Difference between revisions of "Neurons - Anatomy & Physiology"

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==Introduction==
 
==Introduction==
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[[Image:neurondiagram.jpg|thumb|right|600px|Diagram of a neuron. LadyofHats, '''Wikimedia Commons''', 2007]]
 
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[[Image:Aspinall Slide4.JPG|thumb|right|600px|<small>Image from [http://www.elsevierhealth.co.uk/veterinary-nursing/spe-60136/ Aspinall, The Complete Textbook of Veterinary Nursing], Elsevier Health Sciences, ''All rights reserved''</small>]]
 
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Nerves allow electrical impulses to propagate along their elongated cell extensions and facilitate the transfer of information throughout the body. Neural tissue is found within the [[Nervous and Special Senses - Anatomy & Physiology#Central Nervous System (CNS)|central nervous system (CNS)]] and the [[PNS Structure - Anatomy & Physiology|peripheral nervous system (PNS)]] and the composition and constituent parts of neurones and their surrounding cells differ only slightly.
 
  
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===CNS Neurons===
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* There are two major cell types within the CNS:
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Neural tissue found in the CNS is composed of [[Central Nervous System - Anatomy & Physiology#Grey Matter|'''gray''']] and [[Central Nervous System - Anatomy & Physiology#White Matter|'''white''' matter]]. The gray matter includes nerve cell bodies and some short branches from these cell bodies. The white matter is composed of the long extensions of the nerve, the nerve fibres.
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#'''[[#Neurons|Neurons]]'''
 +
#* These are the "functional" cells of the central nervous system.
 +
#'''Glial cells:'''
 +
#* These play a supporting role within the CNS, and include:
 +
#** [[#Astrocytes|Astrocytes]]
 +
#** [[#Oligodendrocytes|Oligodendrocytes]]
 +
#** [[#Microglial Cells|Microglial cells]]
 +
#** [[#Ependymal Cells|Ependymal Cells]]
 +
#** [[#Choroid Plexus Epithelial Cells|Choroid plexus epithelial cells]]
  
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Within the central nervous system there are two major cell types; '''neurons''' which are the "functional" cells of the central nervous system and '''glial cells''' which play a supporting role within the CNS. Within the CNS there are also a number of other cells that play a supportive role to the neurone and these include [[#Astrocytes|astrocytes]], [[#Oligodendrocytes|oligodendrocytes]], [[#Microglial Cells|microglial cells]], [[#Ependymal Cells|ependymal cells]] and [[#Choroid Plexus Epithelial Cells|choroid plexus epithelial cells]].
+
==Neurons==
  
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===PNS Neurons===
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* Neurons are the functional cells of both the CNS and PNS.
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The PNS contains a series of paired nerves that extend from the spinal nerves originating from the spinal cord and [[Cranial Nerves - Anatomy & Physiology|cranial nerves]] that originate from the brain stem. Neurones found in the PNS are similar to those found in the CNS aside from a few subtle differences. Supportive cells that surround the nerves in the PNS are called [[Schwann Cell|Schwann cells]] (rather than Glial cells). The main neurotransmitter found in the PNS is [[Neurotransmitters - Anatomy & Physiology#Other SMTs|acetylcholine]].
+
** They are highly specialised to process and transmit signals.  
  
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==Neurons==
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Structure:
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A typical neuron can be exemplified by a motor neuron in which the cell body of the nerve is located within the gray matter of the [[Spinal Cord - Anatomy & Physiology|spinal cord]] and the nerve fibre, or axon, extends to the [[Muscles - Anatomy & Physiology|muscle]]. Nerve '''axons can be very long''' permitting electrical impulses to be sent over long distances throughout the body. The information below specifically regarding neurons is interchangeable between the CNS and PNS and therefore links have been provided where appropriate on this page to the [[PNS Structure - Anatomy & Physiology|'''PNS Structure''']].
 
  
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===Basic Nerve Structure===
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====Soma====
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The basic structure of a nerve is that of a '''cell body (soma)''', which has a '''single long nerve fibre (axon)''', attached at one end to the cell body and at the other end to another nerve cell body or to a structure requiring nerve impulses such as skeletal muscle. The interface between two nerves or a nerve and another structure is called the '''nerve synapse'''. The cell body of the nerve itself also contains numerous '''dendrites''' which increase its surface area enabling other nerve axons to connect with the cell body. The cell body usually has connections with many other axons from other nerve cells with many synapses on the cell body. The axon of the nerve cell is usually surrounded by some form of insulating protection, or '''myelination'''. This protective layer is called the myelin sheath.
 
  
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===Soma (Cell Body)===
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* The soma, or cell body is the central part of the neuron.
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The soma, or cell body, is the central part of the neuron and contains the nucleus of the cell, the rough and smooth endoplasmic reticulum, ribosomes and golgi apparatus. Similar processes that would be undertaken by any cell occur within the soma and due to the organelles it contains, the soma is where most protein synthesis occurs.
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* The soma contains:
 +
** The nucleus
 +
** Smooth and rough endoplasmic reticulum
 +
** Ribosomes
 +
** Golgi apparatus
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** Due to the organelles it contains, the soma is where most protein synthesis occurs.
  
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===Dendrites===
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====Dendrites====
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The neuron's dendrites are effectively branching extensions of the cell body. Collectively, these branching structures of the dendrites are known as the "dendritic tree". The dendritic tree is the site where '''input to the neuron''' occurs via synapses with axons from other nerve cells. In most cases nerve impulses travel from other nerve cells to the cell body and are then conducted along the nerve cell's own axon to other cell bodies. However, dendrites themselves are unable to propagate nerve impulses in the manner of axons as dendrites are unable to secrete [[Neurotransmitters - Anatomy & Physiology|neurotransmitters]]. Similarly axons do not possess the chemoreceptors that are found within the dendrites and are therefore unable to receive nerve impulses. Nerve impulses are therefore conducted in one direction only.
 
  
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===Axon (Nerve Fibre)===
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* The neuron's dendrites are branching extensions of the cell body.
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The axon is a very fine projection that can measure up to thousands of times the diameter of the soma in length. The axon carries nerve signals away from the soma. The structure and function of the axon is very similar between the CNS and the PNS.  
+
** Collectively, branching structure of the dendrites is known as the "dendritic tree".
 +
* The dendritic tree is the site where input to the neuron occurs. [[Image:neurondiagram.jpg|thumb|right|150px|Diagram of a neuron. LadyofHats, '''Wikimedia Commons''', 2007]]
 +
* Information outflow can also occur in the reverse direction, from the soma to the dendrites.
 +
** However, information is not transmitted from the dendrites in this way, because axons do not possess chemoreceptors and dendrites cannot secrete neurotransmitter chemicals.
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*** Nerve impulses are therefore conducted in one direction only.
  
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Axon has been covered in detail within the [[PNS Structure - Anatomy & Physiology#Structure|PNS Structure and Anatomy]] page.
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====Axon====
  
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==Glial Cells==
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* The axon is a very fine projection that can measure up to thousands of time the soma diameter in length.
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Glial cells make up approximately 50% of the volume of the nervous system and the number of glial cells out-numbers the nerve cells by a factor of ten to one. The primary function of glial cell is to provide '''support to the neuronal cells'''. There are several different types of glial-style cell; in the PNS these are referred to as '''[[Schwann Cell|Schwann cells]]''' and whilst in the CNS they are referred to as '''Oligodendrocytes'''.  
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* The axon carries nerve signals away from the soma.
 +
* Most neurons have only one axon.
 +
** However, this will usually undergo extensive branching to communicate with many target cells.
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* The axon emerged from the soma at the axon hillock
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**  The axon hillock has the highest density of voltage-dependent Na<sup>+</sup>channels, making it the most easily-excited part of the neuron.  
 +
** The axon and axon hillock are generally involved in information outflow.
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*** However, this region can also receive input from other neurons.
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* At the end of the axon, there are synapses.
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** Neurotransmitter chemicals are released here in order to communicate with target neurons.
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* The axon is dependent on the cell body for all its metabolic functions.
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** The axon has no Nissl substance: it cannot make its own protein, and so the soma must undertake this function.
 +
** The soma must also dispose of the axon's waste.
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** Death or damage to the cell body will therefore result in axon compromise or degeneration
  
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Glial cells form a protective networked layer around the neuron within the CNS and also help to maintain the fluid content of the tissue surrounding the nerve. This protective layer is referred to as the '''myelin sheath''' and axons that are wrapped within this myelin sheath are able to conduct nerve impulses at higher speeds than those that are not wrapped. During foetal development the glial cells wrap around axons numerous times and as the glial cell matures it loses the majority of its cytoplasm. The remaining cellular structure consists of many layers of tightly packed lipid membranes around the axon.
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Function:
−
 
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Periodically along the surface of the wrapped glial cell there are gaps approximately every 1-2mm. Each gap allows the environment external to the myelin sheath to be exposed to the axon for approximately 1-2υm. These gaps in the sheath are called the '''Nodes of Ranvier''' and are important in the conduction of impulses along the axon.
 
  
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===Schwann Cell===
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* Neurons are the functional cells of both the CNS and PNS.
−
Details regarding the Schwann cell have been covered within the [PNS Structure - Anatomy & Physiology#Schwann Cell|PNS Structure and Anatomy]].
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** They are highly specialised to process and transmit signals.
 +
* Neurons in different parts of the nervous system perform different functions.
 +
** There is therfore variation in the size, shape, structure, metabolism, function shape and electrochemical properties between locations.
  
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===Oligodendrocytes===
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==Astrocytes==
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[[Image:oligodendrocyte.jpg|thumb|right|250px|Oligodendrocyte]]
 
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The term "oligodendrocyte" literally means a "cell with many branches". Oligodendrocyte precursors are found throughout the CNS and these precursors are post-mitotic, meaning oligodendrocytes can be replaced. Oligodendrocytes can [[PNS Structure - Anatomy & Physiology#The Process of Myelination|myelinate]] up to 40 axons at a time. The myelin sheath formed by oligodendrocytes has numerous functions including; decreasing any ion leakage from the axon lowering the capacitance of the cell membrane and increasing impulse speed along the axon by allowing saltatory conduction of action potentials between the nodes of Ranvier.
 
  
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==Nerve Impulse Propagation==
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* Astrocytes, or astroglia, are star-shaped glial cells in the brain.
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Nerves are able to create, amplify and propagate electrical impulses that run along their axon. These nerve impulses are a form of action potential that is carried by ions. The nerve impulse is in effect an electrical difference between the inside and outside of the axon and is caused by ion movements across the membrane. Nerve impulses can occur from a number of sources including sensory cells, action potentials from other connected nerves or spontaneous depolarisation of the nerve cell membrane.
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* Astrocytes have several functions:
 +
** Biochemical support of the endothelial cells forming the blood-brain barrier
 +
** Provision of nutrients to nervous tissue
 +
** Repair and scarring in the brain
  
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===Unmyelinated Axons===
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Structure:
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Conduction of an action potential along an unmyelinated axon, i.e. an axon not covered by some form of glial cell, is a '''much slower''' form of nerve impulse propagation than that of a myelinated axon.  
+
[[Image:astrocyte.jpg|thumb|right|150px|Astrocyte (immunofluorescence staining). Image courtesy of Alliance Européenne Dana pour le Cerveau (EDAB)]]
 +
* Astrocytes are "star-shaped".
 +
** Their many processes envelope synapses made by neurons.  
 +
* Astrocyte expression of glial fibrillary acidic protein (GFAP) allows their identification histologically.
  
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At a given moment when the action-potential threshold is reached '''voltage-gated sodium channels''' within the membrane of the axon will open resulting in an influx of ions following their electro-chemical gradient. This Na<sup>+</sup> influx causes the axon to accumulate a positive charge and results in '''cellular depolarisation'''. The '''extracellular area''' of the axon therefore looses its positive charge, becoming '''more negative''' resulting in a current of positive charge that flows  through the tissue towards the axon. The membrane of the axon is not a perfect insulator and at some regions on the axon, particularly within the area of the axon in front of the action potential, the voltage-dependant ion channels have not activated yet. This means that '''some of the positive charge is able to flow out of the axon membrane''' in these regions and this positive charge outflow is mainly via '''potassium'''. This outward leakage of potassium results in the current within the axon only being able to travel a short distance before the nerve impulse decays. However the effect of the current locally on the voltage-gated channels means that the nerve impulse is able to open voltage-gated channels within its immediate vicinity and this is enough for the signal to propagate along the nerve.
+
Function:
−
 
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The axon membrane at rest has a slightly negative charge and the local current of positive charge during an action potential reduces this negative charge. This process is called '''depolarisation''' and when this has progressed sufficiently, it results in the membrane potential reaching threshold level allowing the voltage-gated sodium channels to open. These channels only stay open for approximately 0.5ms and when they close the membrane is depolarised for a further 0.1-0.5ms. These voltage-gated channels are unable to open again for a short period post depolarisation. This is called the '''refractory period'''.
 
−
 
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The '''internal electrical resistance''' within the axon is the determining factor regarding how fast the local positive current impulse can pass through. The less the internal resistance, the higher the conduction velocity of the nerve impulses due to the locally positive current being able to depolarise the membrane to threshold level over a greater distance. This internal resistance decreases as a function of axon diameter an therefore '''thicker axons are able to conduct nerve impulses more rapidly'''.
 
  
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===Myelinated Axons===
+
* '''Structural'''
−
The addition of the myelin sheath to nerve axons greatly '''enhances the speed''' with which they are able to conduct nerve impulses. Unmyelinated nerves are able to conduct up to speeds to 25m/s whilst myelinated nerves are capable of up to 100m/s. For an unmyelinated nerve to conduct impulses at the same rate, it would have to have a diameter of approximately 100 times that of the myelinated nerve.  
+
** Astrocytes are involved in the physical structuring of the brain:
−
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*** Interplay between astrocytes and mesenchymal cells is important in sculpting sulci in development.
−
As with unmyelinated axons, an area of locally positive charge flows through the cytosol from the activated area within the axon. The major difference with myelinated nerves is that where the current effectively leaks away in unmyelinated nerves, here the current is only able to leak away through '''nodes of Ranvier''' (see above). Therefore even a weak current is able to depolarise the axon membrane to the threshold level. There is a small loss of current but as these nodes are relatively small the local current is able to travel much further in myelinated axons resulting in a much higher conduction velocity.  
+
*** Astrocytes delineate the  CNS  by creating the CNS-PNS interface.
−
+
*** The cells cover the surface of the brain.
−
The influx of sodium ions via voltage-gated channels is only possible at these nodes and therefore the nerve impulse is able to effectively 'jump' from node to node. This type of impulse propagation is called '''saltatory conduction'''. Myelination of axons in mammals means that the nervous system can sustain a large number of high velocity axons within a relatively small space.
+
**** In this situation, the astrocytes' surface is coated by a basal lamina.
 +
*** Astrocytes take over from fibroblasts to play an important role in abscess formation, as it is difficult to induce fibrosis in the CNS.
 +
*** Astrocytes are the only differentiated CNS cell that retains the ability to divide.
 +
* '''Metabolic'''
 +
** Astrocytes may control glucose levels and provide lactate for neurone for energy production
 +
*** This is by the lactate shuttle mechanism.
 +
** Astrocytes have high levels of antioxidants, such as glutathione peroxidase.
 +
*** This protects neurons from damage by reactive oxygen species.
 +
* '''Blood brain barrier'''
 +
** It was originally thought that the astrocyte end-feet encircling endothelial cells aided in the maintenance of the blood-brain barrier.
 +
***  The tight junctions and basal lamina of the endothelial cells are now considered to play a more substantial role in maintaining the barrier.
 +
* '''Transmitter reuptake'''
 +
** Astrocytes:
 +
**# Isolate synapses
 +
**# Control potassium levels
 +
**# Take up transmitters
 +
**#* Astrocytes express transporters for several neurotransmitters, including glutamate, ATP and GABA.
 +
**#** Glutamate is particularly well taken up, and converted to glutamine using glutamine synthase.
 +
* '''Regulation of ion concentration in the extracellular space'''
 +
** Astrocytes release K<sup>+</sup>  when neurons are active, increasing the local extracellular K<sup>+</sup> concentration.  
 +
** Astrocytes have a high density of K<sup>+</sup> channels, allowing them to rapidly clear the excess accumulation in the extracellular space.
 +
*** If this function fails extracellular K<sup>+</sup> concentration rises. This leads neuronal depolarisation, and may be a cause of epilepsy.
 +
* '''Vasomodulation'''
 +
** Astrocytes may serve as intermediaries in neuronal regulation of blood flow.
 +
* '''Promotion of the myelinating activity of oligodendrocytes'''
 +
** Neuronal electrical activity releases ATP, which stimulates myelin formation.
 +
** This stimulation occurs via astrocytes, which secrete LIM in response to the ATP.
 +
*** LIM is a regulatory protein that promotes the myelinating activity of oligodendrocytes.
 +
* '''Control of inflammation'''
 +
** Astrocytes may play a crucial rule in controlling inflammation in the CNS, by:
 +
*** Secreting chemokines
 +
***  Playing an important role in the entry of inflammatory cells to the CNS.
 +
** When astrocytes are lost or destroyed, the neurons become very vulnerable in the face of inflammation.
  
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==Synapses==
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==Oligodendrocytes==
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The synapses found at the end of axons are fundamental to the functioning of the nervous system as they facilitate communication between nerves and provide an interconnected network for many of the complex processes required by organisms. Synapses are required as the lipid bi-layer of the cell membrane has a relatively large electrical resistance making electrical impulse propagation directly between cells difficult.
+
Structure:
−
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[[Image:oligodendrocyte.jpg|thumb|right|150px|Oligodendrocyte (immunofluorescence staining), Jurjen Broeke, '''WikiMedia Commons''']]
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The most common form of nerve synapse is the chemical synapse which utilises [[Neurotransmitters - Anatomy & Physiology|neurotransmitters]]. When a nerve impulse reaches a synapse, neurotransmitters are released by the presynaptic terminal of the synapse and these transmitters diffuse to the membrane of the post-synaptic membrane where they bind to receptors. These receptors cause an inhibition or excitement in that nerve resulting in either blocking further electrical impulses or the further propagation of a signal.
 
  
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===Neuromuscular Synapses===
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* "Oligodendrocyte" literally means a "cell with many branches".
−
These chemical synapses provide connection '''between nerves and skeletal muscle cells'''. These synapses are most commonly found residing within groups of muscle cells where each neuron is in contact with several muscle cells but each muscle cell is only ever connected to one neuron. The nerve axon branches out prior to the muscle cells allowing multiple synapses with muscle cells from a single nerve axon. A '''synaptic cleft''' of approximately 30-50nm is found between the nerve synapse and the muscle cell.  
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* Oligodendrocyte precursors are found throughout the CNS.
 +
** These precursors are post-mitotic, meaning oligodendrocytes can be replaced.
  
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The nerve terminal membrane or pre-synaptic membrane contains numerous vesicles that contain the neurotransmitter, in this case most commonly [[Neurotransmitters - Anatomy & Physiology#Other SMTs|acetylecholine (ACh)]]. Once released from the vesicles the ACh diffuses across the synaptic cleft and binds to receptors in the muscle cell membrane. This binding causes ligand-gated ion channels to open which are permeable to both Na<sup>+</sup> and K<sup>+</sup>. The movement of potassium is relatively small due to there being only a small electrochemical gradient between the extra and intracellular environment. However there is a large influx of sodium into the cell and consequently this causes depolarisation in the muscle cell thus propagating the impulse into mechanical movement. Within the neuromuscular junction, the release of vesicles is facilitated by an influx of calcium into the pre-synaptic nerve just prior to exocytosis. The calcium enters the pre-synaptic nerve via voltage-gated Ca<sup>2+</sup> channels. There are several mechanisms that reduce the intracellular concentration of calcium once vesicles begin to be released to ensure that the neurotransmitter release is brief to prevent hyperpolarisation.
+
Function:
  
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The particular neurotransmitter ACh is heavily recycled within the synaptic cleft via endocytosis and over time the levels of endocytosis and exocytosis balance one-another resulting in a stable pre-synpatic membrane. Depolarisation within the muscle cell will last as long as the ACh is present in sufficient quantities within the synaptic cleft. In reality this only lasts a few milliseconds as the synaptic cleft also contains the enzyme '''acetylcholinesterase''' which hydrolyses the ACh into acetate and choline. Due to this mechanisms, the calcium restriction and the endocytosis, only one action potential is generated within the muscle fibre.
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* Oligodendrocytes can [[PNS Structure - Anatomy & Physiology#The Process of Myelination|myelinate]] up to 40 axons at a time.
 +
** The myelin sheath functions to:
 +
**# Decrease ion leakage and lower the capacitance of the cell membrane.  
 +
**# Increase impulse speed by allowing saltatory conduction of action potentials between the nodes of Ranvier.
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**#* Nodes of Ranvier are spaces along the axon in between oligodendrocytes.
  
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===Inter-neuron Synapses===
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==Microglial Cells==
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The interaction between nerves within the synapse is fairly similar to that within the neuromuscular junction but there are several key differences. Firstly the receiving neuron will be receiving information from multiple other nerves rather than just one nerve as per a muscle cell. Secondly there are many more types of neurotransmitters utilised by inter-neuron synapses than just ACh. There are also a wider range of types of synapses between neurons that include both excitatory which will propagate a nerve impulse, but also inhibitory which will prevent another synapse on the same nerve propagating a nerve impulse.
+
Structure:
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<br />
 
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<br />
 
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'''Excitatory synapses''' will lead to depolarisation of the target neuron and if this depolarisation reaches the threshold level, an impulse will be propagated. This depolarisation is referred to as '''excitatory post-synaptic potential (EPSP)''' as it brings the membrane potential of the target cell closer to the threshold for an action potential. If several excitatory synapses on the same nerve excite simultaneously, the target cell will receive the sum of all these individual synaptic potentials. Therefore multiple e
 
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active excitatory synapses are more likely to cause a threshold depolarisation.
 
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<br />
 
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<br />
 
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'''Inhibitory synapses''' between neurons also have many synapses from other nerves connecting to a single nerve providing a network of connections. However, the neurotransmitter molecules within an inhibitory synapse usually open ligand-gated channels for chloride-ions and potassium ions. K<sup>+</sup> will then flow out of the cell whilst Cl<sup>-</sup> flows into the cell resulting in the cytosol of the cell becoming more negative. This adjustment in membrane potential is called a '''inhibitory postsynaptic potential (IPSP)''' as it takes the membrane potential of the cell further from the threshold level required to generate an action potential.
 
  
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==Astrocytes==
+
* Microglia are the CNS representatives of mononuclear phagocyte system.
−
Astrocytes, or astroglia, are star-shaped glial cells within the brain which have many processes that envelope synapses made between neurons. Astrocytes have several functions including the biochemical support of the endothelial cells in forming the blood-brain barrier, provision of nutrients to nervous tissues and to form scar tissue during repair of the brain. The fluorescent image of astrocytes on the right is possible because astrocytes express glial fibrillary acidic protein (GFAP) which facilitates their identification. The following paragraphs provide a more detailed overview of the many functions of the astrocyte within the brain.
+
** As such, they are derived from [[Bone Marrow - Anatomy & Physiology|bone marrow]], and are first seen in brain just before vascularistion.
−
[[Image:astrocyte.jpg|thumb|right|250px|Astrocyte (immunofluorescence staining). Image courtesy of Alliance Européenne Dana pour le Cerveau (EDAB)]]
+
* There are two populations of microglial cells:
−
Another key role of astrocytes within the brain where they play an important structural role as well as performing the roles described above. Astrocytes are involved in the physical structuring of the brain and the interplay between astrocytes and mesenchymal cells is important in sculpting [[Forebrain_-_Anatomy_%26_Physiology#Cerebral_Cortex|sulci]] in development. Astrocytes also delineate the  CNS from the PNS by creating the CNS-PNS interface. The cells cover the surface of the brain and where this occurs the astrocytes' surface is coated by a basal lamina.
+
*# '''Extrinsic cells'''
−
<br />
+
*#* Located perivascularly and in the meninges.  
−
<br />
+
*# '''Intrinsic cells'''
−
===Astrocyte Functions===
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*#* Located within the substance of the neural tissue.
−
Where astocytes are involved in repair within the brain they take over from fibroblasts to play an important role in abscess formation, as it is difficult to induce fibrosis within the CNS. It is due to this role that astrocytes are the only differentiated CNS cell that retains the ability to divide.
 
−
<br />
 
−
<br />
 
−
Within the metabolic role that astrocytes play within the brain, it has been found that they may control glucose levels and provide lactate for neurone for energy production. The lactate is believed to be produced via the lactate shuttle mechanism. Astrocytes also have high levels of antioxidants, such as glutathione peroxidase which protects the neurons from damage by reactive oxygen species.
 
−
<br />
 
−
<br />
 
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Perhaps one of the most important roles that is undertaken by astrocytes is that of the '''blood brain barrier (BBB)'''. It was originally thought that the astrocyte "end-feet" that encircle adjacent endothelial cells within the BBB aided in the maintenance of the blood-brain barrier. However, the tight junctions and basal lamina of the endothelial cells are now considered to play a more substantial role in maintaining the barrier and within this astrocytes play a key role.
 
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<br />
 
−
<br />
 
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Some classifications of neurotransmitters, namely [[Neurotransmitters_-_Anatomy_%26_Physiology#Small-molecule_Neurotransmitters|small-molecule neurotransmitters]] are reabsorbed and subsequently recycled. Astrocytes represent an element of the system that enables this neurotransmitter re-uptake. Astrocytes are able to effectively isolate synapses and control their potassium levels as well as expressing several types of transporter for the numerous types of neurotransmitters, including glutamate, ATP and GABA. Glutamate is particularly well taken up, and converted to glutamine using glutamine synthase. The role played in controlling potassium ion concentrations within the extracellular spaces involves astrocytes releasing K<sup>+</sup>  when neurons are active, increasing the local extracellular K<sup>+</sup> concentration. Astrocytes have a high density of K<sup>+</sup> channels, allowing them to rapidly clear the excess accumulation in the extracellular space. If this function fails extracellular K<sup>+</sup> concentration rises leading to neuronal depolarisation, and may be a cause of epilepsy.
 
−
<br />
 
−
<br />
 
−
Astrocytes may serve as intermediaries in the neuronal regulation of blood flow by promoting the myelinating activity of oligodendrocytes. Neuronal electrical activity releases ATP, which has been shown to stimulate myelin formation. This stimulation occurs via astrocytes, which secrete LIM in response to the ATP. LIM is a regulatory protein that promotes the myelinating activity of oligodendrocytes.
 
−
<br />
 
−
<br />
 
−
Astrocytes may play a crucial rule in controlling inflammation in the CNS by secreting chemokines. Astrocytes have also been shown to be able to restrict the entry of inflammatory cells to the CNS. When astrocytes are lost or destroyed, the surrounding neurons become very vulnerable in the face of inflammation.
 
  
−
==Microglial Cells==
+
Function:
−
[[Image:microglia.jpg|thumb|right|250px|Microglia]]
+
[[Image:microglia.jpg|thumb|right|150px|Microglia (immunohistochemical staining.) Grzegorz Wicher, '''WikiMedia Commons''']]
−
Microglia are the CNS version of a mononuclear phagocyte system as the normal phagocytic white blood cells are unable to cross the blood-brain barrier. Microglial cells are derived from [[Bone Marrow - Anatomy & Physiology|bone marrow]] and are first seen in the brain just prior to the vascularistion stage of brain development. There are two populations of microglial cells; extrinsic cells which are located perivascularly and in the meninges, and intrinsic cells which are located within the substance of the neural tissue.
+
* Microglia act as macrophages.
−
<br />
+
** They rapidly respond to any type of injury.  
−
<br />
+
* However, microglia differ from other representatives of mononuclear phagocyte system by their low expression of MHC-2.
−
The microglia effectively act as macrophages where they rapidly respond to any type of injury. However, microglia differ from other representatives of mononuclear phagocyte system by their low expression of MHC-2. When undertaking a phagocytic response, MHC-2 can be easily up-regulated. Each type of microglia has a different turnover rate; perivascular and meningeal microglia turnover fairly rapidly whilst intrinsic microglia have a very slow turnover.
+
** MHC-2 is easily up-regulated though.
 +
* The different types of microglia have different turnover rates.
 +
** Perivascular and meningeal microglia turnover fairly rapidly.
 +
** Intrinsic microglia have a very slow turnover.
  
 
==Ependymal Cells==
 
==Ependymal Cells==
−
[[Image:Forkingaqueduct.jpg|thumb|right|250px|Forking aqueduct, showing ependymal cells. Image courtesy of BioMed Archive]]
+
 
−
Ependymal cells make up the epithelial membrane which is the lining of the ventricular system within the brain and spinal cord. Ependymal cells are are involved in the production of [[Cerebral_Spinal_Fluid_-_Anatomy_%26_Physiology|cerebrospinal fluid]]. The cells line the CSF-filled ventricles in the brain and the central canal of the spinal cord, and form part of the [[Cerebral_Spinal_Fluid_-_Anatomy_%26_Physiology|choroid plexus]]. The apical surfaces of ependymal cells are covered with
+
* Ependymal cells make up the is the epithelial membrane lining the ventricular system of the brain and the spinal cord.
−
a layer of cilia and microvilli.
+
* They are involved in the production of cerebrospinal fluid
−
<br />
+
 
−
<br />
+
Structure:
−
Within the ventricles, modified ependymal cells and capillaries form a system called the choroid plexus which produces CSF. The ependymal cells beat thier cilia to circulate CSF around the central nervous system and the microvilli absorb CSF.
+
[[Image:Forkingaqueduct.jpg|thumb|right|150px|Forking aqueduct, showing ependymal cells. Image courtesy of BioMed Archive]]
 +
* Ependymal cells line the CSF-filled ventricles in the brain and the central canal of the spinal cord, and form part of the choroid plexus.  
 +
* The cells are cuboidal/columnar.
 +
* The apical surfaces of ependymal cells are covered with:
 +
** A layer of cilia.  
 +
** Microvilli.
 +
 
 +
Function:
 +
 
 +
* Within the ventricles, modified ependymal cells and capillaries form a system called the choroid plexus.
 +
** The choroid plexus which produces the CSF.
 +
* Ependymal cells beat thier cilia to circulate CSF around the central nervous system.
 +
* The microvilli absorb CSF.
  
 
==Endothelial Cells and the [[Blood Brain Barrier - Anatomy & Physiology|Blood Brain Barrier]]==
 
==Endothelial Cells and the [[Blood Brain Barrier - Anatomy & Physiology|Blood Brain Barrier]]==
−
The walls of capillaries throughout the body and within the CNS are formed by endothelial cells, however, within the CNS the capillary endothelial cells are very tightly packed. This density of cells restricts the passage of substances out of the bloodstream much more than endothelial cells in capillaries elsewhere in the body. This is the basis of the [[Blood Brain Barrier - Anatomy & Physiology|blood brain barrier]].
+
 
−
==Neuronal Transport Mechanisms==
+
* The walls of capillaries in the CNS are formed by endothelial cells, as for capillaries elsewhere in the body.
−
Neurons are often quite large when compared to other cell types within the body and therefore the transport of molecules, organelles and other elements important to the cell becomes more difficult and equally more important. Neurons have developed a number of mechanisms to transport material throughout the cell.
+
* However, in the CNS capillary endothelial cells are very tightly packed.
−
===Anterograde Transport===
+
** This density of cells restricts passage of substances out of the bloodstream much more than endothelial cells in capillaries elsewhere in the body.
−
Anterograde transport facilitates the movement of substances in the direction of the cell body to the axon. This type of neuronal transport has been covered in detail within the PNS Structure and Anatomy page, please click [[PNS_Structure_-_Anatomy_%26_Physiology#Anterograde_Transport|here]].
+
*** This is the basis of the [[Blood Brain Barrier - Anatomy & Physiology|blood brain barrier]]
−
===Retrograde Transport===
+
 
−
Retrograde transport is the opposite of anterograde in that it returns materials from the axon terminal to the cell body, either for degradation or restoration and reuse. This type of neuronal transport has been covered in detail within the PNS Structure and Anatomy page, please click [[PNS_Structure_-_Anatomy_%26_Physiology#Retrograde_Transport|here]].
+
Structure and Function:
−
==Neuron Blood Supply==
+
[[Image:WIKIVETperipheralnervestructure.jpg|thumb|right|150px|Peripheral Nerve Structure © Sophie Stenner, RVC, 2008]]
−
The blood supply to nerves is called the '''vasa nervorum'''. The blood supply to neurons has been covered in detail within the PNS Structure and Anatomy, please click [[PNS_Structure_-_Anatomy_%26_Physiology#Blood_Supply|here]].
+
*Nerve fibres reside in a connective tissue matrix called the '''endoneurium''' and are gathered together into bundles or fascicles defined by a second connective tissue layer called the '''perineurium'''.
 +
*Groups of fascicles are then gathered together in a third connective tissue layer called the '''epineurium'''.
 +
*Thus, peripheral nerves have a '''three-tiered hierarchical arrangement of connective tissue'''.
 +
*Renaut bodies are loose, cigar-shaped whorls of extracellular matrix within fascicles. They are common in [[Equine Alimentary System  - Anatomy & Physiology|horse]] nerves and may also occur in human and rat peripheral nerves at points of stress or compression.
 +
 
 +
==Nerve Fibre==
 +
*The nerve fibre consists of the impulse-carrying axon, which is surrounded by an ensheathing cell, the [[#The Schwann cell|Schwann cell]], which in turn is surrounded by an acellular basal lamina that is continuous along the length of the nerve.
 +
*Nerve fibres come in various discrete diameter groups, which are reflected in their conduction velocities.
 +
*The larger the diameter the more rapid the rate of impulse conduction.
 +
*Particular targets or receptors are associated with axons of a particular diameter, for example:
 +
**Those connected to muscles spindles have a large diameter (20 um) and conduct at 120 m/s
 +
**The smallest myelinated fibres are about 1um and conduct at around 6 m/s.
 +
**The smallest fibres of all are the unmyelinated fibres (the high-threshold sensory afferents, or C-fibres, and post-ganglionic autonomies) and have a diameter of between 1 and 0.1 um. These fibres do not conduct by saltatory conduction and have very slow conduction rates of around 0.5 m/s.
 +
 
 +
===The Axon===
 +
*An outer membrane called the axolemma,
 +
*Within this there is the axoplasm which is continuous with the cytoplasm of the neuron.
 +
*There are no ribosomes, either free or attached to endoplasmic reticulum in axons and therefore, no protein synthesis.
 +
*''Protein synthesis takes place within the cell body and some dendrites.''
 +
**All protein replacement required for the maintenance of the axon depends on proteins being imported from the cell body.
 +
*A critical feature of the axon is its '''cytoskeleton''', which consists of two key elements:
 +
#'''''The Neurofilaments'''''
 +
#*Neurofilaments are intermediate filaments of about 10 nm diameter, and belong to the same class as other cytoskeletal proteins such as keratin, desmin, vimentin, or GFAP of astrocytes.
 +
#*Neurofilaments are formed from a triplet of polypeptide subunits of heavy (~ 200 kD), medium (~ 150 kD) and low (~ 60 kD) molecular weights.
 +
#*Typically, these subunits are heavily phosphorylated and are more numerous than microtubules, especially in large diameter axons, having a pivotal role in determining axon diameter.
 +
#*They are formed in the cell body, transported down the axon by axoplasmic transport and degraded in the terminals by Ca<sup>2+</sup> activated proteases.
 +
#*In other words, there is a constant turnover of neurofllament within the healthy axon.
 +
#'''''The microtubules'''''
 +
#*Micro tubules within axons are similar to microtubules elsewhere, consisting of polymerised dimers of alpha and beta tubulin arranged as a hollow tube of about 28 nm.
 +
#*They are relatively abundant in smaller diameter axons, and are also synthesised in the cell body.
 +
#*An important component of the cytoskeleton are the '''''microtubule associated proteins''''' or MAP's and the tau protein.
 +
#**These proteins are important in microtubule assembly and stability.
 +
#**Different classes of MAP's occur in the dendrites and the axons, and to some extent account for the different ultrastructural features that distinguish these two types of neuronal process.
 +
#**They form cross links between adjacent microtubules but also connect to neurofilaments and actin microfilaments, implying complex interactions between the various components of the axon skeleton.
 +
 
 +
 
 +
===The Schwann Cell===
 +
*Myelination in the PNS is achieved by the Schwann cell, a derivative of neural crest cells, which bud off from the neuroepithelium at a very early stage of neurogenesis.
 +
*During development, Schwann cells engage many small axons.
 +
*As axonal diameter increases, Schwann cells eventually relate with only a single axon c.f [[Neurons - Anatomy & Physiology#Oligodendrocytes|oligodendrocytes]].
 +
**This single axon is enveloped in a trough by the Schwann cell processes that engulf it.
 +
**As the processes come together, an inner mesaxon is formed.
 +
**The leading-edge process continues to move over the axon forming a spiral.
 +
*Myelination, an extremely complex molecular process, occurs when the cytoplasm within the process is extruded allowing the internal surfaces of the membrane to come together as the '''major dense line''', the outer membrane apposition constituting the intraperiod line.
 +
*The alternating pattern of these two form the lamellae of compacted myelin.
 +
*The myelin sheath is attached to, and is an integral part of, the Schwann cell on which it is dependent for its maintenance.
 +
 
 +
*A single Schwann cell forms a single myelin sheath or internode.
 +
*There is a reasonably constant relationship between the myelin thickness and the internodal length, which in turn is associated with axon calibre.
 +
*''Large axons have long, thick myelin sheaths'', and therefore also conduct more rapidly.
 +
*The internodes do not abut one another but are separated by an exposed area of axon called the '''node of Ranvier'''.
 +
*If the axons remain of small diameter, then a Schwann cell will continue to associate with many axons, although none of them are myelinated.
 +
**Thus, ''even unmyelinated axons retain a Schwann cell ensheathment''.  
 +
**These non-myelinating Schwann cells are sometimes referred to as ''Remak cells.''
 +
 
 +
===Axoplasmic Transport===
 +
*Neurons are very large cells and most of a neurons cytoplasm is present in its processes while most of the cells RNA is located in cell body (Nissl substance).
 +
*These cells have therefore evolved mechanisms to transport large macromolecules and organelles up and down processes.
 +
 
 +
====Anterograde Transport====
 +
*Two basic forms of anterograde transport can be recognised:
 +
'''''Fast anterograde transport'''''
 +
*Transports all membranous organelles such as synaptic vesicles
 +
*Occurs at a rate of around 400mm/day (recent evidence suggests that there are many form of fast anterograde transport, mediated by different kinesins).
 +
*Fast anterograde transport depends critically on oxidative metabolism, and is, in fact, independent of the cell body.
 +
*The "motor" molecule is an ATPase called '''kinesin'''
 +
 
 +
 
 +
'''''Slow anterograde transport'''''
 +
*Deals with cytoskeletal elements and large soluble proteins.
 +
*Slow anterograde transport can be further sub-divided into:
 +
**A slow component, which occurs at about 2mm/day (neurofilament, rubulin, actin)
 +
**A fast component, which occurs at around 4 mm/day, transporting all other proteins (eg myosin, clathrin).
 +
 
 +
====Retrograde Transport====
 +
 
 +
*Retrograde transport returns materials from the axon terminal to the cell body, either for degradation or restoration and reuse.  
 +
*As with fast anterograde transport, particles move along microtubules.  
 +
*The motor molecule for retrograde transport is '''''dynein''''' which is a microtubule-associated ATPase.
 +
*The retrograde transport system is important not only for returning material to the cell body, but also provides the means whereby target-derived trophic factors, such as nerve growth factor (NGF) for dorsal root ganglion neurons, are conveyed to the cell body where they promote cell survival.
  
  
−
{{OpenPages}}
 
 
[[Category:Nervous System - Anatomy & Physiology]]
 
[[Category:Nervous System - Anatomy & Physiology]]
−
[[Category:To Do - Review]]
 

Revision as of 20:57, 24 November 2010

Introduction

  • There are two major cell types within the CNS:
  1. Neurons
    • These are the "functional" cells of the central nervous system.
  2. Glial cells:

Neurons

  • Neurons are the functional cells of both the CNS and PNS.
    • They are highly specialised to process and transmit signals.

Structure:

Soma

  • The soma, or cell body is the central part of the neuron.
  • The soma contains:
    • The nucleus
    • Smooth and rough endoplasmic reticulum
    • Ribosomes
    • Golgi apparatus
    • Due to the organelles it contains, the soma is where most protein synthesis occurs.

Dendrites

  • The neuron's dendrites are branching extensions of the cell body.
    • Collectively, branching structure of the dendrites is known as the "dendritic tree".
  • The dendritic tree is the site where input to the neuron occurs.
    Diagram of a neuron. LadyofHats, Wikimedia Commons, 2007
  • Information outflow can also occur in the reverse direction, from the soma to the dendrites.
    • However, information is not transmitted from the dendrites in this way, because axons do not possess chemoreceptors and dendrites cannot secrete neurotransmitter chemicals.
      • Nerve impulses are therefore conducted in one direction only.

Axon

  • The axon is a very fine projection that can measure up to thousands of time the soma diameter in length.
  • The axon carries nerve signals away from the soma.
  • Most neurons have only one axon.
    • However, this will usually undergo extensive branching to communicate with many target cells.
  • The axon emerged from the soma at the axon hillock
    • The axon hillock has the highest density of voltage-dependent Na+channels, making it the most easily-excited part of the neuron.
    • The axon and axon hillock are generally involved in information outflow.
      • However, this region can also receive input from other neurons.
  • At the end of the axon, there are synapses.
    • Neurotransmitter chemicals are released here in order to communicate with target neurons.
  • The axon is dependent on the cell body for all its metabolic functions.
    • The axon has no Nissl substance: it cannot make its own protein, and so the soma must undertake this function.
    • The soma must also dispose of the axon's waste.
    • Death or damage to the cell body will therefore result in axon compromise or degeneration

Function:

  • Neurons are the functional cells of both the CNS and PNS.
    • They are highly specialised to process and transmit signals.
  • Neurons in different parts of the nervous system perform different functions.
    • There is therfore variation in the size, shape, structure, metabolism, function shape and electrochemical properties between locations.

Astrocytes

  • Astrocytes, or astroglia, are star-shaped glial cells in the brain.
  • Astrocytes have several functions:
    • Biochemical support of the endothelial cells forming the blood-brain barrier
    • Provision of nutrients to nervous tissue
    • Repair and scarring in the brain

Structure:

Astrocyte (immunofluorescence staining). Image courtesy of Alliance Européenne Dana pour le Cerveau (EDAB)
  • Astrocytes are "star-shaped".
    • Their many processes envelope synapses made by neurons.
  • Astrocyte expression of glial fibrillary acidic protein (GFAP) allows their identification histologically.

Function:

  • Structural
    • Astrocytes are involved in the physical structuring of the brain:
      • Interplay between astrocytes and mesenchymal cells is important in sculpting sulci in development.
      • Astrocytes delineate the CNS by creating the CNS-PNS interface.
      • The cells cover the surface of the brain.
        • In this situation, the astrocytes' surface is coated by a basal lamina.
      • Astrocytes take over from fibroblasts to play an important role in abscess formation, as it is difficult to induce fibrosis in the CNS.
      • Astrocytes are the only differentiated CNS cell that retains the ability to divide.
  • Metabolic
    • Astrocytes may control glucose levels and provide lactate for neurone for energy production
      • This is by the lactate shuttle mechanism.
    • Astrocytes have high levels of antioxidants, such as glutathione peroxidase.
      • This protects neurons from damage by reactive oxygen species.
  • Blood brain barrier
    • It was originally thought that the astrocyte end-feet encircling endothelial cells aided in the maintenance of the blood-brain barrier.
      • The tight junctions and basal lamina of the endothelial cells are now considered to play a more substantial role in maintaining the barrier.
  • Transmitter reuptake
    • Astrocytes:
      1. Isolate synapses
      2. Control potassium levels
      3. Take up transmitters
        • Astrocytes express transporters for several neurotransmitters, including glutamate, ATP and GABA.
          • Glutamate is particularly well taken up, and converted to glutamine using glutamine synthase.
  • Regulation of ion concentration in the extracellular space
    • Astrocytes release K+ when neurons are active, increasing the local extracellular K+ concentration.
    • Astrocytes have a high density of K+ channels, allowing them to rapidly clear the excess accumulation in the extracellular space.
      • If this function fails extracellular K+ concentration rises. This leads neuronal depolarisation, and may be a cause of epilepsy.
  • Vasomodulation
    • Astrocytes may serve as intermediaries in neuronal regulation of blood flow.
  • Promotion of the myelinating activity of oligodendrocytes
    • Neuronal electrical activity releases ATP, which stimulates myelin formation.
    • This stimulation occurs via astrocytes, which secrete LIM in response to the ATP.
      • LIM is a regulatory protein that promotes the myelinating activity of oligodendrocytes.
  • Control of inflammation
    • Astrocytes may play a crucial rule in controlling inflammation in the CNS, by:
      • Secreting chemokines
      • Playing an important role in the entry of inflammatory cells to the CNS.
    • When astrocytes are lost or destroyed, the neurons become very vulnerable in the face of inflammation.

Oligodendrocytes

Structure:

Oligodendrocyte (immunofluorescence staining), Jurjen Broeke, WikiMedia Commons
  • "Oligodendrocyte" literally means a "cell with many branches".
  • Oligodendrocyte precursors are found throughout the CNS.
    • These precursors are post-mitotic, meaning oligodendrocytes can be replaced.

Function:

  • Oligodendrocytes can myelinate up to 40 axons at a time.
    • The myelin sheath functions to:
      1. Decrease ion leakage and lower the capacitance of the cell membrane.
      2. Increase impulse speed by allowing saltatory conduction of action potentials between the nodes of Ranvier.
        • Nodes of Ranvier are spaces along the axon in between oligodendrocytes.

Microglial Cells

Structure:

  • Microglia are the CNS representatives of mononuclear phagocyte system.
    • As such, they are derived from bone marrow, and are first seen in brain just before vascularistion.
  • There are two populations of microglial cells:
    1. Extrinsic cells
      • Located perivascularly and in the meninges.
    2. Intrinsic cells
      • Located within the substance of the neural tissue.

Function:

Microglia (immunohistochemical staining.) Grzegorz Wicher, WikiMedia Commons
  • Microglia act as macrophages.
    • They rapidly respond to any type of injury.
  • However, microglia differ from other representatives of mononuclear phagocyte system by their low expression of MHC-2.
    • MHC-2 is easily up-regulated though.
  • The different types of microglia have different turnover rates.
    • Perivascular and meningeal microglia turnover fairly rapidly.
    • Intrinsic microglia have a very slow turnover.

Ependymal Cells

  • Ependymal cells make up the is the epithelial membrane lining the ventricular system of the brain and the spinal cord.
  • They are involved in the production of cerebrospinal fluid

Structure:

Forking aqueduct, showing ependymal cells. Image courtesy of BioMed Archive
  • Ependymal cells line the CSF-filled ventricles in the brain and the central canal of the spinal cord, and form part of the choroid plexus.
  • The cells are cuboidal/columnar.
  • The apical surfaces of ependymal cells are covered with:
    • A layer of cilia.
    • Microvilli.

Function:

  • Within the ventricles, modified ependymal cells and capillaries form a system called the choroid plexus.
    • The choroid plexus which produces the CSF.
  • Ependymal cells beat thier cilia to circulate CSF around the central nervous system.
  • The microvilli absorb CSF.

Endothelial Cells and the Blood Brain Barrier

  • The walls of capillaries in the CNS are formed by endothelial cells, as for capillaries elsewhere in the body.
  • However, in the CNS capillary endothelial cells are very tightly packed.
    • This density of cells restricts passage of substances out of the bloodstream much more than endothelial cells in capillaries elsewhere in the body.

Structure and Function:

Peripheral Nerve Structure © Sophie Stenner, RVC, 2008
  • Nerve fibres reside in a connective tissue matrix called the endoneurium and are gathered together into bundles or fascicles defined by a second connective tissue layer called the perineurium.
  • Groups of fascicles are then gathered together in a third connective tissue layer called the epineurium.
  • Thus, peripheral nerves have a three-tiered hierarchical arrangement of connective tissue.
  • Renaut bodies are loose, cigar-shaped whorls of extracellular matrix within fascicles. They are common in horse nerves and may also occur in human and rat peripheral nerves at points of stress or compression.

Nerve Fibre

  • The nerve fibre consists of the impulse-carrying axon, which is surrounded by an ensheathing cell, the Schwann cell, which in turn is surrounded by an acellular basal lamina that is continuous along the length of the nerve.
  • Nerve fibres come in various discrete diameter groups, which are reflected in their conduction velocities.
  • The larger the diameter the more rapid the rate of impulse conduction.
  • Particular targets or receptors are associated with axons of a particular diameter, for example:
    • Those connected to muscles spindles have a large diameter (20 um) and conduct at 120 m/s
    • The smallest myelinated fibres are about 1um and conduct at around 6 m/s.
    • The smallest fibres of all are the unmyelinated fibres (the high-threshold sensory afferents, or C-fibres, and post-ganglionic autonomies) and have a diameter of between 1 and 0.1 um. These fibres do not conduct by saltatory conduction and have very slow conduction rates of around 0.5 m/s.

The Axon

  • An outer membrane called the axolemma,
  • Within this there is the axoplasm which is continuous with the cytoplasm of the neuron.
  • There are no ribosomes, either free or attached to endoplasmic reticulum in axons and therefore, no protein synthesis.
  • Protein synthesis takes place within the cell body and some dendrites.
    • All protein replacement required for the maintenance of the axon depends on proteins being imported from the cell body.
  • A critical feature of the axon is its cytoskeleton, which consists of two key elements:
  1. The Neurofilaments
    • Neurofilaments are intermediate filaments of about 10 nm diameter, and belong to the same class as other cytoskeletal proteins such as keratin, desmin, vimentin, or GFAP of astrocytes.
    • Neurofilaments are formed from a triplet of polypeptide subunits of heavy (~ 200 kD), medium (~ 150 kD) and low (~ 60 kD) molecular weights.
    • Typically, these subunits are heavily phosphorylated and are more numerous than microtubules, especially in large diameter axons, having a pivotal role in determining axon diameter.
    • They are formed in the cell body, transported down the axon by axoplasmic transport and degraded in the terminals by Ca2+ activated proteases.
    • In other words, there is a constant turnover of neurofllament within the healthy axon.
  2. The microtubules
    • Micro tubules within axons are similar to microtubules elsewhere, consisting of polymerised dimers of alpha and beta tubulin arranged as a hollow tube of about 28 nm.
    • They are relatively abundant in smaller diameter axons, and are also synthesised in the cell body.
    • An important component of the cytoskeleton are the microtubule associated proteins or MAP's and the tau protein.
      • These proteins are important in microtubule assembly and stability.
      • Different classes of MAP's occur in the dendrites and the axons, and to some extent account for the different ultrastructural features that distinguish these two types of neuronal process.
      • They form cross links between adjacent microtubules but also connect to neurofilaments and actin microfilaments, implying complex interactions between the various components of the axon skeleton.


The Schwann Cell

  • Myelination in the PNS is achieved by the Schwann cell, a derivative of neural crest cells, which bud off from the neuroepithelium at a very early stage of neurogenesis.
  • During development, Schwann cells engage many small axons.
  • As axonal diameter increases, Schwann cells eventually relate with only a single axon c.f oligodendrocytes.
    • This single axon is enveloped in a trough by the Schwann cell processes that engulf it.
    • As the processes come together, an inner mesaxon is formed.
    • The leading-edge process continues to move over the axon forming a spiral.
  • Myelination, an extremely complex molecular process, occurs when the cytoplasm within the process is extruded allowing the internal surfaces of the membrane to come together as the major dense line, the outer membrane apposition constituting the intraperiod line.
  • The alternating pattern of these two form the lamellae of compacted myelin.
  • The myelin sheath is attached to, and is an integral part of, the Schwann cell on which it is dependent for its maintenance.
  • A single Schwann cell forms a single myelin sheath or internode.
  • There is a reasonably constant relationship between the myelin thickness and the internodal length, which in turn is associated with axon calibre.
  • Large axons have long, thick myelin sheaths, and therefore also conduct more rapidly.
  • The internodes do not abut one another but are separated by an exposed area of axon called the node of Ranvier.
  • If the axons remain of small diameter, then a Schwann cell will continue to associate with many axons, although none of them are myelinated.
    • Thus, even unmyelinated axons retain a Schwann cell ensheathment.
    • These non-myelinating Schwann cells are sometimes referred to as Remak cells.

Axoplasmic Transport

  • Neurons are very large cells and most of a neurons cytoplasm is present in its processes while most of the cells RNA is located in cell body (Nissl substance).
  • These cells have therefore evolved mechanisms to transport large macromolecules and organelles up and down processes.

Anterograde Transport

  • Two basic forms of anterograde transport can be recognised:

Fast anterograde transport

  • Transports all membranous organelles such as synaptic vesicles
  • Occurs at a rate of around 400mm/day (recent evidence suggests that there are many form of fast anterograde transport, mediated by different kinesins).
  • Fast anterograde transport depends critically on oxidative metabolism, and is, in fact, independent of the cell body.
  • The "motor" molecule is an ATPase called kinesin


Slow anterograde transport

  • Deals with cytoskeletal elements and large soluble proteins.
  • Slow anterograde transport can be further sub-divided into:
    • A slow component, which occurs at about 2mm/day (neurofilament, rubulin, actin)
    • A fast component, which occurs at around 4 mm/day, transporting all other proteins (eg myosin, clathrin).

Retrograde Transport

  • Retrograde transport returns materials from the axon terminal to the cell body, either for degradation or restoration and reuse.
  • As with fast anterograde transport, particles move along microtubules.
  • The motor molecule for retrograde transport is dynein which is a microtubule-associated ATPase.
  • The retrograde transport system is important not only for returning material to the cell body, but also provides the means whereby target-derived trophic factors, such as nerve growth factor (NGF) for dorsal root ganglion neurons, are conveyed to the cell body where they promote cell survival.
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