Histology lecture: Connective Tissue

Connective Tissue
 Provides nutrients and blood supply to the
epithelium
 Binds epithelium to other tissues
 Protects underlying tissues (to be discussed
later)
 Immunologic functions (to be discussed later)

Epithelium vs Connective Tissue

 Epithelium is primarily made up of tightly
packed cells with less amount of fibers and
ground substances
 The major constituent of the connective tissue
is the extracellular matrix (fiber and ground
substances)
*Blue arrow: lining epithelium – tightly packed
together
*Yellow arrow: connective tissue – have spaces
between cells

 Just like the rest of the tissues in the body, it is Ground Substance
formed by three classes of components:
o Cells
o Extracellular Matrix
 Fibers
 Ground substance

 Background material where the cells and fibers

of connective tissue are embedded
 Composed of chemicals that binds:
o cells of the connective tissue together
o cells to the fiber
*Topmost arrow: cell o fiber to another fiber
*Second arrow: fiber  Molecules that stabilize the extracellular
*Third arrow: ground substance matrix by binding to receptors on the surface
of cells and other matrix compositions

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Histology lecture: Connective Tissue

 Functions of ground substance

o Fills space between cells and fibers
and links them  The different joints in the body bines articulate
o Provide venue for exchange of with each other and for these bones not to
substances crush each other, the distal end of one bone
o Store water and the proximal end of the other bone should
o Barrier to infection be covered with articular cartilage (blue
colored structure in the picture)
 As the bone articulate against each other they
are creating compression forces against the
cartilage. What enables cartilages to withstand
the compression brought by the articulation, is
because the cartilage is rich in water, and what
stores water in the cartilages is the ground
substance

 Basement membrane – structure that separate

epithelium and connective tissue
 The area where the oxygen and the nutrients
will diffuse into so that they can reach the cells
of the epithelium and the cells of the
connective tissue is the ground substance

 Water, glucose, and amino acids are diffusing

out of the capillary and they are diffusing
towards the cells
 It is in the ground substance where the water,
glucose, and amino acids are flowing into
 Waste products and carbon dioxide are
diffusing away from the cells into the capillary
via the ground susbtance

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Histology lecture: Connective Tissue

 If there is sharp object that caused a break in

the skin, in this case the stratified squamous
keratinized epithelium cannot anymore
perform its function, which is to provide
protection, and there is a subsequent
introduction of microorganisms on the
underlying connective tissue, these organisms
can spread to other tissue in the skin and cause
destruction
 Microorganism introduced into the underlying
connective tissue were retained in the area
where they were introduced because it needs
to wait for the white blood cells to reach the
site of infection and kill them, and this is
because of the ground substance, which retain
the microorganisms to stay where they are
 But there are microorganisms that can break
ground substance, if this happens expect that
the infection will become deep seated

 Glycosaminoglycans are linked to the core

proteins
 Proteoglycans – composed of
glycosaminoglycans linked to the core proteins
6 proteoglycans are linked to linker proteins
 Linker proteins – links proteoglycans to the
hyaluronic acid

Ground Substance Glycosaminoglycans are long polysaccharides

 Composed of (composed of monosaccharides linked together by
o (1)polysaccharides glycosidic bonds) composed repeating
(glycosaminoglycans, proteoglycans) disaccharide units
o (2) proteins  Present monosaccharides (alternately
arranged)
o uronic acid (first)
o hexosamine

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Histology lecture: Connective Tissue

Uronic acid Acid and Hexosamine

 Uronic acid
o Glucoronic acid
o Iduronic acid
 Hexosamine
o Glucosamine
o Galactosamine

Uronic acid plus Hexosamine

 Proteoglycans are composed of

glycosaminoglycans that are covalently
attached to the core proteins
 Glycosaminoglycans present in connective  Attached to the hyaluronic acid by the linker
tissue; proteins
o Chondroitin sulfate
 Glucuronic acid
 galactosamine
o Dermatan sulfate
 Iduronic acid
 galactosamine
o Heparan sulfate
 Glucuronic acid
 glucosamine
o heparin
 iduronic acid
 glucosamine  Glycosaminoglycans
Long polysaccharides composed repeating o Chondroitin sulfate
disaccharide units (uronic acid and hexosamine) o Dermatan sulfate
o Keratan sulfate
o Heparan sulfate
o Hyaluronic acid - largest
o All linked to proteins to form
proteoglycans EXCEPT hyaluronic acid

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Histology lecture: Connective Tissue

o Most abundant cell of the connective

tissue
o Synthesizes fibers and ground substance
of the connective tissue
o Responsible for the production of the
extracellular matrix of the connective
tissue
 Others are from bone marrow
 Don’t contribute to the formation of the
 There is bacterium that can produce connective tissue, they will just migrate from
hyaluronidase that can degrade the ground the bone marrow and establish themselves in
substance the connective tissue
 Hyaluronidase – enzyme that can break down o Mast cells
hyaluronic acid , if this is introduces into the o Macrophages
ground substance of the connective tissue o Plasma cells (B Lymphocytes)
expect a deep seated infection o Neutrophils
Ground Substance
 Composed of
o (1)polysaccharides (glycosaminoglycans,
proteoglycans)
o (2) proteins (Fibronectin) – also attaches
cells to cells or cells to fibers

Fibroblast
 Most abundant cell of the connective tissue
Collagen fiber component of the connective tissue  Cells that have almost the same shape in the
 Fibronectin connective tissue are most likely firbroblast
o links cell to the collagen fiber  Synthesize collagen and elastic fibers,
o similar to glycosaminoglycans and glycosaminoglycan, proteoglycans and
proteoglycans, just that the fibronectin is multiadhesive proteins.
a protein  Migrates through the connective tissue,
secreting the fibers and components of the
ground substance
 Synthesize collagen and elastic fiber,
glycosaminoglycan, proteoglycans and
multiadhesive proteins.
o Must be abundant in rough ER and
Golgi apparatus
o Must have more euchromatin because
of the active DNA transcription

Cells of the Connective Tissue

 Fibroblasts

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Histology lecture: Connective Tissue
Most of them are protein
Nucleus (electron microscope view)
* green = nucleolus
*blue – heterochromatin – contains dna (tightly
wrapped around with histones, that is why it
appears darker)
*red – chromatin – contains dna
Fibroblast (active) – should appear lighter
Fibrocyte (quiescent)
Left: fibroblast | right: fibrocyte
 Higher amounts of euchromatin
 Light stained nucleus
 More developed RER and Golgi complex
 Irregular cytoplasm
Egg cells are produced and released from
ovary and these egg cells are captured by
fallopian tubes. It is in the fallopian tube that
egg is fertilized by the sperm. The cilia lining
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epithelium of the fallopian tube will proper the
fertilized ovum towards the uterine cavity for
implantation. The embryo will develop and it will
produce three important layers and these layers are
responsible for giving rise to the future organs of
the embryo.
1. Ectoderm – most superficial
2. Mesoderm- middle
3. Endoderm – innermost layer
Ectoderm
 Superficial one, so expect that this is where
our skin will be derived from and makes us
feel that our skin is in contact with a rough
surface or if we are holding a hot object is our
nervous system
Endoderm
 Responsible for giving rise to the internal
organs, the stomach, small intestine,
gallbladder, liver, urinary bladder
Mesoderm
 muscle, bones, cartilage, and even the
connective tissue somewhere in the middle of
the body
 They are found between skin and the internal
organs and all of these tissues and organs are
derived from the mesoderm
 Cells in the mesoderm are referred as
mesenchymal cells
Mesenchymal cells of the Mesoderm
 Can become the future chondrocytes of the
cartilage, osteoblast of the bone, myocytes of
the the muscle, adipocyte of the adipose tissue,
the and the fibroblast od the connective tissue
the
the
Histology lecture: Connective Tissue

1. Mesenchymal cells will become the fibroblast

2. Fibroblasts will start producing the
extracellular matrix
3. After producing extracellular matrix and
building gup the connective tissue, the cells
form the bone marrow will start residing in the
connective tissue

The Other cells

Other cells in the connective tissue are all
produced in the bone marrow and migrate towards
the connective tissue and establish residence there
Neutrophils
 First white blood cell that will go out of the
blood vessel and migrate towards the site of
infection
 Most sensitive to chemotactic stimuli, these
chemotactic stimuli are any chemical released
by the microorganisms introduced into the site
of infection
Neutrophils
 First cells to reach the site of infection
 Phagocytosis of bacteria – engulf foreign
material and caused them to be digested
 Neutrophilia = >70% count means bacterial
infection
o Neutrophil count
o Expected 50-70

1-2 days after activation = apoptosis

 60% (50-70) of the white blood cells;

 Most abundant white blood cells
 3-5 nuclear lobes
 Also known as segmenters or
polymorphonuclears
 Nuclei in neutrophils are not spherical instead Neutrophils die after 1-2 days of activation
they are lobulated, they can have 3 to 5 lobes

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Histology lecture: Connective Tissue
 Neutrophils undergo apoptosis after 1-2 days
of activation
o Pus
 REPLACED by monocytes (phagocytes)
Macrophages are derived from monocytes.
Monocytes and macrophages are produced in the
bone marrow. They will emerge out of the bone
marrow and go to the blood circulation as
monocytes. Some of the monocytes will remain
circulating in the blood while some of the them
will migrate towards the tissue and will now be
referred as the macrophages. Therefore,
macrophages is the tissue form of the monocyte.
Monocytes versus Macrophages
Monocyte and macrophages are produced in the
bone marrow. One It will go out the blood it will
be referred to as monocyte. Once it will migrate
towards the tissue and establish residence in there,
it will now be called as macrophage
Monocyte vs Neutrophil
Neutrophil
o Have cytoplasmic granules – rich in
digestive enzyme, meaning, every time the
neutrophil will engulf bacteria, the
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bacteria will surely die because of the
enzyme present in the granules
Monocyte and macrophages
o Agranulocytes – they do not have granules
in their cytoplasm
o Although they engulf bacteria faster than
neutrophil but it does not have granules
which contains digestive enzymes – their
ability to digest is not good as the
neutrophil
o Monocyte and macrophages will be able to
help us fight infection through antigen
presentation
Antigen Presentation
 Monocytes and macrophages will act an
antigen presenting cells, meaning, they will
present the antigen or the microorganisms to
another cell
 Monocyte or macrophage is engulfing a
pathogen or a microorganism
 While inside the cytoplasm of the monocyte
and the macrophage, the monocyte or
macrophage will select the most antigenic or
immunogenic portion of that pathogen
 The pathogen will be broken into pieces -
because of the monocyte or macrophage
 The most immunogenic or antigenic portion of
the pathogen will be displayed on the surface
of the macrophage
 After the antigen is presented on the surface of
the antigen presenting cells or the
monocyte/macrophage, this antigen now will
be presented to T helper cell
Histology lecture: Connective Tissue
 In this picture, there are four bacteria and only
one was engulfed by the antigen presenting
cells or the monocyte/macrophage
 In the cytoplasm of the antigen presenting cell,
one of the antigens will be loaded on a protein
and the antigen and the protein will be
displayed on the surface of the antigen
presenting cell
What happens to the three remaining bacteria?
 A microbe is about to be engulfed by the
antigen presenting cell, while inside the
cytoplasm of the antigen presenting cell one of
the antigens of the microbe will be displayed
on the surface of the antigen presenting cells
and the antigen now will be presented to T
helper cell
The purpose of activating T helper cells:
 T cell will activate another cell, the B cell
 B cell is known to differentiate to form plasma
cells and these cells are now responsible for the
production of antibodies
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 What will cause the death of the three bacteria
is the antibodies produced by the plasma cells
– because the plasma cells were formed by the
b cells – which was activated by the T helper
cell – were activated by the antigen presenting
cells which engulfed one of the four bacteria
Antigen Presentation
 Macrophage or monocytes phagocytose
extracellular microorganisms and present a
part of these organisms to T cells (T helper)
 T helper cells, in turn, stimulate B cells to
become plasma cells and to produce antibodies
MACROPHAGES IN DIFFERENT TISSUES
 Connective tissue – histiocytes
 Liver – Kupffer cells
 Lung – alveolar macrophages/ dust cells
 Bone – osteoclasts
 GloMErulus of the kidney – MEsangial cells
 CNS – microglia or microglials cells
 Skin – Langerhans cells
 Lymph node – Follicular dendritic cells
Plasma cells
 Activated form of b cells
Plasma cells
Histology lecture: Connective Tissue

 Synthesizes antibodies
 Antibodies are proteins
 Thus, plasma cells have high amount of
euchromatin, well developed rough ER and
Golgi apparatus

Basophil
 one of the white blood cells
 Nucleus is generally spherical but  The dark blue granules contain histamine
eccentrically placed with heterochromation  Leave the bone marrow mature, that’s why
alternating with lighter areas of euchromatin they spend most of their life span circulating in
 Within the nucleolus of the plasma cells there the blood
is the alternate arrangement of
heterochromatin and euchromatin, making the Tissue: Mast cells
nucleus appear to have a clock face
appearance
Plasma cells with Eccentric nucleus

 Leave bone marrow immature and will only

become mature once it reside in the tissue
Clock face  Large, oval and round cells
 Alternate arrangement of heterochromatin  Leaves the bone marrow immature and
and euchromatin matures in the tissue
 Th ones labeled 369 and 12 are the hetero  Basophilic Secretory Granules which contain
chromatin, notice that between them are HISTAMINE
light staining euchromatin
Mast Cells
 In old literatures, mast cells are described as
the tissue forms of basophils
 In latest researches and study they found out
that mast cells are different from basophils,
although they contain the same chemicals in
their granules

Pathogenesis of Allergies

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Histology lecture: Connective Tissue

 Allergen will be engulfed by antigen

presenting cells and since these are antigen
presenting cells except that it will present
antigen to T cell particularly
 T cell will activate b cells
 B cell will differentiate to form plasma cells
 In the case of allergens, the antibody produces
is IgE Several more exposures
 If IgE are readily available, they will display
themselves using their tail potion, the tail
portion of the y-shaped antibody antibody
called FC
 The FC or the tail portion of the IgE will bind
to the receptors present on the surface of mast
cells
First exposure

 Crosslinking will trigger the mast cell and

basophil to release the histamine and the
histamine would have different effects on the
different organs in the body

Smooth Muscle Bronchoconstriction

Cell
Small blood Vasodilation and increased
vessel permeability- leakage of fluid –
hypotension – anaphylactic
shock!
Allergies need several re-exposures before
histamine is released Mucous gland Increased secretion – similar to
allergic rhinitis
 Crosslinking of at least 2 IgE’s causes the mast
Sensory-nerve Pruritus
cell to degranulate and to release histamine
endings
 Crosslinking – one allergen will crosslink to 2
IgEs at the same time
Mechanism
1. First exposure to allergen
First exposure
 Allergen stimulates formation of
antibody (Ig E type)
 IgEs are displayed on the surface mast
cells and basophils
2. Second exposure to the same allergen
 Allergen crosslinks at least 2 IgE
Second exposure: No crosslinking molecules leading to mast cell
activation and degranulation of mast
cells and release of mediators

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Histology lecture: Connective Tissue

Eosinophils

 Bilobed nucleus with orange-red colored

granules
 Has affinity to acid dyes (eosin)
 Major basic protein
o Toxic to parasites
o Antiparasitic – eosinophil count will
go high if there is parasitism

 The tail of a large parasitic worm, the
antibodies covering it since the eosinophil
cannot do phagocytosis and engulf the large
parasite.
 The eosinophil will just simply bind to the
antibodies found on the surface of the parasite
and the eosinophil will be triggered to release
the major basic protein component of its
granules
Eosinophils
 Antibody-dependent cell cytotoxicity (ADCC)
o Eosinophils only degranulate if there
is presence of antibody on the surface
of the parasite
Fibers
 Collagen fibers – if the organ need to be tough
and strong
 Reticular fibers – if the organ is soft and
contains a lot of cells
 Elastic fibers – if the organs is always
subjected to bending, stretching, and must go
back to the original size after being stretched
Elastic fibers
 Found in organs that are subjected to bending
and stretching
 Skin, lungs, aorta and its branches and
pulmonary artery
 The lungs must be provided with elastic fiber,
so that the lungs can expand whenever we
inhale air and must be able to go back to its
original size after exhalation

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Histology lecture: Connective Tissue

3 Reticular fibers Provides support

 The ventricle is more muscular than the
atrium, so expect that the ventricle would
generate a more powerful and more forceful
contraction than that of the atrium. So, blood
vessels that should receive the blood coming
from the ventricle should accommodate the
force and the large volume of blood, so, these
blood vessel should dilate and stretch beyond
their original size
 Pulmonary artery – received the blood coming
from the right ventricle
 Aorta – received the blood coming from the
left ventricle
Cut section of the Aorta
to parenchyma of
organs
4 Basement Sheet-forming
membrane collagen
7 Basement Anchors basal
membrane lamina to
reticular lamina
o Collagen type 1 in the skin along with other
types of collagen fibers will contribute to the
structural stability of our skin. If skin would be
wounded, the fibroblast in the connected
connective tissue will proliferate and this
fibroblast will deposit collagen type 1 in
attempt to repair. There will be scar after, the
scar is rich in collagen type 1, that is why
whenever you try to touch a scar in you skin it
would fell a little harder compared to the
normal skin because of the abundance of
collagen type 1.
o Collagen type 2 is also strong type of fiber,
similar to collagen type 1. The difference of
type 2 from type 1 is that these fibers are
found in organs that are tough, strong, and
rigid but at the same time retain and store
water. Ex: Cartilages
Collagen type 1: Tendon
 Tendon functions to connect the skeletal
muscle to the bone if the skeletal muscle will
contract. It will become shorter, and it will pull
the tendon, the tenson, on the other hand, will
pull the bone to generate the expected body
movement.
 Tendon should be reached in collagen type 1
so they will become strong and will not torn
every time it is pulled by the skeletal muscle

Labeled 2 are elastic fibers

Collagen
 Functions to provide rigidity and strength
 Most abundant protein in humans – 30% dry
weight
 Synthesized by the fibroblast of the connective Collagen type 1: Dentin
tissue  The portion of the tooth that you can see when
you smile is the crown
Types of Collagens  The portion of the tooth that is already
Collagen Tissues Function embedded at the level of the gums is the neck
1 Bone, Tendon, Resistance to  The portion of the tooth that is embedded on
Skin, Dentin Tension the sockets in the maxillary or mandibular
2 Cartilage, Resistance to bone is the root
Vitreous body Pressure  The protective layers of the tooth

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Histology lecture: Connective Tissue

oCrown is covered by both enamel and

dentin
o The root and neck are both covered
and protected by cementum and dentin
o Dentin protects all three portions of
the tooth and must be rigid and tough
to do
 Dentin should be provided with collagen type
1

Collagen IV
Collagen type 4 forms part of the basement
membrane particularly of the basal lamina and
reticular lamina

Collagen type 2: Vitreous chamber of the Eye

 Vitreous chamber of the eye – the large cavity
that you can find at the eye, posterior portion
of the eye
 Vitreous chamber contains high amount of
vitreous humor – which is a fluid
 Whenever there is fluid there is water, vitreous
chamber functions to keep the retina, the
yellow colored portion of the eye to attached
to the macula to prevent the occurrence of
blindness and for the vitreous chamber must
be tough and rigid structure, therefore it must
have collagen fiber and since there is water
within the vitreous chamber the collagen fiber
should be present is a collagen type 2

Collagen type 2: Cartilage

Collagen type 7 – functions to anchor basal lamina

to the reticular lamina, also called anchoring fibril
Tendon
Tendons – function to connect muscle to the bone

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Histology lecture: Connective Tissue

Abundant collagen fibers in single direction Dense

Regular Connective

Dura mater must protect the brain from multiple

Ligaments directions because force and trauma can come
Ligaments functions to link bones together from any direction
Dura mater of the brain: Dense irregular
connective tissue

Aponeurosis
Aponeurosis functions to connect muscle to
Capsule of organs
muscle

The aponeurosis in this picture is attaching

occipitalis to the frontalis muscle

Therefore, ligaments, aponeurosis, tendons and

corneal stroma are: Dense Regular Connective
tissue
Liver’s capsule – Glisson’s capsule
Dura mater of the brain
The brain and spinal cords are protected by bones Skin: Reticular layer of dermis
and meninges. One of the layers of the meninges
of the brain and spinal cord is the dura meter

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Histology lecture: Connective Tissue

difference between them is their affinity to H

and E staining

Tunica Albuginea of the Testes

Reticular Fibers
o B = Bone Marrow
o E = Endocrine glands
o L = Lymph Nodes/
o L = Liver
o Pa = Pancreas
o T = Thymus
o S = Spleen

Liver : H and E staining

Liver : Silver staining

Reticular fibers
 Contains a lot of cells
Bone Marrow: H and E
 Also referred to as collagen type 3
 NOT stained with H and E
o Collagen fibers are stained with pink
color by H and E
 Requires special stain – silver – imparts black
color
 Scientist realize reticular fibers are similar to
collagen fibers because they found out that the
chemical composition of the collagen fibers Bone Marrow: Silver Stain
are similar to the reticular fibers and the only

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Histology lecture: Connective Tissue
TYPES OF CONNECTIVE TISSUES
 Embryonic Connective Tissue
o Mesenchyme
o Mucous connective Tissue
 Mature Connective Tissue
Embryonic Connective Tissue
 CELLS <<< GROUND SUBSTANCE >>>
FIBERS
 The ground substance predominates over the
cells and the fiber component
Mesenchyme
 Mesenchyme is characterized morphologically
by a prominent ground substance matrix
containing a loose aggregate of reticular fibers
and unspecialized mesenchymal stem cells
 Mesenchymal cells are usually irregular in
shape
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 Mesenchymal cells give rise to cells that we
can find in the adult connective tissue
 Mesenchymal cells are supported by reticular
fibers
 The loose fluid nature of mesenchyme allows
its cells to migrate easily and play a crucial
role in the origin and development of
morphological structures during the embryonic
and fetal stages
o Mesenchyme directly gives rise to
most of the body’s connective tissues,
from bones and cartilage and muscles
Mucous Connective Tissue
 CELLS <<< GROUND SUBSTANCE >>>
FIBERS
Ground substance predominate
 Contains large amount of ground substances
with sparse amount of collagen fibers and
fibroblasts
 Functions to provide support
 Wharton’s jelly of the umbilical cord
Histology lecture: Connective Tissue

 Th umbilical cord connects the fetal

circulation to the placenta then to the maternal
circulation, so, this umbilical cord helps the
baby get nutrient and oxygen from his/her
mother
 Umbilical cord has 2 arteries and 1 vein – all
surrounded by the Wharton’s jelly – support
the 3 blood vessel

*left: loose connective tissue

*right: dense connective tissue
Dense Connective Tissue
 CELLS = GROUND SUBSTANCE <<<
FIBERS
 Fiber predominate cells and ground substance

Loose Connective Tissue

 CELLS = GROUND SUBSTANCE =/>
FIBERS
 Cells and ground substance predominate fiber

 Lesser amount of Fiber component

 Predominance of Ground Substance and Cells
 Has two types
o AREOLAR
o RETICULAR
o ADIPOSE
Areolar Connective Tissue
CELLS = GROUND SUBSTANCE = FIBER

TYPES OF MATURE CONNECTIVE TISSUE

 Loose connective tissue
 Dense connective tissue

 Contains fibers, ground substances and cells in

roughly equal parts

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Histology lecture: Connective Tissue

o Found underneath the epithelium Connective tissue in the gastrointestinal tract

o Lamina propria of the digestive tract o Mucosa
o Surrounds blood vessels and nerves o Three layers
 Epithelium – in contact
with the food
 Connective tissue –
lamina propria –
composed of areolar
connective tissue
 Muscularis mucosae
o Submucosa – connective tissue – dense
irregular connective tissue

Skin – dense irregular connective tissue

Papillary dermis – loose connective tissue areolar

Reticular Connective Tissue

• CELLS >>> GROUND SUBSTANCE =
FIBERS
Cells predominate

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Histology lecture: Connective Tissue

 Forms the stroma of the bone marrow, and soft Dense Connective Tissue
organs such as endocrine glands, liver, lymph  Very abundant fibrous components and
nodes, pancreas and thymus relatively few cells and ground substance
 Cells in these organs are supported by reticular  3 types
fibers o Elastic Connective Tissue
o Dense irregular CT
o Dense regular CT

Elastic Connective Tissue

Aorta

Adipose Tissue
 Composed primarily of adipose cells which
Dense regular connective tissue
can be found isolated or in groups within loose
connective tissue.  Collagen fibers are oriented parallel to one
another or in ordered arrangement
 Tendons
o Connective tissue connecting muscles
to bone
 Aponeurosis
o Connective tissue connecting muscle
to muscle
 Ligaments
o Connective tissue connecting bone to
bone
 Corneal Stroma

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Histology lecture: Connective Tissue

Dense Irregular Connective Tissue

 Collagen bundle fibers are randomly oriented
 Found in the
o reticular dermis of the skin
o capsules of organs
o tunica albuginea of testes
o dura mater of the brain
o submucosa of the digestive tract

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