Subject: Zoology

Lesson: Connective Tissues

Lesson Developer: Dr. Zubeda
Department: Department of Zoology,
Delhi University

Institute of Lifelong Learning, University of Delhi 1

Table of Contents

INTRODUCTION

GENERAL FEATURES OF CONNECTIVE TISSUE

LOCATIONS OF CONNECTIVE TISSUES

FUNCTIONS OF CONNECTIVE TISSUES

COMPONENTS OF CONNECTIVE TISSUE

(I) Specialized cells in Connective Tissue
1. Fibroblasts
2. Macrophages
3. Plasma cells
4. Mast cells
5. Adipocytes
6. Leukocytes
(II) Extracellular Matrix in Connective Tissue
i. Ground Substance
ii. Connective Tissue Fibers
a. Collagen fibres
b. Elastic fibres
c. Reticular fibres
CLASSIFICATION OF CONNECTIVE TISSUES

A. EMBRYONIC CONNECTIVE TISSUES

i. Mesenchyme

ii. Mucous connective tissue

B. MATURE CONNECTIVE TISSUES

1. CONNECTIVE TISSUE PROPER

I. Loose Connective Tissues

i. Areolar connective tissue

ii. Adipose connective tissue

Institute of Lifelong Learning, University of Delhi 2

 iii. Reticular connective tissue

II. Dense Connective Tissues

i. Dense regular connective tissue
ii. Dense irregular connective tissue
iii. Elastic connective tissue

2. SUPPORTING CONNECTIVE TISSUE

I. Cartilage
i. Hyaline cartilage
ii. Fibrous cartilage
iii. Elastic cartilage

II. Bone Tissue

i. Compact bone

ii. Spongy bone

3. FLUID CONNECTIVE TISSUE

I. Blood Tissue
II. Lymph

Summary

Exercises

Glossary

References

Institute of Lifelong Learning, University of Delhi 3

Learning objectives

 To discuss the common characteristics features of connective tissues.

 To describe the functions and location of connective tissues.
 To describe the different components (cells, fibres, ground substance) of
connective tissues.
 To classify the various types of connective tissues.
 Describe the structure, location and function of each type of connective
tissues.
 Describe the different cells found in connective tissue.

INTRODUCTION

Animal tissue is an assemblage of identical cells derived from the same origin which
is for performing specific function. Tissues of animal are classified into four major
categories: epithelial, connective, muscle and nervous tissues (Fig 1). All these
categories of tissues are organized to form organs and structures of animal body.
Although each of the animals usually has these four kinds of tissues, the appearance
and expression of all these tissues types may vary.

Connective tissue is one of the four major categories of animal tissues that supports,
joins and separate different forms of tissues and organs of the body. Connective
tissue is present all over the body excluding the central nervous system. Cells of the
connective tissue are more broadly apart from each other as compared to those in
epithelial tissues. Connective tissues contains larger amount of extra-cellular matrix
which is considerably absent in epithelial tissue.

Institute of Lifelong Learning, University of Delhi 4

 Fig. 1.
Four basic
types of tissues.
Source: http://cnx.org/content/m47980/latest/Picture

GENERAL FEATURES OF CONNECTIVE TISSUE

Followings are the general features of connective tissues which are different from the
other types of tissues of animals:

 Connective tissues are present in all part of the body excluding central nervous
system.
 These tissues are never exposed to the outer environment.
 In their various forms, connective tissues have a variety of functions.

 Most of the connective tissues contain a lot of blood vessels (means they are
highly vascular).

 They have receptors which can perceive pain, pressure, temperature, and other
sensations.

 Connective tissue fills spaces between organs and tissues, and provides structural
and metabolic support for other tissues and organs.

Institute of Lifelong Learning, University of Delhi 5

 Connective tissues consist of two chief components: extra-cellular matrix and
cells.

 Extra-cellular matrix in connective tissues is the substances located in spaces

between cells.

 The cells of connective tissues secrete extra-cellular fibres and account for the
many of the functional properties of the tissue in addition to controlling the
surrounding watery environment via specific proteoglycan molecules.

LOCATIONS OF CONNECTIVE TISSUES

Connective tissues are the most diverse and widely distributed tissues in the body.
Bone, blood, and fat are well-known connective tissues which perform many different
functions and constitute different properties.

FUNCTIONS OF CONNECTIVE TISSUES

Connective tissues provide a protective structural framework for other tissue types.
Functions of connective tissue are followed as:

1. Support and protection: The cells of connective tissue produce minerals and
fibers that make up the bony structural framework in the body. As a result, it
protects delicate organs, and covers and connects the other types of tissues.

2. Transport: Fluid connective tissues such as blood and lymph efficiently carry
substances from one area to another area of the body.

3. Storage: Adipose cells in the connective tissues store fats a form of energy until
it is required.

4. Defense: It involves defense against microbes through the intercellular

interactions and antibodies production by specialized cells of connective tissues.

COMPONENTS OF CONNECTIVE TISSUE

Main components of connective tissue are (I) specialized cells and (II)
extracellular matrix. The extracellular matrix is made up of fibres in a protein
and polysaccharide matrix, secreted and organized by cells in the extracellular
matrix. While epithelial tissues are mainly composed of cells, major volume of
connective tissues is occupied by the extracellular matrix. Variations in the
composition of the extracellular matrix, determines the properties of the connective
tissue. For example, if the matrix is calcified, it can form bone or teeth. Specialized
forms of extracellular matrix also make up tendons, cartilage, and the cornea of
the eye. General connective tissue is either loose, or dense, depending on the
arrangment of the fibres. The cells sit in a matrix made up of glycoproteins, fibrous
proteins and glycosoaminoglycans, which have been secreted by the fibroblasts, and
the major component of the matrix, is in fact, water.

Institute of Lifelong Learning, University of Delhi 6

(I) Specialized cells in Connective Tissue

Various cells of connective tissues produced from embryonic cells called

mesenchyme. Mesenchyme arises from mesoderm, an embryonic germ layer, and is
composed of stellate or fusiform cells embedded in a gelatinous ground substance.
Mesenchyme is the stem tissue of all the connective tissues of the body.

Each major type of connective tissue contains an immature class of cell with a name
ending in blast, which means to grow. These immature cells are termed fibroblasts in
case of loose and dense connective tissues while it is named as chondroblasts in
cartilage and osteoblasts in bone.

The types of cells in connective tissues vary according to the type of tissue (Fig. 2.)
and include the following:

1. Fibroblasts

These are bulky and flat cells containing branching projections. Fibroblasts contain
large and oval shaped nuclei with one or two conspicuous nucleoli. These are
distributed generally every connective tissues, and usually are the most abundant.
Fibroblasts migrate through the connective tissues, secreting the fibres and certain
components of the ground substance of the extracellular matrix.

In mature tissue, fibroblasts become dormant. When there is injury, they start to
produce new fibers. The active fibroblast contains a lot of rough endoplasmic
reticulum necessary for production of collagen and elastin fibers. Proteoglycan is a
main constituent of the ground substances in connective tissues and is produced
through fibroblasts.

2. Macrophages
It arises from a type of white blood cell termed as monocytes. Macrophages are not
in regular shape and have short branching processes. It is also called histiocyte.
After fibroblasts, macrophages are present in numerous numbers in loose connective
tissue. At first, these cells are not active. After inflammation, they intensively turn
into amoeboid and phagocytic stage termed as angry macrophages. They directly
engulf blood cells, bacteria, dead cells and debris digesting this material with
powerful enzymes. These cells are an important constituent of the reticuloendothelial
system (RES) positioned in the spleen, liver, lymph nodes and other organs.

Fixed macrophages exist in in a particular tissue; example includes alveolar

macrophages in the lungs or spleenic macrophages in the spleen. Wandering
macrophages have the ability to move throughout the tissue and gahter at sites of
infection or inflammation to carry on phagocytosis.

Institute of Lifelong Learning, University of Delhi 7

3. Plasma cells
These are small cells which arise from B lymphocyte, a type of white blood cell.
Plasma cells release antibodies and proteins that attack or reduce the effect of
foreign materials in the body. Although they are found in many places in the body,
most plasma cells reside in connective tissues especially in the gastrointestinal and
respiratory tracts. It plays an important role in body’s immune response.

4. Mast cells
These are large cells (20-30µm) packed with intensely basophilic granules that most
of the time obscure the nucleus. These are generally abundant around blood vessels.
These are very much similar to the blood basophils. So, these cells also have
mediators e. g. histamine, heparin and serotonin for instantaneous hypersensitivity.

5. Adipocytes
These are also called fat cells or adipose cells. These cells store triglycerides the form
of fat. They are prevalent underneath to the skin and around organs for example
heart and kidneys. These are frequently found arranged near to small blood vessels.
In the beginning these cells resemble to fibroblasts except they have abundant
vacuoles of fat droplets.

6. Leukocytes:
These are white blood cells that move along with the connective tissues around blood
vessels. In normal condition, these are less in numbers in connective tissue. But in
some conditions they move from blood into connective tissues. For instance,
neutrophils get together at region of infection and eosinophils move to region of
parasitic invasions and involved in the allergic reactions. Eosinophils are most
prevalent throughout the respiratory, digestive tracts and in active mammary tissue.

Fig. 2. Cells

Institute of Lifelong Learning, University of Delhi 8

in connective tissues.
Source: http://www.mhhe.com/biosci/ap/histology_mh/loosect.gif

(II) Extracellular Matrix in Connective Tissue

The cells of connective tissue are interspersed in a large volume of extra-cellular

matrix. This extra-cellular matrix (ECM) is secreted by the cells. It constitutes
protein fibers interpose in an amorphous mixture of large protein-polysaccharide
molecules termed as ground substance. All molecules found in the ECM share
common domains, and the function of the ECM relies largely on the interactions
between these molecules. Each connective tissue cell secretes a different ratio of
ECM molecules that contribute to the formation of many different architectural
arrangements; therefore, the ECM possesses specific mechanical and biochemical
properties for the tissue in which it is present. For instance, the properties of the
ECM in loose connective tissue are different from those of the ECM in cartilage or
bone. The followings are description of two major components of extracellular
materials: (i) ground substance and (ii) fibres.

i. Ground Substance

Ground substance is the part of the extracellular matrix that occupies the spaces
between the cells and fibers. It may be fluid, semi fluid, jellylike or calcified.
Functions of ground substance are followed as:
 It supports cells and helps in connecting cell to cell.
 Helps in water storage.
 Works as an exchanging medium between the blood and cells.
 It has an important role in which ways tissues develop, migrate,
proliferate, and change shape, and in what ways they perform their
metabolic functions.

Ground substance consists predominately of three groups of molecules:

(1) Proteoglycans (e.g., aggrecan, syndecan), very large macromolecules

composed of a core protein;
(2) glycosaminoglycan molecules (GAGs) (e.g., dermatan sulfate, keratan
sulfate, hyaluronan), which are covalently bound to the proteoglycans; and

(3) multiadhesive glycoproteins (such as fibronectin and laminin). The size

and structure of the three groups of molecules vary enormously (Fig.3).

Physical properties of Glycosaminoglycan (GAGs):

The GAGs are the most abundant heteropolysaccharide components of ground

substance. These molecules represent long-chain unbranched polysaccharides
composed of repeating disaccharide units. The disaccharide units contain either of
two modified sugars—N-acetylgalactosamine or N-acetylglucosamine— and a uronic

Institute of Lifelong Learning, University of Delhi 9

acid such as glucuronate or iduronate. GAGs (except hyaluronan) are synthesized by
connective tissue cells as a covalent, posttranslational modification of proteins called
proteoglycans. For example, heparin is formed by enzymatic cleavage of heparan
sulfate; dermatan sulfate is similarly modified from chondroitin sulfate.

GAGs are highly negatively charged because of the sulfate and carboxyl groups
located on many of the sugars. The high density of the negative charge (polyanions)
attracts water, forming a hydrated gel. The gel-like composition of ground substance
permits rapid diffusion of water-soluble molecules. At the same time, the rigidity of
the GAGs provides a structural framework for the cells. GAGs are located primarily
within the ground substance as well as on the surface of cells within the extracellular
matrix. On the basis of differences in specific sugar residues, the nature of their
linkages, and the degree of their sulfation, a family of seven distinct GAGs is
recognized.

Different molecules of Glycosaminoglycan (GAGs):

 Hyaluronic acid is a gelatinous, jelly-like substance which attaches cells

together, responsible for lubrication of various joints of the body, and assists
in maintaining the shape of the eye balls. White blood cells, sperm cells, and
some bacteria produce hyaluronidase, an enzyme that breaks apart
hyaluronic acid, thus causing the ground substance of connective tissue to
become more liquid. The ability to produce hyaluronidase helps white blood
cells move easily through connective tissues to reach sites of infection and
aids penetration of an oocyte by a sperm cell during fertilization. It also
accounts for the rapid spread of bacteria through connective tissues.

 Chondroitin sulfate gives support and stickiness in cartilage, bone, skin,

and blood vessels.

 The skin, tendons, blood vessels, and heart valves have dermatan sulfate;
bone, cartilage, and the cornea of the eyeball contain keratan sulfate.

Multiadhesive glycoproteins:

Multiadhesive glycoproteins accounts for linking parts of the ground substance

together and to the surface of cells. The main multiadhesive glycoproteins of
connective tissues are the fibronectin, which attaches to collagen fibers and ground
substance, joining them to one another. Fibronectin also attaches cells to the ground
substance.

Institute of Lifelong Learning, University of Delhi 10

Fig.3. Structural arrangement
of ground substance
which consists of

proteoglycans (core protein), glycosaminoglycan (dermatan sulfate,

chondroitin sulfate, hyaluronic acid), multiadhesive glycoproteins (link
protein).
Source: http://classconnection.s3.amazonaws.com/831/flashcards

Value addition: Did you Know

Nutritional Supplements for Joint Diseases
In the recent years, chondroitin sulfate and glucosamine have been used as
nutritional supplements either alone or in combination to encourage and retain the
structure and function of joint cartilage, to give pain relief from osteoarthritis, and to
decrease joint swelling. More research is needed to determine how they act and why
they help some people and not others.

Source: Principles of Anatomy & Physiology- Tortora, G.J. & Derrickson, B.

ii. Connective Tissue Fibers

Three basic kinds of fibres are embedded in the extracellular matrix between the
cells are: (a) collagen fibres; (b) elastic fibres, and (c) reticular fibres (Fig. 2). These
are formed from protein subunits secreted by fibroblasts. They function to
strengthen and support connective tissues.

Institute of Lifelong Learning, University of Delhi 11

 a. Collagen fibres

These fibres are very strong and resist pulling forces, but they are not stiff, which
allows tissue flexibility. The properties of different types of collagen fibres vary from
tissue to tissue. For example, the collagen fibres found in cartilage and bone form
different form different association with surrounding molecules. As a result of these
associations, the collagen fibres in cartilage are surrounded by more water molecules
than those in bone, which gives cartilage a more cushioning effect. It is found in
most types of connective tissues like bone cartilage, tendons and ligaments.
Chemically, collagen fibres consist of the protein collagen, which is the most
abundant protein in our body (approximately 25% of the total).

b. Elastic fibres

Elastic fibers contain the protein elastin surrounded by a glycoprotein named fibrillin,
which adds strength and stability. They are branched and wavy and after stretching
will return to their original length. These fibres are lesser in width than collagen
fibres, branch and unite together to constitute a fibrous network within a connective
tissue.

c. Reticular fibres

Reticular fibers (reticulum, a network), the least common of the three, are thinner
than collagen fibers and commonly form a branching, interwoven framework in
various organs. It consists of collagen arranged in thin bundles having a covering of
glycoprotein, give support in the walls of blood vessels and form a network
surrounding the cells in some tissues, such as areolar connective tissue, adipose
tissue, nerve fibres, and smooth muscle tissue. Reticular fibres are plentiful in
reticular connective tissue.

Value addition: Did you Know

Marfan Syndrome
It is a hereditary disorder arises from the defective fibrillin gene. It results in
abnormal production of elastic fibers. Tissues with much more elastic fibers become
deformed or weaken. Structures affected most seriously are the outer layer of the
bones, the ligament which hold the lens of the eye and the walls of the large
arteries. People with marfan syndrome tend to be tall and have abnormally long
arms, legs, fingers, and toes. A common symptom is blurred vision; arise due to the
dislocation of the eye. Around 90 percent of individuals with Marfan syndrome have
structural abnormalities in their cardiovascular systems. The most dangerous
potential result is that the weakened connective tissues in the walls of major
arteries, such as the aorta, may burst, causing a sudden, fatal loss of blood.

Source: Principles of Anatomy & Physiology- Tortora, G.J. & Derrickson, B.

Institute of Lifelong Learning, University of Delhi 12

CLASSIFICATION OF CONNECTIVE TISSUES

Connective tissue is one of the four basic tissue types. It fills the spaces between
organs and tissues, and provides them with structural and metabolic support.

Basically, Connective tissue can be classified on the basis of their maturity i.e. (A)
Embryonic and (B) Mature connective tissue. Followings are details about each
type of Connective tissue.

A. EMBRYONIC CONNECTIVE TISSUES

Embryonic connective tissues are present basically in the embryo (human which is
being developed during initial two months of pregnancy), and in the fetus (human
which is being developed during pregnancy period of third month till birth). These
are of two types, mesenchyme and mucous connective tissue.

(i) Mesenchyme: These cells are not regular in shape and embedded in semifluid
ground substance which is composed of delicate reticular fibers (Fig. 4).

Location: It is located almost exclusively under skin and along developing bones
of embryo; some in adult connective tissue, especially along blood
vessels.

Function: It forms almost all other types of connective tissue.

Fig. 4. Sectional view of embryonic mesenchyme which forms all other type of
tissue.
Source- http://www.protein-structure.net/images/Classification-Connective-
Tissue.jpg

Institute of Lifelong Learning, University of Delhi 13

(ii) Mucous (or mucoid) connective tissue: It is widely scattered fibroblasts
embedded in gelatinous and characteristically abundant ground substance
consists of fine collagen fibers (Fig. 5). It also is transitorily encountered as a
stage in the differentiation of mesenchyme into mature connective tissue.

Location: It is chiefly located in fetus.

Function: It forms the umbilical cord.

Fig. 5. Sectional view of mucous connective tissue

Source-
http://www.kln.ac.lk/science/depts/zoology/images/stories/zoo/Kumudu/mucous

B. MATURE CONNECTIVE TISSUES

These are the second major subclass of connective tissues. These are present in the
newborn. Their cells develop chiefly from mesenchyme. Based on the physical
properties of their matrix, further mature connective tissues are classified into three
major types which are given in Figure 4.

Institute of Lifelong Learning, University of Delhi 14

Fig. 4. Major types of mature connective tissue.
Source: http://www.extramarks.com/stryde/uploadfiles/image/Tushar%281%29.gif

1. CONNECTIVE TISSUE PROPER

Connective tissue proper consists of many types of cells and fibers within a matrix
containing a syrupy ground substance. Some cells of connective tissue proper are
"permanent residents'; others are not always present because they leave to defend
and repair areas of injured tissue. Connective tissue proper is categorized as either
loose connective tissues or dense connective tissues on the basis of the
relative proportions of cells, fibers, and ground substance.

Details about these types of mature connective tissues are followed as:

I. Loose Connective Tissues

Loose connective tissues are the packing material in human body. The fibres of loose
connective tissues are loosely arranged between cells. Loose connective tissue forms
a layer that separates the skin from underlying muscles, providing both padding and
a considerable amount of independent movement. Pinching the skin of the arm, for
example, does not distort the underlying muscle. The ample blood supply in this
tissue carries wandering cells to and from the tissue and provides for the metabolic
needs (oxygen and nutrients) of nearby epithelial tissue. They also anchor blood
vessels and nerves, store lipids, and provide a route for the diffusion of materials.

Institute of Lifelong Learning, University of Delhi 15

The different kinds of loose connective tissues are areolar connective tissue, adipose
tissue, and reticular connective tissue.

(i) Areolar connective tissue: It is one among the most abundant connective
tissues in the body. It consists of fibers (collagen, elastic, reticular) arranged
randomly and several kind of cells such as fibroblasts, macrophages, plasma cells,
adipocytes, mast cells, and a few white blood cells, embedded in semi-fluid ground
substance viz. hyaluronic acid, chondroitin sulfate, dermatan sulfate, and keratan
sulfate.

Location: It is known as packing material of the body as it is found in nearly

every body structure. It is present in subcutaneous layer deep to skin,
papillary region of dermis, lamina propria of mucous membranes (Fig.
5), near blood vessels, nerves, and body organs.

Function: It provides strength, elasticity and support to the different body parts.

Fig. 5. Areolar Connective Tissue in lamina propria of mucous membranes.

Source:
http://web.uaccb.edu/AcademicDivisions/MathScience/Science/BWheeler/Ess/figs/03
_19Figuredf-L.jpg

(ii) Adipose connective tissue:

Adipose tissue or fat is a loose connective tissue containing large numbers of fat
cells, or adipocytes. It contains cells or adipocytes arise from fibroblasts which are
modified to store triglycerides as conspicuous and centrally located fat droplets. Cell
is packed with a single, large triglyceride droplet, cytoplasm and nucleus are
pressed towards boundary of cell. There are two types of adipose tissue:

(a) White Adipose tissue: This composes 20 to 25 % of the body weight

in healthy adults. The amount of adipose tissue in an individual is determined
the balance between energy intake and expenditure. It is found supporting

Institute of Lifelong Learning, University of Delhi 16

 the kidneys and the eyes, between muscle fibres and under the skin, where it
acts as a thermal insulator and energy store.

(b) Brown Adipose tissue: This (BAT) is characteristically found in the

fetus and newborn. It includes most abundant capillary network and several
pigmented mitochondria that are responsible for aerobic cellular respiration.
When brown tissue is metabolised, it produce less energy and considerably
more heat than other fat, contributing to the maintenance of body
temperature. In some adults it is present in small amounts.

Location: It is present in subcutaneous layer inner to skin, surrounding heart

(Fig. 6) and kidneys, yellow bone marrow, stuffing around joints. It
fills bony socket behind the eyeball. It dominates extensive areas of
loose connective tissue in the pericardial and peritoneal (abdominal)
cavities.

Function: It provides insulation that slows heat loss through skin.

It provides another source of padding and shock absorption for the
body.
It stores energy in the form of fat; maintains and protects organs.
In newborns, BAT produces heat to maintain proper body temperature.

Fig. 6. Adipose connective tissue around heart.

Source: http://classroom.sdmesa.edu/eschmid/T04.02a-b.150.jpg

Value addition: Did you Know

Weight Control and Liposuction
Adipocytes are metabolically active cells; their lipids are continually being broken
down and replaced. When nutrients are scarce, adipocytes deflate like collapsing
balloons. This deflation occurs during a weight-loss program. In adults, adipocytes

Institute of Lifelong Learning, University of Delhi 17

cannot divide. Though, an excess of circulating lipids can stimulate the division of
connective tissue stem cells, which then differentiate into additional fat cells. As a
result, areas of loose connective tissue can become adipose tissue after chronic
overeating. In the procedure known as liposuction, unwanted adipose tissue is
surgically removed. Because adipose tissue can regenerate through differentiation of
stem cells, liposuction provides only a temporary and potentially risky solution to the
problem of excess weight. Post surgical complications that may develop include fat
that may enter blood vessels broken during the procedure obstruct blood flow,
infection, loss of feeling in the area, fluid depletion, injury to internal structures, and
severe pain.

Source: Principles of Anatomy & Physiology- Tortora, G.J. & Derrickson, B.

(iii) Reticular connective tissue:

It consists of fine interlocking network of reticular fibres (thin form of collagen fiber)
and reticular cells (Fig. 7).

Location: It is present in liver, spleen, lymph nodes; red bone marrow; reticular
lamina of basement membrane; around blood vessels and muscles.

Function: It forms stroma (supporting framework) of organs; binds smooth

muscle tissue cells; filters and eliminates damaged blood cells on
spleen and microorganisms in lymph nodes.

Fig. 7. Reticular connective tissue in spleen.

Source:
http://web.uaccb.edu/AcademicDivisions/MathScience/Science/BWheeler/Ess/figs

II. Dense Connective Tissues

Institute of Lifelong Learning, University of Delhi 18

It contain more fibres, which are thicker and more densely packed, but have
noticeably less number cells than loose connective tissues. Dense connective tissues
consist mostly of collagen fibers; they may also be called fibrous, or collagenous
tissues. Dense connective tissues are tough, strong, and durable. They resist tension
and distortion and interconnect bones and muscles. Dense connective tissue also
forms a thick fibrous layer, called a capsule that surrounds internal organs (e.g.
liver, kidneys, and spleen) and encloses joint cavities.

There are three kinds: dense regular connective tissue, dense irregular connective
tissue, and elastic connective tissue.

(i) Dense regular connective tissue: (Fig. 8).

In dense regular connective tissue, the collagen fibers are parallel to each other,
packed tightly, and aligned with the forces applied to the tissue. Tendons are cords
of dense regular connective tissue that attach skeletal muscles to bones. Their
collagen fibers run along the length of the tendon and transfer the pull of the
contracting muscle to the bone. Ligaments resemble tendons but connect one bone
to another. Ligaments often contain elastic fibers as well as collagen fibers and thus
can tolerate a modest amount of stretching. Aponeuroses are layers of tendon like
materials which join muscle to muscle and muscle to bone.

Location: It is present as Tendons (Fig. 8), Ligaments and Aponeuroses in the

body.

Function: It provides strong attachment between various structures. Tissue

structure withstands pulling (tension) along long axis of fibers.

Fig. 8. Dense regular connective tissue in tendon.

Source: http://classroom.sdmesa.edu/eschmid/T04.02c-d.150.jpg

(ii) Dense irregular connective tissue: (Fig. 9).

Institute of Lifelong Learning, University of Delhi 19

Dense irregular connective tissue contains an interwoven meshwork of collagen
fibers with a few fibroblasts. This structural pattern provides support to areas
subjected to stresses from many directions and gives skin its strength.

Location: It occurs in sheets, such as fasciae (tissue beneath skin and around
muscles and other organs), reticular (deeper) region of dermis of skin,
fibrous pericardium of heart, periosteum of bone, perichondrium of
cartilage, joint capsules, membrane capsules around various organs
(kidneys, liver, testes, lymph nodes, valves of heart.

Function: It provides pulling (tensile) strength in many directions.

Fig. 9. Dense irregular connective tissue from dermis.

Source:
http://www.highlands.edu/academics/divisions/scipe/biology/faculty/harnden/2121

(iii) Elastic connective tissue: (Fig. 10).

It is predominantly elastic fibers with fibroblasts between fibers and yellowish in

colour.

Location: It is found in lung tissue, walls of elastic arteries, trachea, bronchial

tubes, vocal cords, suspensory ligaments of penis and some ligaments
between vertebrae.

Function: It allows stretching of various organs.

It is sturdy and move back to original shape after being stretched.
Elasticity is important for normal functioning of lung tissue and elastic
arteries.

Institute of Lifelong Learning, University of Delhi 20

Fig. 10. Elastic connective tissue from aorta.
Source: http://www.kln.ac.lk/science/depts/zoology/images/stories/zoo/Kumudu

2. SUPPORTING CONNECTIVE TISSUE

Supporting connective tissues includes cartilage and bone. These tissues constitute a
strong framework which supports the rest parts of the body. These connective
tissues are composed of the extra-cellular matrix rich in fibers. In some cases, it
contains deposits of insoluble calcium salts. Different characteristic features of
cartilage and bone are compared in tabulated form also (see Table 1).

I. Cartilage

Cartilage composed of a dense network of collagen and elastic fibers which are
embedded in the chondroitin sulfate, a jelly like ground substance. Cartilage can
tolerate noticeably more stress than loose and dense connective tissues. The
strength of cartilage is owing to its collagen fibers, and its resilience (ability to move
back its original shape after being stretched) is owing to chondroitin sulfate.

Cells of mature cartilage, termed chondrocytes, prevail singly or in groups within

spaces termed as lacunae in the extracellular matrix. A covering of dense irregular
connective tissue termed as perichondrium surrounds the surface of most cartilage.
Perichondrium has blood vessel and nervous system and is the source of cartilage
cells. Because cartilage contains no blood supply, it repairs slowly subsequent to an
injury.

Value addition: Did you Know

Antiangiogenesis factor
Cartilage does not have a blood supply because it secretes an antiangiogenesis
factor. It is a substance that prevents blood vessel growth. Because of this property,

Institute of Lifelong Learning, University of Delhi 21

antiangiogenesis factor is being studied as a possible cancer treatment. If the cancer
cells can be stopped from promoting new blood vessel growth, their rapid rate of cell
division and expansion can be showed or even halted.
Source: Principles of Anatomy & Physiology- Tortora, G.J. & Derrickson, B.

Three main kinds of cartilage are hyaline cartilage, elastic cartilage, and fibrous
cartilage.

(i) Hyaline cartilage:

It consists of a resilient gel as ground substance and appears in the body as a bluish-
white, shiny substance. It constitutes very thin collagen fibers are not easily visible
in microscope. Prominent chondrocytes are found in lacunae surrounded by
perichondrium (Fig. 11). Exceptions are articular cartilage in joints and cartilage of
epiphyseal plates, where bones lengthen during growth.

Location: Most abundant cartilage in body. It is prevalent at ends of long bones,

anterior end of ribs, nose, parts of larynx, trachea, bronchi, bronchial
tubes, embryonic and fetal skeleton.

Function: It forms smooth surfaces for movement at joints

It provides flexibility and support.
It is weakest type of cartilage.

Fig. 11. Hyaline cartilage from fetus.

Source: http://classroom.sdmesa.edu/eschmid/T04.02g-h.150

(ii) Fibrous cartilage: (Fig. 12).

Cells of fibrous cartilage are chondrocytes that are scattered between visibly thick
bundles of collagen fibers within the extracellular matrix. It lacks perichondrium.

Institute of Lifelong Learning, University of Delhi 22

Location: It is prevalent in pubic symphysis, intervertebral discs, menisci
(cartilage pads) of knee, portions of tendons that insert into cartilage.

Function: It support and joins structures together.

Strength and rigidity compose it as the strongest type of cartilage.

Fig. 12. Fibrous cartilage from knee.

Source: http://classroom.sdmesa.edu/eschmid/T04.02g-h.150

(iii) Elastic cartilage: (Fig. 13).

Chondrocytes are embedded in fine network of elastic fibers within extracellular

matrix. It has perichondrium. It is also termed as yellow cartilage.

Location: It forms lid on top of larynx, parts of external ear (auricle), auditory
tubes.

Function: It gives strength and elasticity and maintains shape of certain

structures.

Institute of Lifelong Learning, University of Delhi 23

Fig. 13. Elastic cartilage from auricle of ear.
Source:
http://www.vetmed.vt.edu/education/curriculum/vm8054/Labs/Lab7/IMAGES

Value addition: Did you Know

Cartilages and Joint Injuries
Numerous complex joints in our body (e. g. knee) have both hyaline cartilage and
fibrous cartilage. The hyaline cartilage wrap bony surfaces and fibrous cartilage
cushion in the joint keep away from bone-to-bone contact when movements are
underway. Damage in these joints can produce tears in the fibrous cartilage cushion
that does not or poorly repair as the cartilages are avascular. This loss of cushioning
places more strain on the cartilages within joints and leads to further joint damage.
Eventually, joint mobility is severely reduced. Surgery usually results in only a
temporary or incomplete repair.

Source: Principles of Anatomy & Physiology- Tortora, G.J. & Derrickson, B.

Repair and Growth of Cartilage

Metabolically, cartilage is a relatively inactive tissue that grows slowly. When injured
or inflamed, cartilage repair proceeds slowly, in large part because cartilage is
avascular.The growth of cartilage follows two basic patterns: interstitial growth and
appositional growth.

Interstitial growth: In this pattern of growth, there is growth from inside the
tissue. When cartilage grows by interstitial growth involves two events:
(i) Cartilage increases rapidly in size due to the cell division of prevailing
chondrocytes and
(ii) Continuous deposition of increasing amount of extracellular matrix by
the chondrocytes.

Appositional growth: In this pattern of growth, there is growth at the external

layer of the tissue. When cartilage grows by appositional growth, cells in the inner
cellular layer of the perichondrium differentiate into chondroblasts.

II. Bone Tissue

Institute of Lifelong Learning, University of Delhi 24

Cartilage, joint, and bones make up the skeletal system. The skeletal system
supports soft tissues, protects delicate structures, and works with skeletal muscles to
generate movement. Bones store calcium and phosphorus, contain red bone marrow,
which generates blood cells and contain yellow bone marrow, a storage site for
triglycerides. Bones are organs comprised of many different connective tissues.
These include bone or osseous tissue, periosteum, red and yellow bone marrow and
endosteum. Bone tissue is classified as either compact or spongy, depending on how
its extracellular matrix and cells are organized (Fig. 14).

(i) Compact bone

Compact bone comprised of tightly packed osteons or haversian systems. The

osteon contains a central canal termed as osteonic (haversian) canal that is
enclosed by concentric rings (lamellae) of matrix. Mature bone cells (osteocytes)
are placed in spaces between the rings of matrix termed as lacunae. Small channels
termed as canaliculi radiate from the lacunae to the central osteonic (haversian)
canal to give passageways by the hard matrix. The osteonic canals constitute blood
vessels and nerves which are parallel to the long axis of the bone. These blood
vessels interlinked through the way of perforating canals with vessels on the surface
of the bone. In compact bone, the haversian systems are packed closely together to
form a solid mass.

(ii) Spongy (cancellous) bone

Spongy bone is lighter and less dense than compact bone. It contains plates
(trabeculae) and bars of bone neighbouring to small and irregular cavities which
have red bone marrow. The canaliculi attach to the neighbouring cavities, instead of
a central haversian canal, to receive their blood supply. The trabeculae are organized
to give maximum strength. The trabeculae of spongy bone follow the lines of stress
and can realign if the direction of stress changes.

Location: Compact and spongy bones tissue constitute the various parts of
bones of the body.

Function: It provides support, protection and storage.

It contains blood forming tissue.
It provides levers that act with muscle tissue to enable movement.

Institute of Lifelong Learning, University of Delhi 25

 Fig
.
14. Sectional view of several osteons ( haversian systems ) of bone.
Source: http://cdn.intechopen.com/pdfs-wm/44658.pdf

Table 1: Comparison between features of Bone and Cartilage

FEATURES BONE CARTILAGE

STRUCTURAL
FEATURES
Cells Osteocytes housed in lacunae Chondrocytes located in
within matrix lacunae within matrix

Ground substance A small volume of liquid Chondroitin sulfate (Protein-

surrounding insoluble crystals of polysaccharide gel) and water
calcium salts (calcium
phosphate and calcium
carbonate)
Fibers Collagen fibers predominate Collagen, elastic, reticular fibers
(proportions vary)

Blood supply Contains blood vessels Has no blood vessels

Outer covering of Periosteum (except at joints), a Perichondrium (except at

tissue fibrous membrane containing joints); composed of outer layer
nerves, lymphatic vessels and of dense connective tissue and
capillaries inner layer of cells that
differentiate into chondrocytes;
fibrocartilage has no
perichondrium

Institute of Lifelong Learning, University of Delhi 26

Strength Strong: resists distortion until Limited: bends easily but
breaking point is reached difficult to break

METABOLIC
FEATURES
Oxygen demands High Low

Nutrient delivery By diffusion through cytoplasm By diffusion through selectively

and fluid in canaliculi permeable intercellular matrix

Growth Appositional growth only Interstitial and appositional

growth
Repair capabilities Extensive Limited

3. FLUID CONNECTIVE TISSUE

Fluid connective tissues include blood and lymph which are characteristic collections
of cells in a fluid extra-cellular matrix. Under normal conditions, the proteins
dissolved in this watery matrix do not form large insoluble fibers. In blood, the
watery matrix is called plasma.

I. Blood Tissue

Blood tissue is a connective tissue having liquid extra-cellular matrix and formed
elements. The extra-cellular matrix is termed as blood plasma. The blood plasma is
pale yellow fluid which consists generally of water with a wide variety of dissolved
materials (e.g. nutrients, wastes, enzymes, plasma proteins, hormones, respiratory
gases, and ions). Suspended in the blood plasma are formed elements—red blood
cells, white blood cells and platelets. Red blood cells carry oxygen to various body
cells and remove some carbon dioxide from them. White blood cells are responsible
for phagocytosis, immunity, and allergic responses. Platelets participate in clotting of
blood (Fig. 15).

Location: It is located within blood vessels, within chambers of heart

Function: (i) Red blood cells carry oxygen and few carbon dioxide.
(ii) White blood cells carry on phagocytosis and mediate allergic
reactions and immune system responses.

(iii) Platelets are essential for blood clotting.

Institute of Lifelong Learning, University of Delhi 27

Fig. 15. - Blood smear of human body.
Source: http://classroom.sdmesa.edu/eschmid

II. Lymph

Lymph is the extra-cellular fluid that moves in lymphatic vessels. This is the
connective tissue which comprised of various forms of cells in a transparent liquid
extracellular matrix that resembles to blood plasma although contains much less
protein. The composition of lymph varies from one part of body to another. For
example, lymph leaving lymph nodes includes many lymphocytes, a type of white
blood cell, in contrast to lymph from the small intestine, which has a high content of
newly absorbed dietary lipids.
The three main forms of lymphocyte are thymus cells (T cells), B cells (bursa-derived
cells) and natural killer (NK) cells. Lymphocytes have characteristically large nucleus
(Fig. 16).

Location: It is located within lymphatic vessels.

Function: It facilitates the immune system to the body.

Institute of Lifelong Learning, University of Delhi 28

Fig. 16. A lymphocyte (stained) surrounded by red blood cells.

Source: http://en.wikipedia.org/wiki/Lymphocyte

Summary

 Connective tissue is one of the four general classes of animal tissues (those are
epithelial, muscle, and nervous tissues). It fills the spaces between organs and
tissues, and provides structural and metabolic support for other tissues and
organs. Connective tissues consist of two basic elements: extracellular matrix
and cells.

 The types of cells in connective tissues are fibroblasts, macrophages, plasma

cells, mast cells, adipocytes and leukocytes which are components of different
types of connective tissues.

 Every kind of connective tissues has distinctive properties, depending on the

specific extracellular materials comprises of ground substance and fibres,
between the cells. Ground substance consists predominately of three groups of
molecules: Proteoglycans (core protein), glycosaminoglycan molecules (GAGs)
which are covalently bound to the proteoglycans; and multiadhesive
glycoproteins. Basic types of fibres are embedded in the extracellular matrix
between the cells are collagen fibres, elastic fibres and reticular fibres.

 These are of two types of embryonic connective tissues, mesenchyme and

mucous connective tissue. Mesenchyme forms almost all other types of
connective tissue. Mucous connective tissue support the umbilical cord of fetus.

 Connective tissue proper is categorized as either loose connective tissues or

dense connective tissues on the basis of the relative proportions of cells, fibers,
and ground substance.

Institute of Lifelong Learning, University of Delhi 29

 The fibres of loose connective tissues are loosely arranged between cells. It
forms a layer that separates the skin from underlying muscles, providing both
padding and a considerable amount of independent movement. The kinds of
loose connective tissues are areolar connective tissue, adipose tissue, and
reticular connective tissue.

 Dense connective tissues contain more fibres, which are thicker and more
densely packed. They resist tension and distortion and interconnect bones and
muscles. There are three forms of dense regular connective tissue, dense
irregular connective tissue, and elastic connective tissue.

 Supporting connective tissues include cartilage and bone they constitute a

strong framework that supports the rest parts of the body.

 Cartilage comprised of a dense network of collagen fibres and elastic fibers

embedded in the chondroitin sulfate, a gelatinous like ground substance. The
three main forms of cartilage are hyaline cartilage, elastic cartilage, and fibrous
cartilage.

 Bone tissue is categorized as compact and spongy bone, depending on how its
extracellular matrix and cells are arranged. The fundamental component of
compact bone is an osteon or haversian system. Spongy bone lacks osteons.

 Blood and lymph are fluid connective tissues which have characteristic
assembly of cells in a fluid extracellular matrix. Blood tissues are important in
transport of oxygen and immune system responses. Lymph tissues facilitate the
immune system to the body.

Institute of Lifelong Learning, University of Delhi 30

Exercises

1. Explain the general features of connective tissues.

2. Describe the various functions of connective tissues.

3. What are the components of connective tissues? Explain their role in making
connective tissue with example.

4. Give the classification of connective tissues with their location and functions.

5. Describe difference between bone and cartilage.

6. Cartilage heals very slowly. Explain the reason.

7. Define the following

 Mesenchyme
 Perichondrium
 Osteons

8. Explain two types of Adipose tissue with their location and function

9. Explain the difference between interstitial and appositional growth of cartilage

10. Describe the structure and function of the various types of loose and dense
connective tissues.

11. Write short notes on

 Connective Tissue Fibers

 Lymph
 Glycosaminoglycan (GAGs)

12. Describe the haversian systems of bone with diagram. Explain difference
between compact and spongy bone.

13. Describe types of the fluid connective tissues with the diagram.

Institute of Lifelong Learning, University of Delhi 31

Glossary
Tissue: It is an assembly of similar cells arises from the same origins which carries
out specific function.

Connective tissue: It is a type of animal tissue which supports, connects or

separates various forms of tissues and organs of the body.

Extracellular matrix: It is made up of fibres in a protein and polysaccharide matrix,

secreted and organised by cells of connective tissue.

Adipocytes: These are cells of connective tissue which reserves triglycerides and
are also called fat cells or adipose cells.

Leukocytes: These are white blood cells which move along with the connective
tissues adjacent to blood vessels.

Histamine: It is a chemical that widens small blood vessels as a component of the

inflammatory response.

Ground substance: It is the constituent of a connective tissue lie between the cells
and fibres which is perhaps fluid, semi fluid, viscous or calcified.

Glycosaminoglycan (GAGs): These are heteropolysaccharide components of

ground substance with high density of the negative charge (polyanions) to attracts
water, forming a hydrated gel in order to permit rapid diffusion of water-soluble
molecules.

Mesenchyme: It is a type of embryonic connective tissues which are present

basically in embryo forms almost all other types of connective tissue.

Cartilage: It is supporting connective tissue which contains a dense network of

collagen fibres and elastic fibers lie within the chondroitin sulfate.

Osteon: It forms compact bone and contains central canal termed the osteonic or
haversian canal, which is enclosed by concentric rings (lamellae) of matrix.

Blood tissue: It is a fluid connective tissue composed of liquid extracellular matrix

and formed elements i.e. red blood cells, white blood cells and platelets.

References

Institute of Lifelong Learning, University of Delhi 32

 1. Histology: A Text and Atlas by Michael H. Ross and Wojciech Pawlina, 6th
edition.

2. Anatomy And Physiology: In health and illness. Ross and Wilson (Tenth
Edition)

3. Tortora, G.J. & Grabowski, S. Principles of Anatomy & Physiology. 13th

Edition.

4. Hill, R. W., Wyse, G. A. and Anderson, M. (2006). Animal Physiology. p.355.

5. Randall, D., Burggren W. and French, Kathleen (2001). Eckert Animal

Physiology.

6. Widmaier, E.P., Raff, H. and Strang, K.T. (2008). Vander’s Human Physiology,
XI Edition, McGraw Hill.

7. Textbook of Physiology by Prof. A.K. Jain.

8. Guyton, A.C. and Hall, J.E. (2011). Textbook of Medical Physiology, XII
Edition, Harcourt Asia Pvt. Ltd. W.B. Saunders Company.

Institute of Lifelong Learning, University of Delhi 33