General Neuroanatomy

8 General Neuroanatomy


8.1 Development of the Central Nervous System (CNS)



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A Development of the brain


a Embryo with a greatest length (GL, see p. 4) of 10 mm, at the beginning of the 2nd month of development. Even at this stage we can see the differentiation of the neural tube into segments that will generate various brain regions (see C):


Medulla oblongata (gray)


Pons (gray)


Cerebellum (light blue)


Midbrain (mesencephalon, dark blue)


Interbrain (diencephalon, yellow)


Forebrain (telencephalon, red)


b Embryo with a GL of 27 mm, near the end of the 2nd month of development (end of the embryonic period). The olfactory bulb is developing from the telencephalon, part of the pituitary anlage (neurohypophysis) from the diencephalon.


c Fetus with a GL of 53 mm, in approximately the 3rd month of development. By this time the telencephalon has begun to overgrow the other brain areas. The insula is still on the surface but will subsequently be covered by the cerebral hemispheres (compare with d).


d Fetus with a crown–rump length (CRL, see p. 4) of 27 cm (270 mm), in approximately the 7th month of development. The brain has begun to develop conspicuous gyri and sulci.


Note: The telencephalon grows over all the other brain structures as development proceeds.



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B Brain vesicles and their derivatives


The cranial end of the neural tube expands to form three primary brain vesicles: the forebrain (prosencephalon), the midbrain (mesencephalon), and the hindbrain (rhombencephalon). The forebrain (telencephalon) and interbrain (diencephalon) develop from the prosencephalon. The mesencephalon gives rise to the superior and inferior colliculi and related structures. The rhombencephalon differentiates into the pons, cerebellum, and medulla oblongata. The pons and cerebellum are also known collectively as the metencephalon. Some important structures of the adult brain are listed at far right to illustrate the derivatives of the brain vesicles. They can be traced back in the diagram to their developmental precursors.




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D Differentiation of the neural tube in the spinal cord region during development


Cross section, cranial view.


a Early neural tube, b intermediate stage, c adult spinal cord.


The neurons that form in the basal plate are efferent (motor) neurons, while those that form in the alar plate are afferent (sensory) neurons.


The area between them—the future thoracic, lumbar, and sacral cord—is another zone that gives rise to preganglionic autonomic neurons. The roof plate and floor plate do not form neurons. A knowledge of how these neuron populations are distributed is helpful in understanding the structure of the hindbrain (rhombencephalon, see E).



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E Embryonic migratory movements of neuron populations and their effect on the location of the cranial nerve nuclei


Cross section, cranial view. (Visual aid: If we compare the spinal cord to a book, it would be closed in a and open in b and c.)


a In the initial stage, the motor neurons are ventral (anterior), and the sensory neurons are dorsal (posterior). The arrows indicate the directions of migration.


b In the early embryonic stage, the neurons of the alar plate migrate laterally and ventrally (anteriorly).


c In the adult brain (medulla oblongata and pons, derivatives of the rhombencephalon), we can distinguish four nuclear columns (after His and Herrick) that contain functionally distinct cranial nerve nuclei (from medial to lateral):


1. Somatomotor column (lilac)


2. Visceromotor column (orange and green stripes)


3. Viscerosensory column (light blue)


4. Somatosensory column (dark blue)


8.2 Neural Crest Derivatives and the Development of the Peripheral Nervous System (PNS)



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A Development of the neural crest cells


At 3 weeks’ development, the notochord induces surface ectoderm in the medial embryonic disc to thicken and form the neural plate (neuroectoderm). The neural plate differentiates to form the primordia of the nervous system. Neural folds are raised on each side of the neural plate, and a median groove develops between them—the neural groove. This groove subsequently deepens and closes to form the neural tube, which sinks below the ectoderm. Portions of the folds that do not contribute to neural tube formation differentiate to form the neural crest. While neural crest cells in the future head region start to migrate even before the neural tube is closed, crest migration in the trunk is delayed until tube closure. Cells destined to form the neural crest detach from the ectoderm at the fusing margins of the neural tube and undergo an epithelio-mesenchymal transition, diving into underlying mesoderm, where they begin a long and tortuous migration. As the neural tube differentiates into the central nervous system, migrating neural crest cells settle in different locations and develop into sensory and autonomic ganglia, endocrine glands, melanocytes, cartilage, and other structures (see B and C) (after Wolpert).



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B Main migratory pathways and derivatives of the neural crest (after Christ and Wachtler)


Neural crest cells originated in different regions have different migration paths and fates. Those from cranial levels contribute to cartilage and bone in the head and neck, and to cranial parasympathetic ganglia (see C). Those from thoracolumbar levels do not become skeletal cells, but generate peripheral neurons, endocrine cells, melanocytes, and Schwann cells. This diagram shows the migration of the neural crest cells in the trunk of the early embryo (4 weeks’ development, see p. 7). They follow three main migratory pathways:


Dorsolateral pathway (melanoblasts, which differentiate into melanocytes)


Ventrolateral pathway (ganglioblasts, which differentiate into sensory nerve cells in the dorsal (posterior) root [spinal] ganglia)


Ventral pathway (cells differentiate into neurons and associated cells of the paravertebral sympathetic ganglia, into chromaffin cells of the adrenal medulla, and into autonomic plexuses in the gastrointestinal tract)


Thus, the neural crest can develop into a variety of seemingly unrelated nonneuronal cells as well as peripheral ganglion cells. These unusual characteristics of the neural crest—its pluripotential capacity and wide-ranging migration—have consequences when its differentiation or migration is defective. Disruption of neural crest development may deprive organs of their autonomic innervation (Hirschsprung disease). Tumors derived from neural crest cells tend to be highly malignant and difficult to treat. (See D.)



D Diseases of neural crest derivatives (selected examples)
































Neural crest


Disease


Parasympathetic visceral ganglia


Neuroblastoma


(malignant childhood tumor)


Enteric nervous system


Hirschsprung disease


(aganglionic colon)


Glial cells (Schwann cells, satellite cells)


Neurofibromatosis


(Recklinghausen disease)


Melanocytes


Malignant melanoma, albinism


Adrenal medulla


Pheochromocytoma (adrenal gland tumor)


Endocrine cells of the lung and heart


Carcinoids (malignant tumors with endocrine activity)


Parafollicular cells (C cells) of the thyroid gland


Medullary thyroid carcinoma



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E Development of a peripheral nerve


Afferent (blue) and efferent (red) axons sprout separately from the neuron cell bodies during early development (a). Primary afferent (sensory) neurons develop in the dorsal (posterior) root (spinal) ganglia, and primary motor neurons develop from the basal plate of the spinal cord (b). Interneurons (black), which connect sensory ganglia and motor neurons, develop later.



8.3 Topography and Structure of the Nervous System





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C Terms of location and direction in the CNS


Midsagittal section, right lateral view.


Note two important axes:


① The almost vertical brainstem axis (corresponds approximately to the body axis).


② The horizontal axis through the diencephalon and telencephalon. Keep these reference axes in mind when using directional terms in the CNS.



8.4 Cells of the Nervous System



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A The nerve cell (neuron)


The neuron is the smallest functional unit of the nervous system. Neurons communicate with other nerve cells through synapses. The synapses that end at nerve cells usually do so at dendrites (as seen here). The transmitter substance that is released at the synapses to act on the dendrite membrane may have an excitatory or inhibitory action, meaning that the transmitter either increases or decreases the local action potential at the nerve cell membrane. All of the excitatory and inhibitory potentials of a nerve cell are integrated in the axon hillock. If the excitatory potentials predominate, the stimulus exceeds the excitation threshold of the neuron, causing the axon to fire (transmit an impulse) according to the all-or-nothing rule.



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B Electron microscopy of the neuron


Neurons are rich in rough endoplasmic reticulum (protein synthesis, active metabolism). This endoplasmic reticulum (known also as Nissl substance) is easily demonstrated by light microscopy using cationic dyes, which bind to the phosphodiester backbone of the ribosomal RNAs. The distribution pattern of the Nissl substance is used in neuropathology to evaluate the functional integrity of neurons. Neurotubules and neurofilaments are referred to collectively as neurofibrils in light microscopy, as they are too fine to be identified as separate structures under a light microscope. Neurofibrils can be demonstrated in light microscopy by impregnating the nerve tissue with silver salts. This is of interest in neuropathology because the clumping of neurofibrils is an important histologic feature of Alzheimer disease.



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C Basic forms of the neuron and its functionally adapted variants


The horizontal line marks the region of the axon hillock, which represents the initial segment of the axon. (The structure of a peripheral nerve, consisting only of axons and sheath tissue, is shown on p. 77.)


a Multipolar neuron (multiple dendrites) with a long axon (= long transmission path). Examples are projection neurons such as α-motoneurons in the spinal cord.


b Multipolar neuron with a short axon (= short transmission path). Examples are interneurons like those in the gray matter of the brain and spinal cord.


c Pyramidal cell: Dendrites are present only at the apex and base of the tridentate cell body, and the axon is long. Examples are efferent neurons of the cerebral motor cortex.


d Purkinje cell: An elaborately branched dendritic tree arises from a circumscribed site on the cell body. The Purkinje cell receives many synaptic contacts from afferents to the cerebellum and is also the efferent cell of the cerebellar cortex.


e Bipolar neuron: The dendrite branches in the periphery. Examples are bipolar cells of the retina.


f Pseudounipolar neuron: The dendrite and axon are not separated by a cell body. An example is the primary afferent (= first sensory) neuron in the spinal ganglion (see p. 91).




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E Electron microscopy of synapses in the CNS


Synapses are the functional connection between two neurons. They consist of a presynaptic membrane, a synaptic cleft, and a postsynaptic membrane. In a spine synapse (1), the presynaptic knob (bouton) is in contact with a specialized protuberance (spine) of the target neuron. The side-by-side synapse of an axon with the flat surface of a target neuron is called a parallel contact or bouton en passage (2). The vesicles in the presynaptic expansion contain the neurotransmitters that are released into the synaptic cleft by exocytosis when the axon fires. From there the neurotransmitters diffuse to the postsynaptic membrane, where their receptors are located. A variety of drugs and toxins act upon synaptic transmission (antidepressants, muscle relaxants, toxic gases, botulinum toxin).



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F Cells of the neuroglia in the CNS


Neuroglial cells surround the neurons, providing them with structural and functional support (see G). Various staining methods are available in light microscopy for selectively demonstrating different portions of the neuroglial cells:


a Cell nuclei demonstrated with a basic stain.


b Cell body demonstrated by silver impregnation.


G Summary: cells of the CNS and PNS and their functional importance
































Type of cell


Function


Neurons (CNS and PNS)


1. Impulse formation


2. Impulse conduction


3. Information processing


Glial cells


 


Astrocytes (CNS only)


1. Maintain a constant internal milieu in the CNS


2. Contribute to the structure of the blood–brain barrier (see p. 95)


3. Phagocytize dead synapses


4. Form scar tissue in the CNS (e.g., in multiple sclerosis or following a stroke)


Microglial cells (CNS only)


Phagocytosis (“macrophages of the brain”)


Oligodendrocytes (CNS only)


Myelin sheath formation in the CNS


Schwann cells (PNS only)


Myelin sheath formation in the PNS


Satellite cells (PNS only)


Modified Schwann cells; surround the cell body of neurons in PNS ganglia


8.5 Structure of a Spinal Cord Segment



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A Structure of a spinal cord segment with its spinal nerve


Superior view. The spinal cord is made up of 31 consecutive segments arranged one above the other (see B). An anterior root and a posterior root emerge from the sides of each segment. The anterior root consists of efferent (motor) fibers, while the posterior root consists of afferent (sensory) fibers. Both roots from one segment unite in the intervertebral foramen to form the spinal nerve. The afferent (sensory) fibers and efferent (motor) fibers intermingle at this junction, so that the branches into which the spinal nerve divides (see below) contain motor and sensory elements (except for the meningeal branch, which is purely sensory). This division into branches (rami) occurs shortly after the anterior and posterior roots unite to form the spinal nerve. Consequently, the spinal nerve itself is only about 1 cm long.


The principal branches of the spinal nerve have the following functions:


The anterior ramus innervates the anterior and lateral body wall and the limbs.


The posterior ramus innervates the skin of the back and the intrinsic back muscles.


The meningeal branch reenters the spinal canal, providing sensory innervation to the spinal membranes and other structures.


The white ramus communicans carries white (= myelinated) fibers to the ganglion of the sympathetic trunk.


The gray ramus communicans carries gray (= unmyelinated) fibers from the sympathetic ganglion back to the spinal nerve (the functional significance of this is described on p. 97). The anterior and posterior rami of the spinal nerves subdivide into further branches.

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Jul 25, 2021 | Posted by in ANATOMY | Comments Off on General Neuroanatomy

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