Sunday, April 28, 2013
Saturday, April 27, 2013
Degeneration & Regeneration of Nerve Tissue-Histology of Nervous System
Mammalian neurons usually do not divide, and their degeneration represents a permanent loss. Peripheral nerve fibers can regenerate if their perikaryons are not destroyed.
In contrast to nerve cells, neuroglia of the central nervous system—and Schwann cells and ganglionic satellite cells of the peripheral nervous system—are able to divide by mitosis. Spaces in the central nervous system left by nerve cells lost by disease or injury are invaded by neuroglia.
Because nerves are widely distributed throughout the body, they are often injured. When a nerve axon is transected, degenerative changes take place, followed by a reparative phase.
In a wounded nerve fiber, it is important to distinguish the changes occurring in the proximal segment from those in the distal segment. The proximal segment maintains its continuity with the trophic center (perikaryon) and frequently regenerates. The distal segment, separated from the nerve cell body, degenerates.
Axonal injury causes several changes in the perikaryon: chromatolysis, ie, dissolution of Nissl substances with a consequent decrease in cytoplasmic basophilia; an increase in the volume of the perikaryon; and migration of the nucleus to a peripheral position in the perikaryon.
In the nerve stub distal to the injury, both the axon and the myelin sheath degenerate completely, and their remnants, excluding their connective tissue and perineurial sheaths, are removed by macrophages. While these regressive changes take place, Schwann cells proliferate within the remaining connective tissue sleeve, giving rise to solid cellular columns. These rows of Schwann cells serve as guides to the sprouting axons formed during the reparative phase.
After the regressive changes, the proximal segment of the axon grows and branches, forming several filaments that progress in the direction of the columns of Schwann cells. Only fibers that penetrate these Schwann cell columns will continue to grow and reach an effector organ.
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Choroid Plexus & Cerebrospinal Fluid-Histology of Nervous System
The choroid plexus consists of invaginated folds of pia mater, rich in dilated fenestrated capillaries, that penetrate the interior of the brain ventricles. It is found in the roofs of the third and fourth ventricles and in part in the walls of the lateral ventricles.
The choroid plexus is composed of loose connective tissue of the pia mater, covered by a simple cuboidal or low columnar epithelium made of ion-transporting cells .
The main function of the choroid plexus is to elaborate cerebrospinal fluid, which contains only a small amount of solids and completely fills the ventricles, central canal of the spinal cord, subarachnoid space, and perivascular space. Cerebrospinal fluid is important for the metabolism of the central nervous system and acts as a protective device against mechanical shocks.
Cerebrospinal fluid is clear, has a low density (1.004–1.008 g/ mL), and is very low in protein content. A few desquamated cells and two to five lymphocytes per milliliter are also present. Cerebrospinal fluid is continuously produced and circulates through the ventricles, from which it passes into the subarachnoid space. There, arachnoid villi provide the main pathway for absorption of cerebrospinal fluid into the venous circulation.
The Blood-Brain Barrier-Histology of Nervous System
The blood–brain barrier is a functional barrier that prevents the passage of some substances, such as antibiotics and chemical and bacterial toxic matter, from the blood to nerve tissue.
The blood–brain barrier results from the reduced permeability that is characteristic of blood capillaries of nerve tissue. Occluding junctions, which provide continuity between the endothelial cells of these capillaries, represent the main structural component of the barrier. The cytoplasm of these endothelial cells does not have the fenestrations found in many other locations, and very few pinocytotic vesicles are observed. The expansions of neuroglial cell processes that envelop the capillaries (astrocytes) are partly responsible for their low permeability.
Meninges-Histology of Nervous System
The skull and the vertebral column protect the central nervous system. It is also encased in membranes of connective tissue called the meninges . Starting with the outermost layer, the meninges are the dura mater, arachnoid, and pia mater. The arachnoid and the pia mater are linked together and are often considered a single membrane called the pia-arachnoid.
Dura Mater
The dura mater is the external layer and is composed of dense connective tissue continuous with the periosteum of the skull. The dura mater that envelops the spinal cord is separated from the periosteum of the vertebrae by the epidural space, which contains thin-walled veins, loose connective tissue, and adipose tissue. The dura mater is always separated from the arachnoid by the thin subdural space.
Arachnoid
The arachnoid has two components: a layer in contact with the dura mater and a system of trabeculae connecting the layer with the pia mater. The cavities between the trabeculae form the subarachnoid space, which is filled with cerebrospinal fluid and is completely separated from the subdural space. This space forms a hydraulic cushion that protects the central nervous system from trauma. The subarachnoid space communicates with the ventricles of the brain. The arachnoid is composed of connective tissue devoid of blood vessels. In some areas, the arachnoid perforates the dura mater, forming protrusions that terminate in venous sinuses in the dura mater. These protrusions, which are covered by endothelial cells of the veins, are called arachnoid villi. Their function is to reabsorb cerebrospinal fluid into the blood of the venous sinuses.
Pia Mater
The pia mater is a loose connective tissue containing many blood vessels. Although it is located quite close to the nerve tissue, it is not in contact with nerve cells or fibers. Between the pia mater and the neural elements is a thin layer of neuroglial processes, adhering firmly to the pia mater and forming a physical barrier at the periphery of the central nervous system. This barrier separates the central nervous system from the cerebrospinal fluid. The pia mater follows all the irregularities of the surface of the central nervous system and penetrates it to some extent along with the blood vessels. Squamous cells of mesenchymal origin cover pia mater. Blood vessels penetrate the central nervous system through tunnels covered by pia mater—the perivascular spaces. The pia mater disappears before the blood vessels are transformed into capillaries. In the central nervous system, the blood capillaries are completely covered by expansions of the neuroglial cell processes.
The Ganglia-Histology of Nervous System
Ganglia are ovoid structures containing neuronal cell bodies and glial cells supported by connective tissue. Because they serve as relay stations to transmit nerve impulses, one nerve enters and another exits from each ganglion. The direction of the nerve impulse determines whether the ganglion will be a sensory or an autonomic ganglion.
Sensory Ganglia
Sensory ganglia receive afferent impulses that go to the central nervous system. Two types of sensory ganglia exist. Some are associated with cranial nerves (cranial ganglia); others are associated with the dorsal root of the spinal nerves and are called spinal ganglia. A connective tissue framework and capsule support the ganglion cells. The neurons of these ganglia are pseudounipolar and relay information from the ganglion's nerve endings to the gray matter of the spinal cord via synapses with local neurons.
Autonomic Ganglia
Autonomic ganglia appear as bulbous dilatations in autonomic nerves. Some are located within certain organs, especially in the walls of the digestive tract, where they constitute the intramural ganglia. These ganglia are devoid of connective tissue capsules, and their cells are supported by the stroma of the organ in which they are found. Autonomic ganglia usually have multipolar neurons. As with craniospinal ganglia, autonomic ganglia have neuronal perikaryons with fine Nissl bodies.A layer of satellite cells frequently envelops the neurons of autonomic ganglia. In intramural ganglia, only a few satellite cells are seen around each neuron.
The Nerves-Histology of Nervous System
In the peripheral nervous system, the nerve fibers are grouped in bundles to form the nerves. Except for a few very thin nerves made up of unmyelinated fibers, nerves have a whitish, homogeneous, glistening appearance because of their myelin and collagen content.
Nerves have an external fibrous coat of dense connective tissue called epineurium, which also fills the space between the bundles of nerve fibers. Each bundle is surrounded by the perineurium, a sleeve formed by layers of flattened epitheliumlike cells. The cells of each layer of the perineurial sleeve are joined at their edges by tight junctions, an arrangement that makes the perineurium a barrier to the passage of most macromolecules and has the important function of protecting the nerve fibers from aggression. Within the perineurial sheath run the Schwann cell-sheathed axons and their enveloping connective tissue, the endoneurium. The endoneurium consists of a thin layer of reticular fibers, produced by Schwann cells.
The nerves establish communication between brain and spinal cord centers and the sense organs and effectors (muscles, glands, etc). They possess afferent and efferent fibers to and from the central nervous system. Afferent fibers carry the information obtained from the interior of the body and the environment to the central nervous system. Efferent fibers carry impulses from the central nervous system to the effector organs commanded by these centers. Nerves possessing only sensory fibers are called sensory nerves; those composed only of fibers carrying impulses to the effectors are called motor nerves. Most nerves have both sensory and motor fibers and are called mixed nerves; these nerves have both myelinated and unmyelinated axons .
Peripheral Nervous System-Histology of Nervous System
The main components of the peripheral nervous system are the nerves, ganglia, and nerve endings. Nerves are bundles of nerve fibers surrounded by connective tissue sheaths.
Nerve Fibers
Nerve fibers consist of axons enveloped by a special sheath derived from cells of ectodermal origin. Groups of nerve fibers constitute the tracts of the brain, spinal cord, and peripheral nerves.
Myelinated Fibers
In myelinated fibers of the peripheral nervous system, the plasmalemma of the covering Schwann cell winds and wraps around the axon . The layers of membranes of the sheath cell unite and form myelin. Myelin consists of many layers of modified cell membranes. These membranes have a higher proportion of lipids than do other cell membranes. The myelin sheath shows gaps along its path called the nodes of Ranvier; these represent the spaces between adjacent Schwann cells along the length of the axon. Interdigitating processes of Schwann cells partially cover the node. The distance between two nodes is called an internode and consists of one Schwann cell. The length of the internode varies between 1 and 2 mm. There are no Schwann cells in the central nervous system; there, the processes of the oligodendrocytes form the myelin sheath. Oligodendrocytes differ from Schwann cells in that different branches of one cell can envelop segments of several axons .
Unmyelinated Fibers
In both the central and peripheral nervous systems, not all axons are sheathed in myelin. In the peripheral system, all unmyelinated axons are enveloped within simple clefts of the Schwann cells. Unlike their association with individual myelinated axons, each Schwann cell can sheathe many unmyelinated axons. Unmyelinated nerve fibers do not have nodes of Ranvier, because abutting Schwann cells are united to form a continuous sheath.
The Central Nervous System-Histology of Nervous System
The central nervous system consists of the cerebrum, cerebellum, and spinal cord. It has almost no connective tissue and is therefore a relatively soft, gel-like organ.
When sectioned, the cerebrum, cerebellum, and spinal cord show regions that are white (white matter) and that are gray (gray matter). The differential distribution of myelin in the central nervous system is responsible for these differences: The main component of white matter is myelinated axons and the myelin-producing oligodendrocytes. White matter does not contain neuronal cell bodies.
Gray matter contains neuronal cell bodies, dendrites, and the initial unmyelinated portions of axons and glial cells. This is the region at which synapses occur. Gray matter is prevalent at the surface of the cerebrum and cerebellum, forming the cerebral and cerebellar cortex , whereas white matter is present in more central regions. Aggregates of neuronal cell bodies forming islands of gray matter embedded in the white matter are called nuclei.
In the cerebral cortex, the gray matter has six layers of cells with different forms and sizes. Neurons of some regions of the cerebral cortex register afferent (sensory) impulses; in other regions, efferent (motor) neurons generate motor impulses that control voluntary movements. Cells of the cerebral cortex are related to the integration of sensory information and the initiation of voluntary motor responses.
The cerebellar cortex has three layers : an outer molecular layer, a central layer of large Purkinje cells, and an inner granule layer. The Purkinje cells have a conspicuous cell body and their dendrites are highly developed, assuming the aspect of a fan. These dendrites occupy most of the molecular layer and are the reason for the sparseness of nuclei. The granule layer is formed by very small neurons (the smallest in the body), which are compactly disposed, in contrast to the less cell-dense molecular layer .
In cross sections of the spinal cord, white matter is peripheral and gray matter is central, assuming the shape of an H. In the horizontal bar of this H is an opening, the central canal, which is a remnant of the lumen of the embryonic neural tube. It is lined with ependymal cells. The gray matter of the legs of the H forms the anterior horns. These contain motor neurons whose axons make up the ventral roots of the spinal nerves. Gray matter also forms the posterior horns (the arms of the H), which receive sensory fibers from neurons in the spinal ganglia (dorsal roots).
Neuroglial Cells-Histology of Nervous System
Glial cells are 10 times more abundant in the mammalian brain than neurons; they surround both cell bodies and their axonal and dendritic processes that occupy the interneuronal spaces.
Nerve tissue has only a very small amount of extracellular matrix, and glial cells furnish a microenvironment suitable for neuronal activity.
- Oligodendrocytes produce the myelin sheath that provides the electrical insulation of neurons in the central nervous system. These cells have processes that wrap around axons, producing a myelin sheath
- Schwann cells have the same function as oligodendrocytes but are located around axons in the peripheral nervous system. One Schwann cell forms myelin around a segment of one axon, in contrast to the ability of oligodendrocytes to branch and serve more than one neuron and its processes.
- Astrocytes are star-shaped cells with multiple radiating processes. Astrocytes bind neurons to capillaries and to the pia mater (a thin connective tissue that covers the central nervous system). Astrocytes with few long processes are called fibrous astrocytes and are located in the white matter; protoplasmic astrocytes, with many short-branched processes, are found in the gray matter. In addition to their supporting function, astrocytes participate in controlling the ionic and chemical environment of neurons. Some astrocytes develop processes with expanded end feet that are linked to endothelial cells. It is believed that through the end feet, astrocytes transfer molecules and ions from the blood to the neurons (Blood-Brain Barrier). Expanded processes are also present at the external surface of the central nervous system, where they make a continuous layer. Furthermore, when the central nervous system is damaged, astrocytes proliferate to form cellular scar tissue. Astrocytes can influence neuronal survival and activity through their ability to regulate constituents of the extracellular environment, absorb local excess of neurotransmitters, and release metabolic and neuroactive molecules. The latter molecules include peptides of the angiotensinogen family, vasoactive endothelins, opioid precursors called enkephalins, and the potentially neurotrophic somatostatin. On the other hand, there is some evidence that astrocytes transport energy-rich compounds from the blood to the neurons and also metabolize glucose to lactate, which is then supplied to the neurons.
- Ependymal cells are low columnar epithelial cells lining the ventricles of the brain and central canal of the spinal cord. In some locations, ependymal cells are ciliated, which facilitates the movement of cerebrospinal fluid.
- Microglia are small elongated cells with short irregular processes. Microglia, phagocytic cells that represent the mononuclear phagocytic system in nerve tissue, are derived from precursor cells in the bone marrow. They are involved with inflammation and repair in the adult central nervous system, and they produce and release neutral proteases and oxidative radicals. When activated, microglia retract their processes and assume the morphological characteristics of macrophages, becoming phagocytic and acting as antigen-presenting cells. Microglia secrete a number of immunoregulatory cytokines and dispose of unwanted cellular debris caused by central nervous system lesions.
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