NEUROHISTOLOGY

The nervous system develops embryologically from ectoderm, which forms the neural plate

Successive growth and folding of the plate results in the formation of the primitive neural tube.

The neuroblasts in the wall of the tube differentiates into 3 cell types:

Neurons:  conduction of impulses

Neuroglial cells: connective tissue and support of CNS

Ependymal cells:  Lines the lumen of the tube.

   - Specialized neuro-ectodermal cells which lines the ventricles of the adult brain

                - Essentially also a neuroglial cell

Basic Unit = neuron

Exhibits irritability (excitability) and conductivity

A typical neurons consists of:

Cell body : Has nucleus (karyon) and surrounding cytoplasm (perikaryon) which contains organelles cell's vitality

Dendrites:  Several short processes

Axon:One large process

Terminates in twig like branches (telodendrons)

May also have collateral branches projecting along its course. These exit at nodes of Ranvier

Axon enveloped in a sheath, and together forms the nerve fiber

Classification:

May be done in different ways, i.e.

Functional = afferent, efferent, preganglionic, postganglionic, etc.

Morphological = shape, processes, etc

A typical morphological classification is as follows

a. Unipolar: Has one process only Not found in man

b. Bipolar (so-called ganglion cell):Has two processes Found in sensory systems, e.g. retina olfactory system

c. Multipolar: Has several process Most common in CNS

Cell bodies vary in shape, e.g.  stellate (star) , pyramidal

d. Pseudo-unipolar: Essentially bipolar neurons, but processes have swung around cb and fused with each other. They therefore enter and leave at one pole of the cell.

Typical neuron:

- Has 2 or more dendrites

Close to the cb the cytoplasm of dendrites has Nissl granules as well as mitochondria

Only one axon Arises from axon hillock, Devoid of Nissl granules, Encased in myelin sheath

No additional covering except for occasional foot processes of neuroglial cells

May branch at right angles

Branches at a node of Ranvier is known as a collateral

Ends of axons break up into tree-like branches, known as telodendria

Axons may be short (Golgi Type II) e.g. internuncial long (Golgi Type I) e.g. pyramidal neuron

Nucleus Central position Large and spherical

Chromatin is extended and thus not seen in LM. This allows the nucleolus to be prominent

Cytoplasm (perikaryon)

Surrounds nucleus  May be large or small, shape may be round, oval, flattened, pyramidal, etc

Contains aggregates Nissl granules(Bodies) which is also sometimes referred to as rhomboid flakes

aggregation of membranes and cisternae of rough endoplasmic reticulum (RER)

numerous ribosomes and polyribosomes scattered between cisternae

(Polyribosome = aggregate of free ribosomes clumped together)

responsible for ongoing synthesis of new cytoplasm and cytoplasmic substances

needed for conduction of impulses

highly active in cell protein synthesis

resultant loss of power to divide which is characteristic of neurons

- Golgi network surrounding nucleus (seen in EM only)

- Fibrils made up of:

- neurofilaments

- microtubules

Tubules involved in:

1. plasmic transport

2. maintenance of cell shape

3. essential for growth and elongation of axons and dendrites

Neurofilament:

1. provide skeletal framework

2. maintenance of cell shape

3. possible role in axonal transport

(Axonal [axoplasmic; plasmic] transport may be antero- or retrograde. Anterograde transport via neurotubules is fast and moves neurotransmitters. Retrograde transport is slow and is the reason why viruses and bacteria can attack and destroy cell bodies. E.g. polio in the ventral columns and syphilis in the dorsal columns).

- Numerous mitochondria

- Neurons lack ability to store glycogen and are dependent for energy on circulating glucose

Impulses are conducted in one direction only

Dendrites conduct towards the cb

Axons conduct away from cb

Synapses:

- Neurons interconnect by way of synapses

- Normally the telodendria of an axon synapse with the dendrites of a succeeding axon

axo-dendritic synapse

This is usually excitatory

- Other types of synapses are:

 axo-axonic

May be excitatory and/or inhibitory

axo-somatic

May be excitatory and/or inhibitory

 dendrodendritic

Usually inhibitory

- Synapses are not tight junctions but maintain a narrow space the so-called synaptic cleft

- The end of an telodendron is usually enlarged (bouton) and contains many synaptic vesicles,

mitochondrion, etc. Its edge that takes part in the synapse is known as the postsynaptic membrane and no

vesicles are seen in this area

- Synapses may be chemical (as above) or electrical as in the ANS supplying smooth muscle cells subjacent to adjacent fibres

Gray and White Matter of Spinal Cord:

- Gray matter contains:

- cb's (somas) of neurons

- neuroglial cells

- White matter contains:

- vast number of axons

- no cb's

- colour of white matter due to myelin that ensheathes axons

Myelin:

- Non-viable fatty material contains phospholipids, cholesterol and some proteins

- Soluble and not seen in H&E-sections because it has become dissolved in the process, thus leaving empty spaces around the axons

- Osmium tetroxide (OsO4) fixes myelin and makes it visible by staining it black. Seen as concentric rings in cross section

- Myelin sheath (neurolemma) is formed by two types of cells

- Within the CNS by Oligodendrocytes

- On the peripheral neurons system by Schwann cells

- Sheath is formed by being wrapped around the axon in a circular fashion by both types of cells

Neuroglial Cells:

- Forms roughly 40% of CNS volume

- May function as: 1. support

2. nurture ("feeding")

3. maintain

Types of glial cells:

Oligodendrocytes:

- Small dark stained dense nucleus

- Analogue of Schwann cell in peripheral nervous system

- Has several processes which forms internodal segments of several fibres (one cell ensheathes more than one axon)

- Provides myelin sheaths in CNS

- Role in nurturing (feeding) of cells

Astrocytes:

Protoplasmic astrocytes:

- found in gray matter

- round cell body

- large oval nucleus with prominent nucleolus

- large thick processes

- processes are short but profusely branched

- perivascular and perineurial foot processes

- sometimes referred to as mossy fibres

Fibrous Astrocytes:

- found in white matter

- polymorphic cells body

- large oval nucleus

- long thin processes

Microglia:

- Neural macrophages

- smallest of the glial cells

- intense dark stained nucleus

- conspicuously fine processes which has numerous short branches

Cerebral Cortex:

Consists of six layers which are best observed in the cortex of the hippocampus

From superficial to deep:

- Molecular layer:

- Has few cells and many fibres of underlying cells

- Outer granular layer:

- Many small nerve cells

- Pyramidal layer:

- Pyramidally-shaped cells bodies

- Inner granular layer:

- Smaller cells and nerve fibres

- Internal (inner) pyramidal layer:

- Pyramidal cells bodies

- Very large in the motor cortex and known as Betz-cells

- Polymorphic layer:

- Cells with many shapes

Cerebellar Cortex:

Consists of three layers

Connections are mainly inhibitory

From superficial to deep

- Outer molecular layer:

- Few cells and many fibres

- Purkinje layer:

- Huge flask-shaped cells that are arranged next to one another

- Inner granular layer:

- Many small nerve cells

Motor endplate:

Seen in periphery on striated muscle fibres

- known as boutons

- has no continuous myelin covering from the Schwann cells

- passes through perimysium of muscle fiber to "synapse"

- multiple synaptic gutter (fold) in sarcoplasma of muscle fiber beneath bouton

- contains numerous synaptic vesicles and mitochondria

Ganglia:

- Sensory Ganglia:

(e.g. trigeminal nerve, ganglia and dorsal root ganglia)

- No synapse (trophic unit)

- pseudo-unipolar neurons

- centrally located nucleus

- spherical smooth border

- conspicuous axon hillock

- Surrounded by cuboidal satellite cells (Schwann cells)

- Covered by spindle shaped capsular cells of delicate collagen which forms the endoneurium

- Visceral and Motor Ganglia (Sympathetic and Parasympathetic):

- Synapse present

- Ratio of preganglionic: postganglionic fibres

1. Sympathetic 1:30

Therefore excitatory and catabolic

2. Parasympathetic 1:2

Therefore anabolic

Except in Meissner and Auerbach's plexuses where ratio is 1:1000 '2 because of parasympathetic component's involvement in digestion

- Preganglionic axons are myelinated (e.g. white communicating rami)

- Postganglionic axon are non-myelinated (e.g. gray communicating rami)

- small multipolar cell body

- excentrally located nucleus

- Inconspicuous axon hillock

- satellite cells few or absent

- few capsular cells

The Hard Palate

• The anterior bony part of the palate is formed by the palatine process of the maxillae and the horizontal plates of the palatine bones.

• Anteriorly and laterally, the hard palate is bounded by the alveolar processes and the gingivae.
• Posteriorly, the hard palate is continuous with the soft palate.

• The incisive foramen is the mouth of the incisive canal.
• This foramen is located posterior to the maxillary central incisor teeth.
• This foramen is the common opening for the right and left incisive canals.
• The incisive canal and foramen transmit the nasopalatine nerve and the terminal branches of the sphenopalatine artery.

• Medial to the third molar tooth, the greater palatine foramen pierces the lateral border of the bony palate.
• The greater palatine vessels and nerve emerge from this foramen and run anteriorly into two grooves on the palate.

• The lesser palatine foramen transmits the lesser palatine nerve and vessels.
• This runs to the soft palate and adjacent structures.

Muscles of the Soft Palate

The Levator Veli Palatini (Levator Palati)

• Superior attachment: cartilage of the auditory tube and petrous part of temporal bone.
• Inferior attachment: palatine aponeurosis.
• Innervation: pharyngeal branch of vagus via pharyngeal plexus.
• This cylindrical muscle runs inferoanteriorly, spreading out in the soft palate, where it attaches to the superior surface of the palatine aponeurosis.
• It elevates the soft palate, drawing it superiorly and posteriorly.
• It also opens the auditory tube to equalise air pressure in the middle ear and pharynx.

The Tensor Veli Palatini (Tensor Palati)

• Superior attachment: scaphoid fossa of medial pterygoid plate, spine of sphenoid bone, and cartilage of auditory tube.
• Inferior attachment: palatine aponeurosis.
• Innervation: medial pterygoid nerve (a branch of the mandibular nerve).
• This thin, triangular muscle passes inferiorly, and hooks around the hamulus of the medial pterygoid plate.
• It then inserts into the palatine aponeurosis.
• This muscle tenses the soft palate by using the hamulus as a pulley.
• It also pulls the membranous portion of the auditory tube open to equalise air pressure of the middle ear and pharynx.

The Palatoglossus Muscle

• Superior attachment: palatine aponeurosis.
• Inferior attachment: side of tongue.
• Innervation: cranial part of accessory nerve (CN XI) through the pharyngeal branch of vagus (CN X) via the pharyngeal plexus.
• This muscle, covered by mucous membrane, forms the palatoglossal arch.
• The palatoglossus elevates the posterior part of the tongue and draws the soft palate inferiorly onto the tongue.

The Palatopharyngeus Muscle

• Superior attachment: hard palate and palatine aponeurosis.
• Inferior attachment: lateral wall of pharynx.
• Innervation: cranial part of accessory nerve (CN XI) through the pharyngeal branch of vagus (CN X) via the pharyngeal plexus.
• This thin, flat muscle is covered with mucous membrane to form the palatopharyngeal arch.
• It passes posteroinferiorly in this arch.
• This muscle tenses the soft palate and pulls the walls of the pharynx superiorly, anteriorly and medially during swallowing.

The Musculus Uvulae

• Superior attachment: posterior nasal spine and palatine aponeurosis.
• Inferior attachment: mucosa of uvula.
• Innervation: cranial part of accessory through the pharyngeal branch of vagus, via the pharyngeal plexus.
• It passes posteriorly on each side of the median plane and inserts into the mucosa of the uvula.
• When the muscle contracts, it shortens the uvula and pulls it superiorly.

Sternum

o    Forms the medial part of the anterior chest wall
o    Manubrium (upper part)-clavicle and first rib articulate with the manubrium .
o    Body (middle blade)-second and tenth ribs articulate with the body via the costal cartilages
o    Xiphoid (blunt cartilaginous tip)

Ribs (12 pairs)

o    Each rib articulates with both the body and the transverse process of its corresponding
o    thoracic vertebra
o    The second to ninth ribs articulate with the body of the vertebra above'
o    Ribs curve outward, forward, and then downward
o    Anteriorly, each of the first seven ribs joins a costal cartilage that attaches to the sternum
o    Next three ribs (eighth to tenth) join the cartilage of the rib above
o    Eleventh and twelfth ribs do not attach to the sternum; are called "floating ribs"

Eye 

At week 4, two depressions are evident on each of the forebrain hemispheres.  As the anterior neural fold closes, the optic pits elongate to form the optic vesicles.  The optic vesicles remain connected to the forebrain by optic stalks. 
The invagination of the optic vesicles forms a bilayered optic cup.  The bilayered cup becomes the dual layered retina (neural and pigmented layer)
Surface ectoderm forms the lens placode, which invaginates with the optic cup.
The optic stalk is deficient ventrally to contain choroids fissure to allow blood vessels into the eye (hyaloid artery).  The artery feeds the growing lens, but will its distal portion will eventually degenerate such that the adult lens receives no hyaloid vasculature.
At the 7th week, the choroids fissure closes and walls fuse as the retinal nerve get bigger.
The anterior rim of the optic vesicles forms the retina and iris.  The iris is an outgrowth of the distal edge of the retina.
Optic vesicles induces/maintains the development of the lens vesicle, which forms the definitive lens.  Following separation of the lens vesicle from the surface ectoderm, the cornea develops in the anterior 1/5th of the eye.
The lens and retina are surrounded by mesenchyme which forms a tough connective tissue, the sclera, that is continuous with the dura mater around the optic nerve.  
Iridopupillary membrane forms to separate the anterior and posterior chambers of the eye.  The membrane breaks down to allow for the pupil
Mesenchyme surrounding the forming eye forms musculature (ciliary muscles and pupillary muscles – from somitomeres 1 and 2; innervated by CN III), supportive connective tissue elements and vasculature.

Eyelids

Formed by an outgrowth of ectoderm that is fused at its midline in the 2nd trimester, but later reopen.

Connective Tissue

Functions of Connective tissue:

→ joins together other tissues

→ supporting framework for the body (bone)

→ fat stores energy

→ blood transports substances

Connective tissue is usually characterized by large amounts of extracellular materials that separate cells from each other, whereas epithelial tissue is mostly cells with very little extracellular material. The extracellular substance of connective tissue consists of protein fibers which are embedded in ground substance containing tissue fluid.

Fibers in connective tissue can be divided into three types:

→ Collagen fibers are the most abundant protein fibers in the body.

→ Elastic fibers are made of elastin and have the ability to recoil to original shape.

→ Reticular fibers are very fine collagen fibers that join connective tissues to other tissues.

Connective tissue cells are named according to their functions:

 → Blast cells produce the matrix of connective tissues

→ Cyte cells maintains the matrix of connective tissues

→ Clast cells breaks down the matrix for remodeling (found in bone)