1.

Neuron (Perikaryon, Nissl bodies, Cytoskeleton)

The functional unit in both the CNS and PNS is the neuron.Most neurons
have three main parts:
■ The cell body (also called the perikaryon or soma),contains the nucleus and
most of the cell’s organelles. serves as the synthetic or trophic center for the
entire neuron.
■ The dendrites are the numerous elongated processes extending from the
perikaryon and specialized to receive stimuli from other neurons at unique
sites called synapses.
■ The axon is a single long process ending at synapses specialized to generate
and conduct nerve impulses to other cells (eg, nerve, muscle, and gland cells).
Axons may also receive information from other neurons, information that
mainly modifies the transmission of action potentials to those neurons.

Neurons can be classified according to the number of processes extending

from the cell body:
■ Multipolar neurons- each with one axon and two or more dendrites, are the
most common.
■ Bipolar neurons, with one dendrite and one axon,comprise the sensory
neurons of the retina, the olfactory epithelium, and the inner ear.
■ Unipolar or pseudounipolar neurons, which include all other sensory
neurons, each have a single process that bifurcates close to the perikaryon,
with the longer branch extending to a peripheral ending and the other toward
the CNS.
■ Anaxonic neurons, with many dendrites but no true axon, do not produce
action potentials, but regulate electrical changes of adjacent CNS neurons.
Because the fine processes emerging from cell bodies are seldom seen in
sections of nervous tissue, it is difficult to classify neurons structurally by
microscopic inspection.

Nervous components can also be subdivided functionally.

■ Sensory neurons are afferent, receiving from receptors throughout the body.
■ Motor neurons are efferent, sending impulses to effector organs such as
muscle fibers and glands.
Somatic motor nerves are under voluntary control and typically innervate
skeletal muscle; autonomic motor nerves control the involuntary or
unconscious activities of glands, cardiac muscle, and most smooth muscle.
Interneurons establish relationships among other neurons, forming complex
functional networks or circuits in the CNS.
Interneurons are either multipolar or anaxonic and comprise 99% of all
neurons in adults.In the CNS most neuronal perikarya occur in the gray
matter, with their axons concentrated in the white matter. Tese terms refer to
 the general appearance of unstained CNS tissue caused in part by the diferent
densities of nerve cell bodies. In the PNS cell bodies are found in ganglia and
in some sensory regions, such as the olfactory mucosa, and axons are bundled
in nerves.

The neuronal cell body contains the nucleus and surrounding cytoplasm,
exclusive of the cell processes. It acts as a trophic center, producing most
cytoplasm for the processes. Most cell bodies are in contact with a great
number of nerve endings conveying excitatory or inhibitory stimuli generated
in other neurons. A typical neuron has an unusu- ally large, euchromatic
nucleus with a prominent nucleolus, indicating intense synthetic activity.
Cytoplasm of perikarya often contains numerous free polyribosomes and
highly developed RER, indicating active production of both cytoskeletal
proteins and proteins for transport and secretion. Histologically these regions
with concentrated RER and other polysomes are basophilic and are
distinguished as chromatophilic substance (or Nissl substance, Nissl bodies)
(Figure 9–3). The amount of this material varies with the type and functional
state of the neu- ron and is particularly abundant in large nerve cells such as
motor neurons (Figure 9–3b). The Golgi apparatus is located only in the cell
body, but mitochondria can be found throughout the cell and are usually
abundant in the axon terminals.
In both perikarya and processes microtubules, actin filaments and
intermediate filaments are abundant, with the latter formed by unique protein
subunits and called neurofilaments in this cell type. Cross-linked with certain
fixatives and impregnated with silver stains, neurofilaments are also referred
to as neurofibrils by light microscopists. Some nerve cell bodies also contain
inclusions of pigmented material, such as lipofus- cin, consisting of residual
bodies left from lysosomal digestion.

2. Dendrites & Axons

Dendrites
Dendrites (Gr. dendron, tree) are typically short, small pro- cesses emerging
and branching of the soma (Figure 9–3). Usually covered with many synapses,
dendrites are the principal signal reception and processing sites on neurons.
Te large number and extensive arborization of dendritesallowa single neu-
ron to receive and integrate signals from many other nerve cells. For example,
up to 200,000 axonal endings can make func- tional contact with the dendrites
of a single large Purkinje cell of the cerebellum.
Unlike axons, which maintain a nearly constant diameter, dendrites become
much thinner as they branch, with cyto- skeletal elements predominating in
these distal regions. In the CNS most synapses on dendrites occur on
dendritic spines, which are dynamic membrane protrusions along the small
dendritic branches, visualized with silver staining (Figure 9–5)and studied by
confocal or electron microscopy. Dendritic spines serve as the initial
processing sites for synaptic signals and occur in vast numbers, estimated to
be on the order of 1014 for cells of the human cerebral cortex. Dendritic spine
mor- phology depends on actin flaments and changes continuously as
synaptic connections on neurons are modifed. Changes in dendritic spines are
of key importance in the constant changes of the neural plasticity that occurs
during embryonic brain development and underlies adaptation, learning, and
memory postnatally.
Axons
Most neurons have only one axon, typically longer than its dendrites. Axonal
processes vary in length and diameter according to the type of neuron. Axons
of the motor neurons that innervate the foot muscles have lengths of nearly a
meter; large cell bodies are required to maintain these axons, which contain
most of such neurons’ cytoplasm. Te plasma mem- brane of the axon is ofen
called the axolemma and its con- tents are known as axoplasm.
Axons originate from a pyramid-shaped region of the perikaryon called the
axon hillock (Figure 9–3), just beyond which the axolemma has concentrated
ion channels thatgenerate the action potential. At this initial segment of the
axon, the various excitatory and inhibitory stimuli impinging on the neuron
are algebraically summed, resulting in the deci- sion to propagate—or not to
propagate—a nerve impulse.
Axons generally branch less profusely than dendrites, but do undergo
terminal arborization (Figure 9–3). Axons of interneurons and some motor
neurons also have major branches called collaterals that end at smaller
branches with synapses infuencing the activity of many other neurons. Each
small axonal branch ends with a dilation called a terminal bouton (Fr. bouton,
button) that contacts another neuron or non-nerve cell at a synapse to initiate
an impulse in that cell.
Axoplasm contains mitochondria, microtubules, neuro- flaments, and
transport vesicles, but very few polyribosomes or cisternae of RER, features
that emphasize the dependence of axoplasm on the perikaryon. If an axon is
severed from its cell body, its distal part quickly degenerates and undergoes
phagocytosis.
Lively bidirectional transport of molecules large and small occurs within
axons. Organelles and macromolecules synthesized in the cell body move by
anterograde trans- port along axonal microtubules via kinesin from the peri-
karyon to the synaptic terminals. Retrograde transport in the opposite
direction along microtubules via dynein carries certain other macromolecules,
such as material taken up by endocytosis (including viruses and toxins), from
the periph- ery to the cell body. Retrograde transport can be used to study the
pathways of neurons: if peroxidase or another marker is
 injected into regions with axon terminals, its later distribution throughout the
neurons serving such regions can be deter- mined histochemically.
Anterograde and retrograde transports both occur fairly rapidly, at rates of
50-400 mm/d. A much slower anterograde stream, moving only a few
millimeters per day, involves move- ment of the axonal cytoskeleton itself. Tis
slow axonal trans- port corresponds roughly to the rate of axon growth.

3. Types of Nerve Cells, MAP – Types, Classification, Cell Types

4. Multipolar Neurons & Bipolar Neurons

5. Neuromediator Types

6. Synapses- Electrical, Chemical, Ultrastructure

Classifi cation depends on the mechanism of conduction of the nerve
impulses and the way the action potential is generated in the target cells. Th
us, synapses may also be classifi ed as the following.
• Chemical synapses. Conduction of impulses is achieved by the release of
chemical substances ( neuro transmitters) from the presynaptic neuron.
Neurotransmitters then diff use across the narrow intercellular space that
separates the presynaptic neuron from the postsynaptic neuron or target cell.
A specialized type of chemical synapses called ribbon synapses are found in
the receptor hair cells of the internal ear and photoreceptor cells of the retina.
Th eir structures and functions are described in Chapter 25).

• Electrical synapses. Common in invertebrates, these synapses contain gap

junctions that permit movement of ions between cells and consequently
permit the direct spread of electrical current from one cell to another. Th ese
synapses do not require neurotransmitters for their function. Mammalian
equivalents of electrical synapses include gap junctions in smooth muscle and
cardiac muscle cells
7. Neuroglia –General Consideration

Glial cells support neuronal survival and activities, and are 10 times more
abundant than neurons in the mammalian brain. Like neurons most glial cells
develop from progenitor cells of the embryonic neural plate. In the CNS glial
cells sur- round both the neuronal cell bodies, which are ofen larger than the
glial cells, and the processes of axons and dendrites occupying the spaces
between neurons. Except around the
 larger blood vessels, the CNS has only a very small amount of connective
tissue and collagen. Glial cells substitute for cells of connective tissue in some
respects, supporting neurons and creating immediately around those cells
microenvironments that are optimal for neuronal activity. Te fbrous
intercellular network of CNS tissue superficially resembles collagen by light
microscopy, but is actually the network of fne cellular pro- cesses emerging
from neurons and glial cells. Such processes are collectively called the
neuropil (Figure 9–8). Tere are six major kinds of glial cells,as shown
schematically in Figure 9–9, four in the CNS and two in the PNS. Teir main
functions, locations, and origins are summarized in Table 9–2.

8. Astrocytes-Types, Functions, (Immunity, Phagocytosis, Synthetic Function,

Glial Fibrillary Acid Protein)

Two main types of astrocytes can be dis- tinguished in both light and electron
mi- croscopy. The fibrous nstrocyte (spider cell) is characterized by its thin,
poorly branched processes, which radiate from the cell body for considerable
distances. These glial ele- ments are often interposed between neurons and
adjacent blood vessels and have promi- nent perivascular end feet (9)(Figs.
6.1, 6.3 and
6.4). Fibrous astrocytes are most numerous in the white matter (Fig. 6.2). With
appropriate stains the cell body and processes are seen to contain many
delicate fibrils. Each intracellu- lar gliofilament is 10 nm in width and of vari-
ous length. Such filaments correspond to the thicker fibrils observed in
muscle and epithlial cells (i.e., myofibrils and tonofibrils) and are part of the
group of intermediate filaments, which form neurofilaments in neurons (see
the section entitled Ne11ro11n/ Cytoskeleton in Chapter 5). In the larger
processes, they are arranged in straight parallel bundles and can be followed
for considerable distances. Such gliofibrils are much less numerous in the
protoplasmic astrocytes described later. Another feature common to both
types of astrocytes are small granular swellings along the pro- cesses called
gliosollies (Figs. 6.3 and 6.4). They occur in the cell body as well, and in
electron micrographs they are seen to be clumps of mitochondria which
contain a dense matrix material.
The protoplns111icnstrocytes (mossy cells) are most numerous and easy to
identify in the gray matter (Fig. 6.2). They have numerous freely branching
processes, perivascular end feet, and often are observed in close proxim- ity to
neuronal soma and dendrites. If fibrous and protoplasmic astrocytes are
examined in Golgi preparations (Fig. 6.4), one can find sharp distinctive cells
 in each category. In the same sections, one can also observe a host of
intermediate and transitional cells that defy a precise morphologic
classification. Electron microscopic studies suggest that they may represent
different forms of the same cell (69,
90). Variations in cell type may in part be a reflection of the cytoarchitectural
differences that exist between the gray and white matterof the brain and
spinal cord. Electron micro- scopic features common to both types of as-
trocytes (Fig. 6.5) are the usual cytoplasmic organelles: dense mitochondria,
scanty rough and smooth endoplasmic reticulum, gliofila- ments, and an
abundant watery cytoplasm. The nucleus is finely granular and only mod-
erately dense, and nuclear pores have been identified, as shown by Palay (82)
and Maxwell and Kruger (69). Fine structural characteristics that identify the
astrocyte alone were described by the latter authors. These include a watery
cytoplasm that con- tains gliofilaments and dense glycogen gran- ules 15--40
nm in diameter. Furthermore, re- cent intracellular HRP experiments by
Sasaki and colleagues have shown that gray matter and white matter
astrocytes possess distinc6 Neuroglia 205tive morphologic features. Their
camera lucida reconstructions clearly show the diffecence between the very
long, threadlike processes of fibrous astrocytes in the white matter compared
with the shorter, sinuous processes with clusters or lamellar ap- pendages of
protoplasmic astrocytes in gray matter (107).

9. Olygodendrocytes –Types, Function, Distribution in CNS

10. Schwann Cells

11. Myelin Sheath-Formation in CNS and PNS, Protein in Myelin

12. Microglia, Brain Macrophages
13. Node of Ranvier, Schmidt-Lanterman’s Cleft
14. Blood-Brain Barrier
The blood-brain barrier (BBB) is a functional barrier that allows much tighter
control than that in most tissues over the passage of substances moving from
blood into the CNS tissue. The main structural component of the BBB is the
capillary endothelium, in which the cells are tightly sealed together with
well-developed occluding junctions, with little or no transcytosis activity, and
surrounded by the basement membrane. The limiting layer of perivascular
astrocytic feet that envelops the basement membrane of capillaries in most
CNS regions contributes to the BBB and further regulates passage of
molecules and ions from blood to brain. The BBB protects neurons and glia
from bacterial toxins, infectious agents, and other exogenous substances, and
helps maintain the stable composition and constant balance of ions in the
interstitial fluid required for normal neuronal function. The BBB is not present
in regions of the hypothalamus where plasma components are monitored, in
the posterior pituitary which releases hormones, or in the choroid plexus
where CSF is produced.

15. Central Nervous System -White Matter, Central Nervous System- Gray
Matter & Spinal Cord

The major structures comprising the CNS are the cerebrum, cerebellum, and
spinal cord. The CNS is completely covered by connective tissue layers, the
meninges, but CNS tissue contains very little collagen or similar material,
making it relatively soft and easily damaged by injuries affecting the
protective skull or vertebral bones. Most CNS neurons and their functional
organization are more appropriately covered in neuroscience rather than
histology courses, but certain important cells and basic topics will be
introduced here. Many structural features of CNS tissues can be seen in
unstained, freshly dissected specimens. Many regions show organized areas
of white matter and gray matter, differences caused by the differential
distribution of lipid-rich myelin. The main components of white matter are
myelinated axons, often grouped together as tracts, and the myelin-producing
oligodendrocytes. Astrocytes and microglia are also present, but very few
neuronal cell bodies. Gray matter contains abundant neuronal cell bodies,
 dendrites, astrocytes, and microglial cells, and is where most synapses occur.
Gray matter makes up the thick cortex or surface layer of both the cerebrum
and the cerebellum; most white matter is found in deeper regions. Deep
within the brain are localized, variously shaped darker areas called the
cerebral nuclei, each containing large numbers of aggregated neuronal cell
bodies.

16. Cerebellum (cortex, neurons, and glial cell types)

The sharply folded cerebellar cortex coordinates muscular activity throughout
the body and is organized with three layers:
■ A thick outer molecular layer has much neuropil and scattered neuronal cell
bodies.
■ A thin middle layer consists only of very large neurons called Purkinje cells
(named for the 19th century Czech histologist Jan Purkinje). These are
conspicuous even in H&E-stained sections, and their dendrites extend
throughout the molecular layer as a branching basket of nerve fibers.
■ A thick inner granular layer contains various very small, densely packed
neurons (including granule cells, with diameters of only 4-5 μm) and little
neuropil. In cross sections of the spinal cord, the white matter is peripheral
and the gray matter forms a deeper, H-shaped mass. The two anterior
projections of this gray matter, the anterior horns, contain cell bodies of very
large motor neurons whose axons make up the ventral roots of spinal nerves.
The two posterior horns contain interneurons which receive sensory fibers
from neurons in the spinal (dorsal root) ganglia. Near the middle of the cord
the gray matter surrounds a small central canal, which develops from the
lumen of the neural tube, is continuous with the ventricles of the brain, is
lined by ependymal cells, and contains CSF.
17. Meninges- Dura Matter
Dura Mater The thick external dura mater consists of dense irregular
connective tissue organized as an outer periosteal layer continuous with the
periosteum of the skull and an inner meningeal layer. These two layers are
usually fused, but along the superior sagittal surface and other specific areas
around the brain they separate to form the blood-filled dural venous sinuses.
Around the spinal cord the dura mater is separated from the periosteum of
the vertebrae by the epidural space, which contains a plexus of thin walled
veins and loose connective tissue. The dura mater may be separated from the
arachnoid by formation of a thin subdural space.
18. Arachnoid and Cerebro-Spinal Fluid Circulation
The arachnoid has two components: (1) a sheet of connective tissue in contact
with the dura mater and (2) a system of loosely arranged trabeculae
composed of collagen and fibroblasts, continuous with the underlying pia
 mater layer. Surrounding these trabeculae is a large, sponge-like cavity, the
subarachnoid space, filled with CSF. This fluid-filled space helps cushion and
protect the CNS from minor trauma. The subarachnoid space communicates
with the ventricles of the brain where the CSF is produced. The connective
tissue of the arachnoid is said to be avascular because it lacks nutritive
capillaries, but larger blood vessels run through it. Because the arachnoid has
fewer trabeculae in the spinal cord, it can be more clearly distinguished from
the pia mater in that area. The arachnoid and the pia mater are intimately
associated and are often considered a single membrane called the
pia-arachnoid. In some areas, the arachnoid penetrates the dura mater and
protrudes into blood-filled dural venous sinuses located there. These
CSF-filled protrusions, which are covered by the vascular endothelial cells
lining the sinuses, are called arachnoid villi and function as sites for
absorption of CSF into the blood of the venous sinuses.

19. Pia Matter

Pia Mater The innermost pia mater consists of flattened, mesenchymally
derived cells closely applied to the entire surface of the CNS tissue. The pia
does not directly contact nerve cells or fibers, being separated from the neural
elements by the very thin superficial layer of astrocytic processes (the glial
limiting membrane, or glia limitans), which adheres firmly to the pia mater.
Together, the pia mater and the layer of astrocytic end feet from a physical
barrier separating CNS tissue from CSF in the subarachnoid space. Blood
vessels penetrate CNS tissue through long perivascular spaces covered by pia
mater, although the pia disappears when the blood vessels branch to form the
small capillaries. However, these capillaries remain completely covered by the
perivascular layer of astrocytic processes

20. Chroid plexus (tela choroidea, choroid epithelium)

The choroid plexus consists of highly vascular tissue, elaborately folded and
projecting into the large ventricles of the brain (Figure 9–20a). It is found in
the roofs of the third and fourth ventricles and in parts of the two lateral
ventricular walls, all regions in which the ependymal lining directly contacts
the pia mater. Each villus of the choroid plexus contains a thin layer of
well-vascularized pia mater covered by cuboidal ependymal cells (Figure
9–20b). The function of the choroid plexus is to remove water from blood and
release it as the CSF. CSF is clear, contains Na+, K+, and Cl– ions but very
little protein, and its only cells are normally very sparse lymphocytes. It is
produced continuously and it completely fills the ventricles, the central canal
of the spinal cord, the subarachnoid and perivascular spaces. It provides the
ions required for CNS neuronal activity and in the arachnoid serves to help
absorb mechanical shocks. Arachnoid villi (Figure 9–19) provide the main
 pathway for absorption of CSF back into the venous circulation. There are
very few lymphatic vessels in CNS tissue.

21. Cerebrospinal Fluid

22. Nerve Fibers

23. Myelinated and Unmyelinated Fibers
24. Connective Tissue Sheets
25. Ganglia (sensory, autonomic ganglia)
26. Autonomic Nervous System

27. Merkel’s Cell

28. Vater-Pachinian Corpuscle
29. Meissnerian Corpuscle
30. Epidermis Organization
31. Keratinocytes Types
32. Melanocytes and Development of Melanin Granule
33. Skin Macrophages- Lagerhans’ Cell
34. Cells of Dermis
35. Dermal Fibrous Tissue Types ( elastic, collagen, distribution in body)
36. Subcutaneus Tissue-Hairs
37. Subcutaneus Tissue- Sebaceous Glands
38. Sweat Gland Types ( eccrine, apocrine, innervations, cell types, distribution
in body)
39. Nail Matrix, Nail Plate, Nail Bed
40. Photoreceptor System –The Eye, External Layers of Eye (Fibrous)
Eyes (Figure 23–1) are highly developed photosensitive organs for analyzing
the form, intensity, and color of light reflected from objects and providing the
sense of sight. Protected within the orbits of the skull which also contain
adipose cushions, each eyeball consists externally of a tough, fibrous globe,
which maintains an eye’s overall shape. Internally the eye contains
transparent tissues that refract light to focus the image, a layer of
photosensitive cells, and a system of neurons that collect, process, and
transmit visual information to the brain. Each eye is composed of three
concentric tunics or layers (Table 23–1):
■ A tough external fibrous layer consisting of the sclera and the transparent
cornea;
■ A middle vascular layer consisting of the choroid, ciliary body, and iris; and
■ An inner sensory layer, the retina, which communicates with the cerebrum
through the posterior optic nerve

41. Sclera and Corneal Epithelium

Sclera The fibrous, external layer of the eyeball protects the more delicate
internal structures and provides sites for muscle insertion (see Table 23–1).
The white posterior five-sixths of this layer is the sclera (Figure 23–1), which
encloses a portion of the eyeball about 22 mm in diameter in adults. The sclera
averages 0.5 mm in thickness and consists mainly of dense connective tissue,
with flat bundles of type I collagen parallel to the organ surface but
intersecting in various directions; microvasculature is present near the outer
surface. Tendons of the extraocular muscles which move the eyes insert into
the anterior region of the sclera. Posteriorly the sclera thickens to
approximately 1 mm and joins with the epineurium covering the optic nerve.
Where it surrounds the choroid, the sclera includes an inner suprachoroid
lamina, with less collagen, more fibroblasts, elastic fibers, and melanocytes.

Cornea In contrast to the sclera, the anterior one-sixth of the eye—the

cornea—is transparent and completely avascular (Figure 23–1). A section of
the cornea shows five distinct layers:
■ An external corneal epithelium, which is a stratified squamous epithelium;
■ An anterior limiting membrane (Bowman membrane), which is the
basement membrane beneath the corneal epithelium;
■ The thick stroma;
■ A posterior limiting membrane (Descemet’s membrane), which is the
basement membrane of the endothelium; and
■ An inner simple squamous endothelium.
The corneal epithelium is nonkeratinized, five or six cell layers thick, and
comprises about 10% of the corneal thickness (Figure 23–3). The basal cells
have a high proliferative capacity important for renewal and repair of the
corneal surface and emerge from stem cells in the corneoscleral limbus
encircling the cornea. The flattened surface cells have microvilli protruding
into a protective tear film of lipid, glycoprotein, and water. As another
protective adaptation, the corneal epithelium also has one of the richest
sensory nerve supplies of any tissue. The basement membrane of this
epithelium, often called Bowman membrane, is very thick (8-10 μm) and
contributes to the stability and strength of the cornea, helping to protect
against infection of the underlying stroma. The stroma, or substantia propria,
makes up 90% of the cornea’s thickness and consists of approximately 60
layers of parallel collagen bundles aligned at approximately right angles to
each other and extending almost the full diameter of the cornea. The uniform
orthogonal array of collagen fibrils contributes to the transparency of this
avascular tissue. Between the TABLE 23–1 Tunics of the eye. Structures
Components Function Fibrous tunic Sclera Cornea Retina Pigmented layer
Neural layer Vascular tunic Iris Ciliary body Choroid Fibrous Tunic (External
Layer) Sclera Dense irregular connective tissue Supports eye shape Protects
 delicate internal structures Extrinsic eye muscle attachment site Cornea Two
layers of epithelium with organized connective tissue in between Protects
anterior surface of the eye Refracts (bends) incoming light Vascular Tunic or
Uvea (Middle Layer) Choroid Areolar connective tissue; highly vascularized
Supplies nourishment to retina Pigment absorbs extraneous light Ciliary body
Ciliary smooth muscle and ciliary processes; covered with a secretory
epithelium Holds suspensory ligaments that attach to the lens and change
lens shape for far and near vision Epithelium secretes aqueous humor Iris
Two layers of smooth muscle (sphincter pupillae and dilator pupillae) and
connective tissue, with a central pupil Controls pupil diameter and thus the
amount of light entering the eye Retina (Internal Layer) Pigmented layer
Pigmented epithelial cells Absorbs extraneous light Provides vitamin A for
photoreceptor cells Neural layer Photoreceptors, bipolar neurons, ganglion
cells, and supporting Müller cells Detects incoming light rays; light rays are
converted to nerve signals and transmitted to the brain collagen lamellae are
cytoplasmic extensions of flattened fibroblast-like cells called keratocytes
(Figure 23–3). The ground substance around these cells contains
proteoglycans such as lumican, with keratan sulfate and chondroitin sulfate,
which help maintain the precise organization and spacing of the collagen
fibrils.
42. Descemet’s Membrane, Corneoscleral Junction or Limbus
The posterior surface of the stroma is bounded by another thick basement
membrane, called Descemet’s membrane, which supports the internal simple
squamous corneal endothelium (Figure 23–3). This endothelium maintains
Descemet’s membrane and includes the most metabolically active cells of the
cornea. Na+/K+ ATPase pumps in the basolateral membranes of these cells
are largely responsible for regulating the proper hydration state of the corneal
stroma to provide maximal transparency and optimal light refraction. Limbus
Encircling the cornea is the limbus, a transitional area where the transparent
cornea merges with the opaque sclera (Figures 23–1 and 23–4). Here
Bowman’s membrane ends and the surface epithelium becomes more
stratified as the conjunctiva that covers the anterior part of the sclera (and
lines the eyelids). As mentioned previously, epithelial stem cells located at the
limbus surface give rise to rapidly dividing progenitor cells, which then move
centripetally into the corneal epithelium. The stroma becomes vascular and
less well-organized at the limbus, as the collagen bundles merge with those of
the sclera. Also at the limbus Descemet’s membrane and its simple
endothelium are replaced with a system of irregular endothelium-lined
channels called the trabecular meshwork (Figure 23–5). These penetrate the
stroma at the corneoscleral junction and allow slow, continuous drainage of
aqueous humor from the anterior chamber. This fluid moves from these
channels into the adjacent larger space of the scleral venous sinus, or canal of
 Schlemm (Figures 23–1, 23–4, and 23–5), which encircles the eye. From this
sinus aqueous humor drains into small blood vessels (veins) of the sclera.

43. Vascular or Middle Layer –Choroid, Bruch’s Membrane

Choroid Located in the posterior two-thirds of the eye, the choroid consists of
loose, well-vascularized connective tissue and contains numerous
melanocytes (Figure 23–6). These form a characteristic black layer in the
choroid and prevent light from entering the eye except through the pupil.
Two layers make up the choroid (Figure 23–6):
■ The inner choroidocapillary lamina has a rich microvasculature important
for nutrition of the outer retinal layers.
■ Bruch membrane, a thin extracellular sheet, is composed of collagen and
elastic fibers surrounding the adjacent microvasculature and basal lamina of
the retina’s pigmented layer.
44. Middle Layer of Eye – Ciliary body, Ciliary Processes, Iris
The ciliary body, the anterior expansion of the uvea that encircles the lens, lies
posterior to the limbus (Figures 23–1 and 23–4). Like the choroid, most of the
ciliary body rests on the sclera. Important structures associated with the
ciliary body include the following:
■ Ciliary muscle makes up most of the ciliary body’s stroma and consists of
three groups of smooth muscle fibers. Contraction of these muscles affects the
shape of the lens and is important in visual accommodation (see Lens).
■ Ciliary processes are a radially arranged series of about 75 ridges extending
from the inner highly vascular region of the ciliary body. These provide a
large surface area covered by a double layer of low columnar epithelial cells,
the ciliary epithelium (Figure 23–7). The epithelial cells directly covering the
stroma contain much melanin and correspond to the anterior projection of the
pigmented retina epithelium. The surface layer of cells lacks melanin and is
contiguous with the sensory layer of the retina.
■ Cells of this dual epithelium have extensive basolateral folds with
Na+/K+-ATPase activity and are specialized for secretion of aqueous humor.
Fluid from the stromal microvasculature moves across this epithelium as
aqueous humor, with an inorganic ion composition similar to that of plasma
but almost no protein. As shown in Figure 23–8, aqueous humor is secreted by
ciliary processes into the posterior chamber, flows through the pupil into the
anterior chamber, and drains at the angle formed by the cornea and the iris
into the channels of the trabecular meshwork and the scleral venous sinus,
from which it enters venules of the sclera.
■ The ciliary zonule is a system of many radially oriented fibers composed
largely of fibrillin-1 and -2 produced by the nonpigmented epithelial cells on
the ciliary processes. The fibers extend from grooves between the ciliary
 processes and attach to the surface of the lens (Figure 23–9), holding that
structure in place.
45. Lens, Vitreous Body, Cavum Vitreum
Lens The lens is a transparent biconvex structure suspended immediately
behind the iris, which focuses light on the retina (Figure 23–1). Derived from
an invagination of the embryonic surface ectoderm (see Figure 23–2), the lens
is a unique avascular tissue and is highly elastic, a property that normally
decreases with age. The lens has three principal components:
■ A thick (10-20 μm), homogeneous lens capsule composed of proteoglycans
and type IV collagen surrounds the lens (Figure 23–11) and provides the place
of attachment for the fibers of the ciliary zonule (Figure 23–10). This layer
originates as the basement membrane of the embryonic lens vesicle.
■ A subcapsular lens epithelium consists of a single layer of cuboidal cells
present only on the anterior surface of the lens (Figure 23–11). The epithelial
cells attach basally to the surrounding lens capsule and their apical surfaces
bind to the internal lens fibers. At the posterior edge ofthis epithelium, near
the equator of the lens, the epithelial cells divide to provide new cells, which
differentiate as lens fibers. This process allows for growth of the lens and
continues at a slow, decreasing rate near the equator of the lens throughout
adult life.
■ Lens fibers are highly elongated, terminally differentiated cells that appear
as thin, flattened structures (Figure 23–11). Developing from cells in the lens
epithelium, lens fibers typically become 7-10 mm long, with crosssection
dimensions of only 2 by 8 μm. The cytoplasm becomes filled with a group of
proteins called crystallins, and the organelles and nuclei undergo autophagy.
Lens fibers are packed tightly together and form a perfectly transparent tissue
highly specialized for light refraction. The lens is held in place by fibers of the
ciliary zonule, which extend from the lens capsule to the ciliary body (Figures
23–1 and 23–9). Together with the ciliary muscles, this structure allows the
process of visual accommodation, which permits focusing on near and far
objects by changing the curvature of the lens (Figure 23–12). When the eye is
at rest or gazing at distant objects, ciliary muscles relax and the resulting
shape of the ciliary body puts tension on the zonule fibers, which pulls the
lens into a flatter shape. To focus on a close object the ciliary muscles contract,
causing forward displacement of the ciliary body, which relieves some of the
tension on the zonule and allows the lens to return to a more rounded shape
and keep the object in focus. In the fourth decade of life presbyopia (Gr.
presbyter, elder + L. opticus, relating to eyes) normally causes the lenses to
lose elasticity and their ability to undergo accommodation.
46. Retina –Neuronal Types and Functions

47. Photoreceptor, Pigment Epithelium

48. Retinal Histophysiology
49. Accessory Structures of The Eye
50. The Audio Receptor System - External Ear
51. Middle Ear- Tympanic Cavity
52. Internal Ear – Organ of Corti
53. Sensory Cells in Cochlear Duct and Labyrinth –Hair Cell Types,
Differentiation Patterns
54. Saccule and Utricle
55. Spiral Lamina- Endolymph, Perilymph, Contacts with CNS Meninges
56. Cochlear Duct - Hair Cells, Supporting Cell Types