This document provides information about the structure and function of neurons. It begins by stating the learning objectives, which are to define neurons and their processes, identify different neuron types, review neuron cell biology and functions, and explain transport within neurons and the role of synapses. It then defines neurons and their key structures like the cell body, dendrites, and axon. The rest of the document describes the internal structures of the neuron in detail and classifies neurons based on their morphology and size.
This document provides information about the structure and function of neurons. It begins by stating the learning objectives, which are to define neurons and their processes, identify different neuron types, review neuron cell biology and functions, and explain transport within neurons and the role of synapses. It then defines neurons and their key structures like the cell body, dendrites, and axon. The rest of the document describes the internal structures of the neuron in detail and classifies neurons based on their morphology and size.
This document provides information about the structure and function of neurons. It begins by stating the learning objectives, which are to define neurons and their processes, identify different neuron types, review neuron cell biology and functions, and explain transport within neurons and the role of synapses. It then defines neurons and their key structures like the cell body, dendrites, and axon. The rest of the document describes the internal structures of the neuron in detail and classifies neurons based on their morphology and size.
This document provides information about the structure and function of neurons. It begins by stating the learning objectives, which are to define neurons and their processes, identify different neuron types, review neuron cell biology and functions, and explain transport within neurons and the role of synapses. It then defines neurons and their key structures like the cell body, dendrites, and axon. The rest of the document describes the internal structures of the neuron in detail and classifies neurons based on their morphology and size.
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By:
Dr. Maria Simplicia E. Flores
At the end of this lesson, the student should be able to: 1. To define the neuron and name its processes 2. To learn the varieties of neurons and identify them in different parts of the nervous system 3. To review the cell biology of a neuron and understand the function of a nerve cell and its processes 4. To review the structure of the plasma membrane as it is related to its physiology 5. To learn the transport of materials from the cell body to the axon terminals 6. To understand the structure and function of synapses and neurotransmitters 7. To review the supporting function of the neuroglial cells for nerve cells and the possible role that they play in neuronal metabolism, function, and neuronal death Neuron - nerve cell and all its processes. Excitable cells that are specialized for the reception of stimuli and the conduction of the nerve impulse. Varies in size and shape, one or more neurites project from each cell body ◦ Neurites responsible for receiving information and conducting it toward the cell body are dendrites. ◦ The single long tubular neurite that conducts impulses away from the cell body is the axon. Dendrites and axons are commonly referred to as nerve fibers.
Neurons are found in the
brain and spinal cord and in ganglia. Normal neurons in the mature individual do NOT undergo division and replication. Schematic Drawing of a Motor Neuron Cell body of a neuron may be as small as 5 μm or as large as 135 μm in diameter, but the processes or neurites may extend over a distance of more than 1 m. Neurons can be CLASSIFIED MORPHOLOGICALLY based on the NUMBER, LENGTH, and MODE OF BRANCHING OF THEIR NEURITES. 1. UNIPOLAR neurons have a single neurite that divides a short distance from the cell body into two branches, one proceeding to some peripheral structure and the other entering the CNS. The branches of this single neurite have the structural and functional characteristics of an axon. In this type of neuron, the fine terminal branches found at the peripheral end of the axon at the receptor site are often referred to as the dendrites. Example: posterior root ganglion. 2. BIPOLAR neurons have an elongated cell body, with a single neurite emerging from each end. Examples: found in the retinal bipolar cells and the cells of the sensory cochlear and vestibular ganglia. 3. MULTIPOLAR neurons have a number of neurites arising from the cell body. With the exception of the axon, the remainder of the neurites are dendrites. Examples: neurons of the brain and spinal cord Neurons may also be CLASSIFIED ACCORDING TO SIZE. 1. GOLGI TYPE I neurons have a long axon that can stretch 1 m or more in length in extreme cases. The axons of these neurons form the long fiber tracts of the brain and spinal cord and the nerve fibers of peripheral nerves. Examples: pyramidal cells of the cerebral cortex, Purkinje cells of the cerebellar cortex, and the motor cells of the spinal cord 2. GOLGI TYPE II neurons have a short axon that terminates in the neighborhood of the cell body or is entirely absent. They greatly outnumber the Golgi type I neurons. The short dendrites that arise from these neurons give them a star-shaped appearance. Examples: numerous in the cerebral and cerebellar cortex and are often inhibitory in function. Different Types of Neurons A. Neuronal Structure Cell body - consists essentially of a mass of cytoplasm in which a nucleus is embedded, bounded externally by a plasma membrane. 1. Nerve Cell Body The volume of cytoplasm within the nerve cell body is often far less than the total volume of cytoplasm in the neurites.
A. NUCLEUS - stores the genes
In mature neurons, the chromosomes NO longer duplicate themselves and function only in gene expression. Therefore, the chromosomes exist in an uncoiled state. The large size of the nucleolus probably is due to the high rate of protein synthesis. Barr body – composed of sex chromatin for female chromosome. Nuclear envelope is continuous with the cytoplasmic RER. ◦ The envelope is double layered and possesses fine nuclear pores, through which materials can diffuse into and out of the nucleus. Newly formed ribosomal subunits can be passed into the cytoplasm through the nuclear pores. B. CYTOPLASM - rich in RER and SER, and contains the following organelles and inclusions: (a) Nissl substance (b) Golgi complex (c) mitochondria (d) microfilaments (e) microtubules (f) lysosomes (g) centrioles (h) lipofuscin, melanin, glycogen, and lipid Nissl substance – basophilic in color, responsible for protein synthesis, composed of RER (ribosomes contain RNA) arranged in the form of broad cisternae stacked one on top of the other. Consists of granules that are distributed throughout the cytoplasm of the cell body, except for the region close to the axon hillock. The granular material also extends into the proximal parts of the dendrites but is NOT present in the axon. Fatigue or neuronal damage causes the Nissl substance to move and become concentrated at the periphery of the cytoplasm. The phenomenon which gives the impression that the Nissl substance has disappeared is called chromatolysis. Golgi complex – appears as network of irregular wavy threads around the nucleus; appears as clusters of flattened cisternae and small vesicles made up of SER The protein produced by the Nissl substance is transferred to the inside of the Golgi complex in transport vesicles, where it is temporarily stored and where carbohydrate may be added to the protein to form glycoproteins. Each cisterna of the Golgi complex is specialized for different types of enzymatic reaction. The Golgi complex is also active in lysosome production and in the synthesis of cell membranes. Mitochondria - found scattered throughout the cell body, dendrites, and axons Spherical or rod shaped Walls are double membrane; inner membrane is thrown into folds or cristae It possess many enzymes which take part in the tricarboxylic acid cycle and the cytochrome chains of respiration. Important in the production of energy. Neurofibrils - are numerous and run parallel to each other through the cell body into the neurites. Resolve into bundles of neurofilaments
Neurofilaments - form the main component
of the cytoskeleton. Chemically, neurofilaments are very stable and belong to the cytokeratin family. Microfilaments - measure about 3 to 5 nm in diameter and are formed of actin. Concentrated at the periphery of the cytoplasm just beneath the plasma membrane where they form a dense network. Together with microtubules, these play a key role in the formation of new cell processes and the retraction of old ones. These also assist the microtubules in axon transport. Microtubules - measures about 25 nm in diameter; are found interspersed among the neurofilaments Extends throughout the cell body and its processes.
Microtubules and microfilaments provide a
stationary track that permits specific organelles to move (stop-and-start movement) by molecular motors. Cell transport - involves the movement of membrane organelles, secretory material, synaptic precursor membranes, large dense core vesicles, mitochondria, and SER. Can take place in both directions, in the cell body and its processes. RAPID TRANSPORT (100 to 400 mm/day) is mediated by two motor proteins associated with the microtubule ATPase sites. SLOW TRANSPORT (0.1 to 3.0 mm/day) involves the bulk movement of the cytoplasm, and includes the movement of mitochondria and other organelles RAPID TRANSPORT In anterograde (away from the cell) movement, kinesin-coated organelles are thought to move toward one end of the tubule In retrograde (toward the cell) movement, dynein-coated organelles are thought to move toward the other end of the tubule. The direction and speed of the movement of an organelle are brought by the activation of one of the motor proteins or both simultaneously. SLOW TRANSPORT Slow axonal transport occurs only in the anterograde direction. Due to one of the kinesin families. Lysosomes - membrane-bound vesicles measuring about 8 nm in diameter. Acting as intracellular scavengers and contain hydrolytic enzymes. Formed by the budding off of the Golgi apparatus. Exist in three forms: (1) primary lysosomes (2) secondary lysosomes - contain partially digested material (myelin figures) (3) residual bodies - the enzymes are inactive and the bodies have evolved from digestible materials such as pigment and lipid. Centrioles - small, paired structures found in immature dividing nerve cells. Each centriole is a hollow cylinder whose wall is made up of microtubule bundles. These are associated with the formation of the spindle during cell division and in the formation of microtubules. Involved also in the maintenance of microtubules in mature nerve cells. Lipofuscin - pigment material; occurs as yellowish brown granules within the cytoplasm. Forms as the result of lysosomal activity It represents a harmless metabolic by product. Accumulates with age. Melanin granules - are found in the cytoplasm of cells in certain parts of the brain (e.g., substantia nigra of the midbrain). Their presence may be related to the catecholamine-synthesizing ability of these neurons, whose neurotransmitter is dopamine. C. PLASMA MEMBRANE - forms the continuous external boundary of the cell body and its processes. Site for the initiation and conduction of the nerve impulse. It is about 8 nm thick; appears as two dark lines with a light line between them. Composed of an inner and an outer layer of very loosely arranged protein molecules, each layer being about 2.5 nm thick, separated by a middle layer of lipid about 3 nm thick. Lipid layer is made up of two rows of phospholipid molecules arranged so that their hydrophobic ends are in contact with each other and their polar ends are in contact with the protein layers. Certain protein molecules lie within the phospholipid layer and span the entire width of the lipid layer ◦ Provide the membrane with hydrophilic channels through which inorganic ions may enter and leave the cell. Carbohydrate molecules are attached to the outside of the plasma membrane and are linked to the proteins or the lipids, forming as cell coat or glycocalyx Plasma membrane and the cell coat together form a semipermeable membrane that allows diffusion of certain ions through it but restricts others. B. Nerve Cell Processes Neurites- the processes of a nerve cell (divided into dendrites and an axon). 1. Dendrites - are the short processes of the cell body which conduct the nerve impulse toward the cell body; its diameter tapers as they extend from the cell body, and often branch profusely. The finer branches bear large numbers of small projections called dendritic spines. Dendrites are extensions of the cell body to increase the surface area for the reception of axons from other neurons. The cytoplasm of the dendrites closely resembles that of the cell body and contains Nissl granules, mitochondria, microtubules, microfilaments, ribosomes, and SER. During early embryonic development, dendrites are overproduced. Later, they are reduced in number and size in response to altered functional demand from afferent axons. Dendrites remain plastic throughout life and elongate and branch or contract in response to afferent activity. 2. Axon - the longest process of the cell body. Arises from a small conical elevation on the cell body called axon hillock (no Nissl granules) It may arise occasionally from the proximal part of a dendrite. Tubular and uniform in diameter; it tends to have a smooth surface. It does not branch close to the cell body; collateral branches occurs along their length; branch profusely before their termination. Terminals – enlarged distal ends of the terminal branches of the axons. Varicosities - series of swellings resembling a string of beads near the termination (especially those of autonomic nerves). Axons may be very short (0.1 mm) as seen in many neurons of the CNS, or extremely long (3.0 m) as seen when they extend from a peripheral receptor in the skin to the spinal cord and thence to the brain. Larger axon diameter conduct impulses rapidly, while those smaller diameter conduct impulses very slowly. Axon always conduct impulses away from the cell body EXCEPT for the axons of the sensory posterior root ganglion because it carries the impulse toward the cell body.
Axolemma - plasma membrane bounding the
axon Axoplasm - cytoplasm of the axon; lacks Nissl granules and Golgi complex; RNA and ribosomes are absent (sites for the production of protein) Axonal survival depends on the transport of substances from the cell bodies.
Initial segment of the axon - first 50 to 100 um
after it leaves the axon hillock; most excitable part of the axon and is the site in which an AP originates. Under normal conditions, an AP does not originate on the plasma membrane. 3. Myelin – covers the axon; consists of multiple concentric layers of lipid-rich membrane produced by glial cells namely: Schwann cells (PNS) and oligodendrocytes (CNS). Nodes of Ranvier - small gaps (1 μm long) in the segments of myelin sheaths (axon) where myelin is absent. Smallest axons are unmyelinated. Function of myelin: insulator Myelination - increases the speed of impulse conduction along the axon. C. Axon Transport Materials are transported from the cell body to the axon terminals (anterograde transport) and to a lesser extent in the opposite direction (retrograde transport). Fast anterograde transport (100 to 400 mm/day) - refers to the transport of proteins and transmitter substances or their precursors. Slow anterograde transport (0.1 to 3.0 mm/day) - refers to the transport of axoplasm and includes the microfilaments and microtubules. Retrograde transport - explains how the cell bodies of nerve cells respond to changes in the distal end of the axons. ◦ Examples: activated growth factor receptors, pinocytotic vesicles arising at the axon terminals, worn-out organelles which goes back to the cell body for breakdown by the lysosomes.
Axon transport is brought about by microtubules
assisted by the microfilaments. D. Synapses Nervous system consists of a large number of neurons that are linked together to form functional conducting pathways. Synapse or synaptic junction- the site where two neurons (or a neuron and a skeletal muscle or gland cell) come into close proximity and functional interneuronal communication occurs. Communication between neurons takes place in one direction only, usually occurs from the axon terminal of the transmitting neuron (presynaptic side) to the receptive region of the receiving neuron (postsynaptic side). Synaptic spines – extensions of the surface of a neuron form receptive sites for synaptic contact with afferent boutons.
Synapses occur in a number of forms:
1. between an axon of one neuron and the dendrite or cell body of the second neuron (most common type) 2. Depending on the site of the synapse: axodendritic, axosomatic, or axoaxonic. Different Types of Chemical Synapses Two types of synapses: chemical and electrical. 1. Chemical synapse- most common, in which a chemical substance called neurotransmitter, passes across the narrow space between the cells and becomes attached to a protein molecule in the postsynaptic membrane called the receptor. o Have several distinctive characteristics: synaptic vesicles on the presynaptic side, a synaptic cleft, and a dense thickening of the cell membrane on both the receiving cell and the presynaptic side. ◦ Presynaptic and postsynaptic membranes – are the apposed surfaces of the terminal axonal expansion and the neuron; separated by a synaptic cleft measuring about 20 to 30 nm wide. ◦ The synaptic cleft contains polysaccharides. ◦ Presynaptic terminal contains many small presynaptic vesicles containing neurotransmitters. ◦ One neurotransmitter is usually the principal activator and acts directly on the postsynaptic membrane, while the other transmitters function as modulators and modify the activity of the principal transmitter. ◦ The vesicles fuse with the presynaptic membrane and discharge the neurotransmitter(s) into the synaptic cleft by a process of exocytosis. ◦ The plasticity of synapses may be of great importance in the process of learning and in the development and maintenance of memory. 2. Electrical synapse– are gap junctions containing channels that extend from the cytoplasm of the presynaptic neuron to that of the postsynaptic neuron. ◦ The neurons communicate electrically which involves current that passes directly from cell to cell through specialized junctions, hence chemical transmitter is not present. ◦ The bridging channels permit ionic current flow to take place from one cell to the other with a minimum of delay. ◦ The rapid spread of activity from one neuron to another ensures that a group of neurons perform an identical function acting together. ◦ Electrical synapses are bidirectional; chemical synapses are not. E. Neurotransmitter Presynaptic vesicles containing neurotransmitter substance and mitochondria which provide ATP for the synthesis of new transmitter substance play a key role in the release of neurotransmitter substances at synapses. Most neurons produce and release only one principal transmitter at all Neurotransmitters: ACh, norepinephrine, epinephrine, dopamine, glycine, serotonin, GABA, enkephalins, substance P, and glutamic acid. ACh - found at the neuromuscular junction of all skeletal muscle, in autonomic ganglia, and at parasympathetic nerve endings.
Norepinephrine - found at sympathetic nerve
endings.
Dopamine - found in high concentration in
different parts of the CNS, such as in the basal nuclei (ganglia). Neuroglia – are nonexcitable cells supporting the neurons of CNS. Generally smaller than neurons and outnumber them by 5 to 10 times; comprise about half the total volume of the brain and spinal cord. FOUR types namely: (1) astrocytes (2) oligodendrocytes (3) microglia (4) ependyma 1. ASTROCYTES - have small cell bodies with branching processes that extend in all directions. Two types: fibrous and protoplasmic ◦ Fibrous astrocytes are found mainly in the white matter, where their processes pass between the nerve fibers ◦ Protoplasmic astrocytes are found mainly in the gray matter, where their processes pass between the nerve cell bodies Functions: 1. form a supporting framework for the nerve cells and nerve fibers 2. for the embryo, this serve as a scaffolding for the migration of immature neurons 3. serve as electrical insulators preventing axon terminals from influencing neighboring and unrelated neurons. 4. barriers for the spread of neurotransmitter substances released at synapses 5. limits the influence of GABA and glutamic acid neurotransmitters secreted by the nerve terminals 6. take up excess K+ ions from the extracellular space 7. store glycogen within their cytoplasm 8. serve as phagocytes by taking up degenerating synaptic axon terminals 9. serve as a conduit for the passage of metabolites or raw materials from blood capillaries to the neurons through their perivascular feet. 10. produce substances that have a trophic influence on neighboring neurons (e.g. cytokines) 11. blood-brain barrier. 2. OLIGODENDROCYTES - have small cell bodies and a few delicate processes; their cytoplasm does not contain filaments. Found in rows along myelinated nerve fibers and surrounds nerve cell bodies Functions: 1. responsible for the formation of the myelin sheath of nerve fibers in the CNS. However, the myelin of peripheral nerves in the PNS is formed from by Schwann cells. provides an insulating coat and greatly increases the speed of nerve conduction 2. Oligodendrocytes unlike Schwann cells, can each form several internodal segments of myelin on the same or different axons ◦ Oligodendrocytes are not surrounded by a basement membrane. ◦ Myelination begins at about the 16th week of intrauterine life and continues postnatally until all the major nerve fibers are myelinated by the time the child is walking. 3. Oligodendrocytes also surround nerve cell bodies (satellite Oligodendrocytes). 4. It influence the biochemical environment of neurons. 3. MICROGLIAL CELLS - embryologically unrelated to the other neuroglial cells; derived from macrophages outside the nervous system. Smallest of the neuroglial cells and are found scattered throughout the CNS Wavy branching processes arise from their small cell bodies that give off numerous spinelike projections. Migrate into the nervous system during fetal life. Microglial cells increases in number in the presence of damaged nervous tissue resulting from trauma and ischemic injury and in the presence of diseases including Alzheimer disease, Parkinson disease, multiple sclerosis, and AIDS. Many of these new cells are monocytes that have migrated from the blood. Functions: 1. Microglial cells in the normal brain and spinal cord appear to be inactive and are sometimes called resting microglial cells. In inflammatory disease of the CNS, they become the immune effector cells. They retract their processes and migrate to the site of the lesion. Here, they proliferate and become antigen-presenting cells, which, together with the invading T lymphocytes, confront invading organisms. 2. Active phagocytic cell; their cytoplasm becomes filled with lipids and cell remnants. The microglial cells are joined by monocytes from neighboring blood vessels. 4. EPENDYMAL CELLS - line the cavities of the brain and the central canal of the spinal cord. They form a single layer of cells that are cuboidal or columnar in shape and possess microvilli and cilia. ◦ The cilia are often motile, and their movements contribute to the flow of the cerebrospinal fluid (CSF). Three groups of Ependymal cells : 1. Ependymocytes - line the ventricles of the brain and the central canal of the spinal cord and are in contact with the CSF. The adjacent surfaces have gap junctions but the CSF is in free communication with CNS intercellular spaces. 2. Tanycytes - line the floor of the third ventricle overlying the median eminence of the hypothalamus. These cells have long basal processes that pass between the cells of the median eminence and place endfeet on blood capillaries. 3. Choroidal epithelial cells - cover the surfaces of the choroid plexuses. The sides and bases of these cells are thrown into folds; near their luminal surfaces, the cells are held together by tight junctions that encircle the cells. The presence of tight junctions prevents the leakage of CSF into the underlying tissues. Functions: 1. Ependymocytes assist in CSF circulation within the cavities of the brain and the central canal of the spinal cord by the movements of the cilia. The microvilli on the free surfaces of the ependymocytes indicate that they also have an absorptive function. 2. Tanycytes are thought to transport chemical substances from the CSF to the hypophyseal portal system. In this manner, they may play a part in the control of the hormone production by the anterior lobe of the pituitary. 3. Choroidal epithelial cells are involved in the production and secretion of CSF from the choroid plexuses. Accounts 20% of the total volume of the brain and spinal cord. Very narrow gap that separates the neurons and the neuroglial cells wherein these gaps are linked together and filled with tissue fluid. It is in direct continuity with the CSF in the subarachnoid space externally and with the CSF in the ventricles of the brain and the central canal of the spinal cord internally. It surrounds the blood capillaries in the brain and spinal cord (CNS does not have lymphatic capillaries). Provides a pathway for the exchange of ions and molecules between the blood and the neurons and glial cells. The plasma membrane of the endothelial cells of most capillaries is impermeable to many chemicals, and this forms the blood—brain barrier. CEREBRAL EDEMA - there is an increase in the bulk of the brain. ◦ It can be either vasogenic (extracellular primarily) or cytotoxic (intracellular primarily). ◦ Because of the limited size of the cranial vault within the skull, cerebral edema must be treated emergently. Wallerian degeneration - if the axon is cut, the part distal to the cut degenerates, because materials for maintaining the axon (mostly proteins) are formed in the cell body and can no longer be transported down the axon (axoplasmic transport). Distal to the level of axonal transection when a peripheral nerve is injured, Schwann cells dedifferentiate and divide. Together with macrophages, they phagocytize the remnants of the myelin sheaths, which lose their integrity as the axon degenerates. Successful axonal regeneration does not commonly occur after injury to the CNS. Many neurons appear to be dependent on connection with appropriate target cells; if the axon fails to regenerate and form a new synaptic connection with the correct postsynaptic cells, the axotomized neuron may die or atrophy. REGENERATION A. Peripheral Nerves Regeneration denotes a nerve’s ability to regrow to an appropriate target, including the reestablishment of functionally useful connections. Nerve crush versus Nerve transection ◦ Crush injury - axons may be severed, but the Schwann cells, surrounding basal lamina, and perineurium maintain continuity through the lesion, facilitating regeneration of axons through the injured nerve. ◦ Nerve transection (nerve is cut) – the continuity of these pathways is disrupted. Regeneration is less likely to occur.
B. Central Nervous System
Axonal regeneration is typically abortive due to glial scar formed by astrocytic processes, properties of the oligodendroglial cells, and inhibitory factor produced by oligodendrocytes, CNS myelin, or both. C. Remyelination In a number of disorders of the PNS (Guillain– Barré syndrome), there is demyelination which interferes with conduction. This condition is often followed by remyelination by Schwann cells, which are capable of elaborating new myelin sheaths. Occurs much more slowly in the CNS. Little remyelination occurs in demyelinated plaques within the brain and spinal cord (Multiple Sclerosis). D. Collateral Sprouting Occurs when an innervated structure has been partially denervated. The remaining axons then form new collaterals that reinnervate the denervated part of the end organ. This kind of regeneration demonstrates that there is considerable plasticity in the nervous system and that one axon can take over the synaptic sites formerly occupied by another. Capability to produce neurons from undifferentiated, proliferative progenitor cells. After pathological insults that result in neuronal death, the number of neurons is permanently reduced.
(Doi 10.1007 - 978!3!662-05204-4) Mirsky, Vladimir M. - (Springer Series On Chemical Sensors and Biosensors) Ultrathin Electrochemical Chemo - and Biosensors Volume 2