 
  Image:Gray666.png|Cross-sections of the spinal cord at varying levels.
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Image:Rabbitspinalcord100x1.jpg|Cross-section of rabbit spinal cord.
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Image:Sus barbatus 01 - Sagittal Section of Vertebral Column - sharp focus.jpg|Sagittal section of pig vertebrae showing a section of the spinal cord.
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File:Slide12ee.JPG|The base of the brain and the top of the spinal cord
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File:Slide1drdr.GIF|Spinal cord. Spinal membranes and nerve roots.Deep dissection. Posterior view.
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File:Slide2drdr.GIF|Spinal cord. Spinal membranes and nerve roots.Deep dissection. Posterior view.
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File:Slide3dsdd.GIF|Spinal cord. Spinal membranes and nerve roots.Deep dissection. Posterior view.
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File:Slide2fer.JPG|Spinal cord. Spinal membranes and nerve roots.Deep dissection. Posterior view.
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File:Slide3fer.JPG|Spinal cord. Spinal membranes and nerve roots.Deep dissection. Posterior view.
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File:Slide4rer.JPG|Spinal cord. Spinal membranes and nerve roots.Deep dissection. Posterior view.
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File:Slide5rer.JPG|Spinal cord. Spinal membranes and nerve roots.Deep dissection. Posterior view.
 
</gallery>
 
</gallery>
  
 
==See also==
 
==See also==
{{Wiktionary}}
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{{commons category|Spinal cord}}
{{commonscat|Spinal cord}}
 
 
* ''[[Cauda equina]]''
 
* ''[[Cauda equina]]''
 
* ''[[Conus medullaris]]''
 
* ''[[Conus medullaris]]''
 
* [[Meninges]]
 
* [[Meninges]]
* [[Spinal nerves]]
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* [[Spinal nerve]]
 
* [[Lumbar puncture]]
 
* [[Lumbar puncture]]
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* [[Neutral spine]]
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* [[Brown–Séquard syndrome]]
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* [[Hereditary spastic paraplegia]] <small>(HSP, or Familial spastic paraplegia – FSP, Strümpell-Lorrain syndrome)</small>
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* [[Poliomyelitis]], [[Post-polio syndrome]]
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* [[Upper-limb surgery in tetraplegia]]
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* [[Redlich-Obersteiner's Zone]]
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 +
==References==
 +
<references/>
  
 
==External links==
 
==External links==
All links retrieved November 21, 2007.
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* [http://biology.clc.uc.edu/fankhauser/Labs/Anatomy_&_Physiology/A&P202/CNS_Histology/Spinal_Cord/Spinal_Cord_Histology.htm Spinal Cord Histology] - A multitude of great images from the [[University of Cincinnati]]
 
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* [http://www.rahulgladwin.com/blog/2006/07/spinal-cord-notes.html Spinal Cord Medical Notes] - Online medical notes on the spinal cord
* [http://biology.clc.uc.edu/fankhauser/Labs/Anatomy_&_Physiology/A&P202/CNS_Histology/Spinal_Cord/Spinal_Cord_Histology.htm Spinal Cord Histology] - A multitude of great Images from the [[University of Cincinnati]]
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* {{cite web|title=The Nervous System: Sensory and Motor Tracts of the Spinal Cord|url=http://www.napavalley.edu/people/briddell/Documents/BIO%20218/15_lecture_presentation.pdf|publisher=Napa Valley College / Southeast Community College  Lincoln, Nebraska|accessdate=20 May 2013}}
* [http://www.rahulgladwin.com/blog/2006/07/spinal-cord-notes.html Spinal Cord Medical Notes] - Online medical notes on the Spinal Cord
 
 
* [http://www.emedicine.com/neuro/topic657.htm eMedicine: Spinal Cord, Topographical and Functional Anatomy]
 
* [http://www.emedicine.com/neuro/topic657.htm eMedicine: Spinal Cord, Topographical and Functional Anatomy]
*WebMD. May 17, 2005. [http://children.webmd.com/tc/Spina-Bifida-Topic-Overview Spina Bifida - Topic Overview] Information about Spina Bifida in fetuses and throughout adulthood. WebMD children's health.
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* WebMD. May 17, 2005. [http://children.webmd.com/tc/Spina-Bifida-Topic-Overview Spina Bifida - Topic Overview] Information about spina bifida in fetuses and throughout adulthood. WebMD children's health. Retrieved March 19, 2007.
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* [http://news.bbc.co.uk/2/hi/uk_news/wales/6274960.stm  Potential for spinal injury repair]  Retrieved February 6, 2008.
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* [http://mousespinal.brain-map.org/ 4000 sets of digital images, showing spatial expression patterns for various genes in adult and juvenile mouse spinal cords] from the [[Allen Institute for Brain Science]]
  
{{nervous system}}
 
{{Spinal cord}}
 
{{Vertebral column and spinal cord}}
 
  
 
[[Category:Life sciences]]
 
[[Category:Life sciences]]
{{credits|171437498}}
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{{credit|Spinal_cord|559968301}}

Revision as of 23:22, 23 June 2013

Spinal cord
Nervous system diagram.png
The spinal cord (in red) connects the brain to the nerves throughout the body
Latin medulla spinalis
Divisions of Spinal Segments
Gray 111 - Vertebral column-coloured.png
Segmental Spinal Cord Level and Function
Level Function
C1-C6 Neck flexors
C1-T1 Neck extensors
C3, C4, C5 Supply diaphragm (mostly C4)
C5, C6 Shoulder movement, raise arm (deltoid); flexion of elbow (biceps); C6 externally rotates the arm (supinates)
C6, C7 Extends elbow and wrist (triceps and wrist extensors); pronates wrist
C7, T1 Flexes wrist
C7, T1 Supply small muscles of the hand
T1 -T6 Intercostals and trunk above the waist
T7-L1 Abdominal muscles
L1, L2, L3, L4 Thigh flexion
L2, L3, L4 Thigh adduction
L4, L5, S1 Thigh abduction
L5, S1, S2 Extension of leg at the hip (gluteus maximus)
L2, L3, L4 Extension of leg at the knee (quadriceps femoris)
L4, L5, S1, S2 Flexion of leg at the knee (hamstrings)
L4, L5, S1 Dorsiflexion of foot (tibialis anterior)
L4, L5, S1 Extension of toes
L5, S1, S2 Plantar flexion of foot
L5, S1, S2 Flexion of toes

The spinal cord is a long, thin, tubular bundle of nervous tissue and support cells that extends from the brain (the medulla oblongata specifically). The brain and spinal cord together make up the central nervous system (CNS). The spinal cord begins at the occipital bone and extends down to the space between the first and second lumbar vertebrae; it does not extend the entire length of the vertebral column. It is around 45 cm (18 in) in men and around 43 cm (17 in) long in women. Also, the spinal cord has a varying width, ranging from 1/2 inch thick in the cervical and lumbar regions to 1/4 inch thick in the thoracic area. The enclosing bony vertebral column protects the relatively shorter spinal cord. The spinal cord functions primarily in the transmission of neural signals between the brain and the rest of the body but also contains neural circuits that can independently control numerous reflexes and central pattern generators. The spinal cord has three major functions: as a conduit for motor information, which travels down the spinal cord, as a conduit for sensory information in the reverse direction, and finally as a center for coordinating certain reflexes. [1]

Structure

The spinal cord is the main pathway for information connecting the brain and peripheral nervous system. The length of the spinal cord is much shorter than the length of the bony spinal column. The human spinal cord extends from the foramen magnum and continues through to the conus medullaris near the second lumbar vertebra, terminating in a fibrous extension known as the filum terminale.

It is about 45 cm (18 in) long in men and around 43 cm (17 in) in women, ovoid-shaped, and is enlarged in the cervical and lumbar regions. The cervical enlargement, located from C3 to T2 spinal segments, is where sensory input comes from and motor output goes to the arms. The lumbar enlargement, located between L1 and S3 spinal segments, handles sensory input and motor output coming from and going to the legs.

The spinal cord is protected by three layers of tissue, called spinal meninges, that surround the canal. The dura mater is the outermost layer, and it forms a tough protective coating. Between the dura mater and the surrounding bone of the vertebrae is a space called the epidural space. The epidural space is filled with adipose tissue, and it contains a network of blood vessels. The arachnoid mater is the middle protective layer. Its name comes from the fact that the tissue has a spiderweb-like appearance. The space between the arachnoid and the underlying pia mater is called the subarachnoid space. The subarachnoid space contains cerebrospinal fluid (CSF). The medical procedure known as a lumbar puncture (or "spinal tap") involves use of a needle to withdraw cerebrospinal fluid from the subarachnoid space, usually from the lumbar region of the spine. The pia mater is the innermost protective layer. It is very delicate and it is tightly associated with the surface of the spinal cord. The cord is stabilized within the dura mater by the connecting denticulate ligaments, which extend from the enveloping pia mater laterally between the dorsal and ventral roots. The dural sac ends at the vertebral level of the second sacral vertebra.

In cross-section, the peripheral region of the cord contains neuronal white matter tracts containing sensory and motor neurons. Internal to this peripheral region is the gray, butterfly-shaped central region made up of nerve cell bodies. This central region surrounds the central canal, which is an anatomic extension of the spaces in the brain known as the ventricles and, like the ventricles, contains cerebrospinal fluid.

The spinal cord has a shape that is compressed dorso-ventrally, giving it an elliptical shape. The cord has grooves in the dorsal and ventral sides. The posterior median sulcus is the groove in the dorsal side, and the anterior median fissure is the groove in the ventral side.

Spinal cord segments

Model of a section of a spine.
A model of segments of the human vertebral column, nerve roots can be seen extending laterally from the (not visible) spinal cord.

The human spinal cord is divided into 31 different segments. At every segment, right and left pairs of spinal nerves (mixed; sensory and motor) form. Six to eight motor nerve rootlets branch out of right and left ventro lateral sulci in a very orderly manner. Nerve rootlets combine to form nerve roots. Likewise, sensory nerve rootlets form off right and left dorsal lateral sulci and form sensory nerve roots. The ventral (motor) and dorsal (sensory) roots combine to form spinal nerves (mixed; motor and sensory), one on each side of the spinal cord. Spinal nerves, with the exception of C1 and C2, form inside intervertebral foramen (IVF). Note that at each spinal segment, the border between the central and peripheral nervous system can be observed. Rootlets are a part of the peripheral nervous system.

In the upper part of the vertebral column, spinal nerves exit directly from the spinal cord, whereas in the lower part of the vertebral column nerves pass further down the column before exiting. The terminal portion of the spinal cord is called the conus medullaris. The pia mater continues as an extension called the filum terminale, which anchors the spinal cord to the coccyx. The cauda equina (“horse’s tail”) is the name for the collection of nerves in the vertebral column that continue to travel through the vertebral column below the conus medullaris. The cauda equina forms as a result of the fact that the spinal cord stops growing in length at about age four, even though the vertebral column continues to lengthen until adulthood. This results in the fact that sacral spinal nerves actually originate in the upper lumbar region. The spinal cord can be anatomically divided into 31 spinal segments based on the origins of the spinal nerves.

Each segment of the spinal cord is associated with a pair of ganglia, called dorsal root ganglia, which are situated just outside of the spinal cord. These ganglia contain cell bodies of sensory neurons. Axons of these sensory neurons travel into the spinal cord via the dorsal roots.

Ventral roots consist of axons from motor neurons, which bring information to the periphery from cell bodies within the CNS. Dorsal roots and ventral roots come together and exit the intervertebral foramina as they become spinal nerves.

The gray matter, in the center of the cord, is shaped like a butterfly and consists of cell bodies of interneurons and motor neurons. It also consists of neuroglia cells and unmyelinated axons. Projections of the gray matter (the “wings”) are called horns. Together, the gray horns and the gray commissure form the “gray H.”

The white matter is located outside of the gray matter and consists almost totally of myelinated motor and sensory axons. “Columns” of white matter carry information either up or down the spinal cord.

Within the CNS, nerve cell bodies are generally organized into functional clusters, called nuclei. Axons within the CNS are grouped into tracts.

There are 33 spinal cord nerve segments in a human spinal cord:

  • 8 cervical segments forming 8 pairs of cervical nerves (C1 spinal nerves exit spinal column between occiput and C1 vertebra; C2 nerves exit between posterior arch of C1 vertebra and lamina of C2 vertebra; C3-C8 spinal nerves through IVF above corresponding cervica vertebra, with the exception of C8 pair which exit via IVF between C7 and T1 vertebra)
  • 12 thoracic segments forming 12 pairs of thoracic nerves (exit spinal column through IVF below corresponding vertebra T1-T12)
  • 5 lumbar segments forming 5 pairs of lumbar nerves (exit spinal column through IVF, below corresponding vertebra L1-L5)
  • 5 sacral segments forming 5 pairs of sacral nerves (exit spinal column through IVF, below corresponding vertebra S1-S5)
  • 3 coccygeal segments joined up becoming a single segment forming 1 pair of coccygeal nerves (exit spinal column through the sacral hiatus).

In the fetus, vertebral segments correspond with spinal cord segments. However, because the vertebral column grows longer than the spinal cord, spinal cord segments do not correspond to vertebral segments in the adult, particularly in the lower spinal cord. For example, lumbar and sacral spinal cord segments are found between vertebral levels T9 and L2, and the spinal cord ends around the L1/L2 vertebral level, forming a structure known as the conus medullaris.

Although the spinal cord cell bodies end around the L1/L2 vertebral level, the spinal nerves for each segment exit at the level of the corresponding vertebra. For the nerves of the lower spinal cord, this means that they exit the vertebral column much lower (more caudally) than their roots. As these nerves travel from their respective roots to their point of exit from the vertebral column, the nerves of the lower spinal segments form a bundle called the cauda equina.

Spinal cord

There are two regions where the spinal cord enlarges:

  • Cervical enlargement - corresponds roughly to the brachial plexus nerves, which innervate the upper limb. It includes spinal cord segments from about C4 to T1. The vertebral levels of the enlargement are roughly the same (C4 to T1).
  • Lumbosacral enlargement - corresponds to the lumbosacral plexus nerves, which innervate the lower limb. It comprises the spinal cord segments from L2 to S3 and is found about the vertebral levels of T9 to T12.

Embryology

The spinal cord is made from part of the neural tube during development. As the neural tube begins to develop, the notochord begins to secrete a factor known as Sonic hedgehog or SHH. As a result, the floor plate then also begins to secrete SHH, and this will induce the basal plate to develop motor neurons. Meanwhile, the overlying ectoderm secretes bone morphogenetic protein (BMP). This induces the roof plate to begin to secrete BMP, which will induce the alar plate to develop sensory neurons. The alar plate and the basal plate are separated by the sulcus limitans.

Medulla spinalis of 8-week-old human embryo

Additionally, the floor plate also secretes netrins. The netrins act as chemoattractants to decussation of pain and temperature sensory neurons in the alar plate across the anterior white commissure, where they then ascend towards the thalamus.

Lastly, it is important to note that the past studies of Viktor Hamburger and Rita Levi-Montalcini in the chick embryo have been further proven by more recent studies which demonstrated that the elimination of neuronal cells by programmed cell death (PCD) is necessary for the correct assembly of the nervous system.

Overall, spontaneous embryonic activity has been shown to play a role in neuron and muscle development but is probably not involved in the initial formation of connections between spinal neurons.

Blood supply

The spinal cord is supplied with blood by three arteries that run along its length starting in the brain, and many arteries that approach it through the sides of the spinal column. The three longitudinal arteries are called the anterior spinal artery, and the right and left posterior spinal arteries.[2] These travel in the subarachnoid space and send branches into the spinal cord. They form anastamoses (connections) via the anterior and posterior segmental medullary arteries, which enter the spinal cord at various points along its length.[2] The actual blood flow caudally through these arteries, derived from the posterior cerebral circulation, is inadequate to maintain the spinal cord beyond the cervical segments.

The major contribution to the arterial blood supply of the spinal cord below the cervical region comes from the radially arranged posterior and anterior radicular arteries, which run into the spinal cord alongside the dorsal and ventral nerve roots, but with one exception do not connect directly with any of the three longitudinal arteries.[2] These intercostal and lumbar radicular arteries arise from the aorta, provide major anastomoses and supplement the blood flow to the spinal cord. In humans the largest of the anterior radicular arteries is known as the artery of Adamkiewicz, or anterior radicularis magna (ARM) artery, which usually arises between L1 and L2, but can arise anywhere from T9 to L5. [3] Impaired blood flow through these critical radicular arteries, especially during surgical procedures that involve abrupt disruption of blood flow through the aorta for example during aortic aneursym repair, can result in spinal cord infarction and paraplegia.

Somatosensory organization

Spinal cord tracts.

Somatosensory organization is divided into the dorsal column-medial lemniscus tract (the touch/proprioception/vibration sensory pathway) and the anterolateral system, or ALS (the pain/temperature sensory pathway). Both sensory pathways use three different neurons to get information from sensory receptors at the periphery to the cerebral cortex. These neurons are designated primary, secondary and tertiary sensory neurons. In both pathways, primary sensory neuron cell bodies are found in the dorsal root ganglia, and their central axons project into the spinal cord.

In the dorsal column-medial leminiscus tract, a primary neuron's axon enters the spinal cord and then enters the dorsal column. If the primary axon enters below spinal level T6, the axon travels in the fasciculus gracilis, the medial part of the column. If the axon enters above level T6, then it travels in the fasciculus cuneatus, which is lateral to the fasiculus gracilis. Either way, the primary axon ascends to the lower medulla, where it leaves its fasiculus and synapses with a secondary neuron in one of the dorsal column nuclei: either the nucleus gracilis or the nucleus cuneatus, depending on the pathway it took. At this point, the secondary axon leaves its nucleus and passes anteriorly and medially. The collection of secondary axons that do this are known as internal arcuate fibers. The internal arcuate fibers decussate and continue ascending as the contralateral medial lemniscus. Secondary axons from the medial lemniscus finally terminate in the ventral posterolateral nucleus (VPL) of the thalamus, where they synapse with tertiary neurons. From there, tertiary neurons ascend via the posterior limb of the internal capsule and end in the primary sensory cortex.

The proprioception of the lower limbs differs from the upper limbs and upper trunk. There is a four-neuron pathway for lower limb proprioception. This pathway initially follows the dorsal spino-cerebellar pathway. It is arranged as follows: proprioceptive receptors of lower limb -> peripheral process -> dorsal root ganglion -> central process -> Clarke's column -> 2nd order neuron -> medulla oblogata (Caudate nucleus) -> 3rd order neuron -> VPL of thalamus -> 4th order neuron -> posterior limb of internal capsule -> corona radiata -> sensory area of cerebrum.

The anterolateral system works somewhat differently. Its primary neurons axons enter the spinal cord and then ascend one to two levels before synapsing in the substantia gelatinosa. The tract that ascends before synapsing is known as Lissauer's tract. After synapsing, secondary axons decussate and ascend in the anterior lateral portion of the spinal cord as the spinothalamic tract. This tract ascends all the way to the VPL, where it synapses on tertiary neurons. Tertiary neuronal axons then travel to the primary sensory cortex via the posterior limb of the internal capsule.

It should be noted that some of the "pain fibers" in the ALS deviate from their pathway towards the VPL. In one such deviation, axons travel towards the reticular formation in the midbrain. The reticular formation then projects to a number of places including the hippocampus (to create memories about the pain), the centromedian nucleus (to cause diffuse, non-specific pain) and various parts of the cortex. Additionally, some ALS axons project to the periaqueductal gray in the pons, and the axons forming the periaqueductal gray then project to the nucleus raphes magnus, which projects back down to where the pain signal is coming from and inhibits it. This helps control the sensation of pain to some degree.

Motor organization

The corticospinal tract serves as the motor pathway for upper motor neuronal signals coming from the cerebral cortex and from primitive brainstem motor nuclei.

Cortical upper motor neurons originate from Brodmann areas 1, 2, 3, 4, and 6 and then descend in the posterior limb of the internal capsule, through the crus cerebri, down through the pons, and to the medullary pyramids, where about 90% of the axons cross to the contralateral side at the decussation of the pyramids. They then descend as the lateral corticospinal tract. These axons synapse with lower motor neurons in the ventral horns of all levels of the spinal cord. The remaining 10% of axons descend on the ipsilateral side as the ventral corticospinal tract. These axons also synapse with lower motor neurons in the ventral horns. Most of them will cross to the contralateral side of the cord (via the anterior white commissure) right before synapsing.

The midbrain nuclei include four motor tracts that send upper motor neuronal axons down the spinal cord to lower motor neurons. These are the rubrospinal tract, the vestibulospinal tract, the tectospinal tract and the reticulospinal tract. The rubrospinal tract descends with the lateral corticospinal tract, and the remaining three descend with the anterior corticospinal tract.

The function of lower motor neurons can be divided into two different groups: the lateral corticospinal tract and the anterior cortical spinal tract. The lateral tract contains upper motor neuronal axons which synapse on dorsal lateral (DL) lower motor neurons. The DL neurons are involved in distal limb control. Therefore, these DL neurons are found specifically only in the cervical and lumbosacral enlargements within the spinal cord. There is no decussation in the lateral corticospinal tract after the decussation at the medullary pyramids.

The anterior corticospinal tract descends ipsilaterally in the anterior column, where the axons emerge and either synapse on lower ventromedial (VM) motor neurons in the ventral horn ipsilaterally or descussate at the anterior white commissure where they synapse on VM lower motor neurons contralaterally . The tectospinal, vestibulospinal and reticulospinal descend ipsilaterally in the anterior column but do not synapse across the anterior white commissure. Rather, they only synapse on VM lower motor neurons ipsilaterally. The VM lower motor neurons control the large, postural muscles of the axial skeleton. These lower motor neurons, unlike those of the DL, are located in the ventral horn all the way throughout the spinal cord.

Spinocerebellar tracts

Proprioceptive information in the body travels up the spinal cord via three tracts. Below L2, the proprioceptive information travels up the spinal cord in the ventral spinocerebellar tract. Also known as the anterior spinocerebellar tract, sensory receptors take in the information and travel into the spinal cord. The cell bodies of these primary neurons are located in the dorsal root ganglia. In the spinal cord, the axons synapse and the secondary neuronal axons decussates and then travel up to the superior cerebellar peduncle where they decussate again. From here, the information is brought to deep nuclei of the cerebellum including the fastigial and interposed nuclei.

From the levels of L2 to T1, proprioceptive information enters the spinal cord and ascends ipsilaterally, where it synapses in Clarke's nucleus. The secondary neuronal axons continue to ascend ipsilaterally and then pass into the cerebellum via the inferior cerebellar peduncle. This tract is known as the dorsal spinocerebellar tract.

From above T1, proprioceptive primary axons enter the spinal cord and ascend ipsilaterally until reaching the accessory cuneate nucleus, where they synapse. The secondary axons pass into the cerebellum via the inferior cerebellar peduncle where again, these axons synapse on cerebellar deep nuclei. This tract is known as the cuneocerebellar tract.

Motor information travels from the brain down the spinal cord via descending spinal cord tracts. Descending tracts involve two neurons: the upper motor neuron (UMN) and lower motor neuron (LMN).[4] A nerve signal travels down the upper motor neuron until it synapses with the lower motor neuron in the spinal cord. Then, the lower motor neuron conducts the nerve signal to the spinal root where efferent nerve fibers carry the motor signal toward the target muscle. The descending tracts are composed of white matter. There are several descending tracts serving different functions. The corticospinal tracts (lateral and anterior) are responsible for coordinated limb movements.[4]

Injury

Spinal cord injuries can be caused by trauma to the spinal column (stretching, bruising, applying pressure, severing, laceration, etc.). The vertebral bones or intervertebral disks can shatter, causing the spinal cord to be punctured by a sharp fragment of bone. Usually, victims of spinal cord injuries will suffer loss of feeling in certain parts of their body. In milder cases, a victim might only suffer loss of hand or foot function. More severe injuries may result in paraplegia, tetraplegia (also known as quadriplegia), or full body paralysis below the site of injury to the spinal cord.

Damage to upper motor neuron axons in the spinal cord results in a characteristic pattern of ipsilateral deficits. These include hyperreflexia, hypertonia and muscle weakness. Lower motor neuronal damage results in its own characteristic pattern of deficits. Rather than an entire side of deficits, there is a pattern relating to the myotome affected by the damage. Additionally, lower motor neurons are characterized by muscle weakness, hypotonia, hyporeflexia and muscle atrophy.

Spinal shock and neurogenic shock can occur from a spinal injury. Spinal shock is usually temporary, lasting only for 24–48 hours, and is a temporary absence of sensory and motor functions. Neurogenic shock lasts for weeks and can lead to a loss of muscle tone due to disuse of the muscles below the injured site.

The two areas of the spinal cord most commonly injured are the cervical spine (C1-C7) and the lumbar spine (L1-L5). (The notation C1, C7, L1, L5 refer to the location of a specific vertebra in either the cervical, thoracic, or lumbar region of the spine.)

Spinal cord injury can also be non traumatic and caused by disease (transverse myelitis, polio, spina bifida, Friedreich's ataxia, spinal cord tumor, spinal stenosis etc.)[citation needed]

Additional images

See also

Commons
Wikimedia Commons has more media related to:
  • Cauda equina
  • Conus medullaris
  • Meninges
  • Spinal nerve
  • Lumbar puncture
  • Neutral spine
  • Brown–Séquard syndrome
  • Hereditary spastic paraplegia (HSP, or Familial spastic paraplegia – FSP, Strümpell-Lorrain syndrome)
  • Poliomyelitis, Post-polio syndrome
  • Upper-limb surgery in tetraplegia
  • Redlich-Obersteiner's Zone

References
ISBN links support NWE through referral fees

  1. Maton, Anthea and Jean Hopkins, Charles William McLaughlin, Susan Johnson, Maryanna Quon Warner, David LaHart, Jill D. Wright (1993). Human Biology and Health. Englewood Cliffs, New Jersey, USA: Prentice Hall, 132–144. ISBN 0-13-981176-1. 
  2. 2.0 2.1 2.2 Moore, Keith and Anne Agur (2007). Essential Clinical Anatomy, Third Edition. Lippincott Williams & Wilkins, 298. ISBN 0-7817-6274-X. 
  3. Biglioli, Paolo and et alia (April 2004). Upper and lower spinal cord blood supply: the continuity of the anterior spinal artery and the relevance of the lumbar arteries. Journal of Thoracic and Cardiovascular Surgery 127 (4): 1188–1192.
  4. 4.0 4.1 Saladin. Anatomy and Physiology, 5th Ed.

External links

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