HISTOLOGY (Study of tissues)

A tissue is a group of similar cells organized into a structural and functional unit. It is a group of
cells of similar structure organized for carrying out a particular function(s).

Characteristics of tissues

 Cells of a tissue are physically linked.

 The cells of a tissue may be interspersed with intercellular substances.
 A tissue may comprise one or more types of cells.
 A tissue is specialized to perform a particular function(s).

ANIMAL TISSUES

Early in development, the cells of the growing embryo differentiate into three fundamental embr
yonic tissues called germ layers.

 Ectoderm which forms the outer layer of the skin and the nervous system.
 Mesoderm which forms muscles, connective tissues, skeleton, kidneys and circulatory
and reproductive organs.
 Endoderm which forms the lining of the respiratory tract and urinary bladder. It also
forms the glands associated with the guts and respiratory tract.

Therefore the germ layers in turn differentiate into different cell types and tissues that are charact
eristic of the vertebrate’s body. Tissues are joined to each other by proteins. The point of connect
ion between two cells is a junction.

In adult vertebrates, there are four principle kinds of tissues which include; epithelial tissue, conn
ective tissue, muscular tissue and nervous tissue

EPITHELIAL TISSUE

This is a collection of closely packed single and multilayered compound sheets of cells covering
the external and internal surface of the body of an organism.

Characteristics of epithelial tissue

1. Cellularity- Epithelial tissue consists of tightly packed cells that are firmly attached to
each other with little intercellular material between them. Epithelial cells are held firmly
together by small amounts of carbohydrate cementing substances and by special
intercellular junctions between the cells.
2. Polarity-The bottom of epithelial cells rests on a basement membrane composed of a
network of fibres which include collagen. The portion of epithelial cells attached to the
basement membrane is called the basal surface, the opposite end facing the external

1
 environment or the lumen of the body cavity is called the apical surface/free surface.
3. Avascularity- There are no blood vessels in the epithelial tissues hence the tissue lacks
vascularity. However, nerve endings may occur in the epithelium.
4. Supported by connective tissue- Epithelial tissue receives nutrients by diffusion from
underlying connective tissue through the basement membrane. As the epithelial cells are
not supplied with blood vessels, they rely on diffusion of nutrients and oxygen from
lymph vessels which run through nearby intercellular spaces.
5. Regeneration- Epithelial cells have a high regeneration capacity due to rapid cell division.
This gives the epithelial tissue quick recovery after any injury or abrasions.
6. Germ cell origin- Epithelial cells are derived from all the three primary layers; ectoderm,
mesoderm and endoderm.

CLASSIFICATION OF EPITHELIAL TISSUE

Epithelial tissues are classified according to the number of cells/layers and the shape of the indivi
dual cells.

Epithelial tissue

Simple epithelium (one cell layer thick) compound (more than one cell layer thick)

Pseudostratified squamous cuboidal columnar ciliated stratified transitional

A. Simple squamous/pavement epithelium

It consists of a single layer of thin and flattened cells. They are so thin that the nucleus causes a b
ulge and it is centrally placed, with a disc shape. Cells are irregular with tapering edges. Simple s
quamous epithelium is found in the following areas: Renal corpuscles of the kidney, lining of the
alveoli of the lungs, lining of the blood vessels where it is referred to as the endothelium, blood
capillary walls and lining of lymphatic vessels.

Illustration

Functions

 Diffusion of materials
 Exchange of gases

2
 Adaptations

i. Thin flattened cells to reduce the distances across which materials diffuse.
ii. Provides smooth lining to allow relatively friction free passage of fluids and materials
through the hollow structures.
B. Simple cuboidal epithelium

The cells are roughly cube shaped and possess a central spherical nucleus. The upper surface of c
uboidal cells is either pentagonal or hexagonal in outline. It is the least specialized of all the epith
elial tissues.

Distribution

 Lining of salivary ducts, pancreatic ducts, convoluted tubules and collecting ducts of the
nephrons of the kidney
 Lining of the salivary glands, sweat glands and thyroid glands
 Lining of the retina

NB: The simple cuboidal epithelial tissue is non-secretory in the proximal convoluted tubule, dis
tal convoluted tubule and pancreatic ducts.

The functions include: Protection, excretion, absorption and secretion

Adaptations

i. Cells are tightly packed together with little intercellular spaces between them to offer
protection from injury and infection.
ii. Possess many Golgi bodies which perform functions of secretion of hormones and
enzymes.
iii. Some possess microvilli which increase the surface area for example reabsorption of
materials from the renal fluids in the kidney tubules.
iv. Cells have numerous mitochondria for energy production to be used in active
reabsorption of materials eg from renal filtrate back into the bloodstream.

3
 C. Simple columnar epithelium

It is a single layer of column like narrow elongated cells at right angles to the basement membran
e. Each cell possesses a nucleus situated at the basal end and it is oval in shape. The epithelium is
often interspersed with goblet cells. The free surface of each columnar cell has microvilli formin
g a brush border.

Distribution

 Lining of the stomach, small intestines, gall bladder, and kidney ducts.
 Lining of the gastric glands, intestinal glands, mammary glands, thyroid glands, salivary
glands.

The functions include: Secretion, protection, absorption and the surface area is increased by havi
ng brush border (microvilli) at the cell free surface.

Adaptations

i) Possess fingerlike projections called microvilli which increase the surface area for
absorption such as digested food in the intestines.
ii) Possess mucus secreting cells which secrete mucus. The mucus protects the gastric
walls from hydrochloric acid and digestive enzymes.
iii) Mucus from goblet cells also lubricates the passage of food in the intestines.

D. Ciliated epithelium

Cells of this tissue are usually columnar in shape but bear numerous cilia at their free surfaces. T
he cells are usually associated with mucus secreting goblet cells.

Distribution

 Lines the oviduct

 Ventricles of the brain, spinal canal and respiratory passages (trachea, bronchi and
bronchioles)

Functions

4
  Mucus protects lining and lubricates the passage of materials.
 Cilia set up currents to move materials in a certain direction.

Adaptations

i) Interspersed with goblet cells which secrete mucus to protect the lining of the gut
from enzyme and acidic action.
ii) Possess cilia which set up currents that move materials from one direction to another.
iii) Possess goblet cells which lubricate the passage.

E. Pseudostratified epithelium

This is a simple epithelium since all cells rest on a basement membrane, but some do not reach th
e free surfaces. This gives an appearance of the epithelium to be on different levels and the nucle
i at different layers. Nevertheless, the epithelium is one layer of cells thick with each cell attache
d to the basement membrane.

Most cells are columnar, thus usually named pseudostratified columnar epithelium. Where the cil
ia appear at the free surface (trachea, bronchi, bronchioles), it is called pseudo stratified columna
r ciliated epithelium.

Pseudo stratified epithelium also lines the urinary tract and ducts of large glands (non-ciliated ps
eudo stratified).

Pseudo stratified columnar

pseudo stratified columnar ciliated

5
 STRATIFIED EPITHELIA

Have more than one layer of cells with only the bottom layer resting on the basement membrane.
These cells form the germinative layer and continue to divide by mitosis and push other areas ou
twards. They are primarily found in areas where the epithelium has protective functions.

a) Stratified squamous epithelium

The cells first formed on the basement membrane are cuboidal in shape, but as they are pushed o
utwards the free surface of the tissue, they become flattened and these cells are called squamous.
It is the thickest of all epithelia and its function is protection.

The cells of the surface layer may have keratin (cornified) or lack keratin (uncornified). Keratin i
s a tough protective protein which prevents water loss, is resistant to friction and repels bacteria.

Non-keratinized stratified squamous epithelium lines wet surfaces subjected to abrasion such as l
ining of the mouth, oesophagus, part of the epiglottis and vagina. Keratinized stratified squamous
epithelium forms the epidermis of the skin.

b) Stratified cuboidal epithelium

6
This epithelium has several layers of epithelial cells but the surface layer of this epithelium is co
mposed of cuboidal cells. It is found in the largest ducts of sweat glands, mammary glands, saliv
ary glands and in parts of the male urethra. Its role is protection and provides strength.

c) Stratified columnar epithelium

This epithelium has several layers of epithelial cells but the surface layer of this epithelium is co
mposed of columnar cells. It is very rare. It lines parts of the urethra, larger ducts of some glands,
portion of the conjunctiva of the eye. Its roles are protection and absorption.

d) Transitional epithelium

It comprises 3 to 4 layers of cells all of similar size and shape except at the free surface where th
ey are more flattened. The cells do not slough off/flake off.

It is found in organs/structures which can expand like the bladder, guddles of women where they
come back to normal after delivery, ureters and part of urethra. It allows for distension of the uri
nary organ.

Because the shape of the cells at the surface is transitory (changes depending on the degree of str
etching of the organ), this epithelium is called transitional. It will look like a stratified squamous
epithelium if it is stretched or stratified cuboidal epithelium if it is unstretched.

Adaptations of the transitional epithelium to its function

i) By changing its shape, the transitional epithelium allows expansion of the organ such
as the urinary bladder. This increases the volume of the organ.
ii) Transitional epithelium is composed of many layers of cells making it impermeable to
water from blood to urine.
iii) Due to its thickness, it prevents urine from escaping to the surrounding tissues.

Glandular epithelium

It is formed by epithelial cells which are frequently interspersed with secretory cells eg goblet cel
ls or aggregates of glandular cells forming multicellular gland.

Illustration of glandular epithelium

There are two types of glands.

 Exocrine glands where the secretion is delivered to the free surface via ducts.

7
  Endocrine glands- Secretions are released and passed into the bloodstream (ductless
glands).

Based on the mode of secretion, the exocrine glands are of three types

a) Merocrine glands

Secretions produced by cells are simply passed through the cell membrane at the cell free surface
without losing any of its cytoplasm. The cell therefore remains intact. Examples include; goblet
cells, pancreatic glands, sweat glands.

b) Apocrine glands

Here, the glandular epithelia shed their secretion by the top part of the cell loaded with the secreti
on breaking away from the rest of the cell eg mammary glands. The cell loses part of the cytopla
sm while releasing its secretion.

c) Holocrine glands

In this case, the whole secretory cell disintegrates and the secretions are from the epithelium eg t
he sebaceous glands of the mammalian skin.

An epithelium containing goblet cells is called a mucus membrane. Multicellular exocrine glands
exist in various forms which include

1. Simple saccular/alveolar gland-located in the skin of frog and other amphibians

2. Simple tubular gland- found in the crypts of lieberkuhn in the walls of the mammalian
small intestines.

3. Coiled tubular gland- eg that found in the sweat glands in the skin of humans

8
4. Simple branched tubular gland- eg brunner’s glands in the walls of the mammalian small
intestines and gastric glands in the walls of the stomach.

5. Simple branched saccular gland- eg the oil secreting sebaceous glands in the mammalian
skin.

6. Compound tubular gland- eg the salivary glands

7. Compound saccular gland- eg part of the pancreas which secretes the digestive enzymes
and mammary glands.

Functions of the epithelial tissue

1. Protection

9
Epithelial tissue basically protects the underlying tissue from injuries by chemicals, pressure abra
sion and infection.

Adaptations of the epithelial tissue for protection

i. Columnar epithelium lining the stomach is interspersed with goblet cells which
secrete mucus. The mucus protects the stomach lining from acidic contents of the
stomach and from digestion by enzymes. Mucus also lubricates the passage of food
thereby protecting the lining from abrasion.
ii. The shorter cells of the pseudo stratified epithelium lining the trachea and bronchi
secretes mucus which traps dust particles and bacteria in the inhaled air and the cilia
on the longer cells beat expelling them in the outward direction.
iii. Keratinized squamous stratified epithelium on external skin surfaces (epidermis) is
highly resistant to mechanical damage due to cornification (addition of keratin)
protecting the underlying tissue from abrasion.
iv. The layers of the non-keratinized stratified squamous epithelium lining the pharynx
and oesophagus are thick, protects the underlying tissue from mechanical damage by
the food that is swallowed.
v. The many layers of the stratified cuboidal epithelium lining the salivary, pancreatic
and sweat ducts protects them against mechanical stress.
vi. The layers of the columnar cells of the stratified columnar epithelium lining the ducts
of the mammary glands, protects them from mechanical and chemical injury.
2. Secretion

A number of epithelial cells are modified to produce secretions such as mucus, hormones etc.

Adaptations of epithelium for secretion

i. Cuboidal epithelium lining the salivary glands, sweat glands and thyroid gland secrete
saliva, sweat and thyroxine respectively.
ii. Columnar epithelium lining the stomach is interspersed with goblet cells which
secrete mucus.
iii. The stratified columnar epithelium lining the ducts of mammary glands secretes a
fluid.
3. Excretion- The epithelial cells of the kidney tubules and sweat glands remove excessive
and toxic metabolic wastes from the body thus helping the body in excretion.

Ciliated cuboidal epithelium lining parts of the nephron have cilia which beat to facilitate flow gl
omerular filtrate.

4. Absorption- Cuboidal and columnar epithelia are modified for absorption.

Adaptation of epithelia to absorption

10
 a) Squamous epithelium lines the renal corpuscles of the kidney, the alveoli of the lungs and
the blood capillary walls where its extreme thinness permits rapid diffusion of materials
through it.
b) Cuboidal epithelial cells lining the proximal convoluted possess extensive microvilli
forming a brush border of microvillus which increases the surface area for absorption of
nutrients.
5. Exchange of materials and gases- Squamous epithelium is extremely thin and flattened
promoting exchange of materials and gases by diffusion such as the alveoli of the lungs.
6. Sensory- epithelia bearing sensory cells and nerve endings are specialized to receive
stimuli as in the skin and retina of the eye.
7. Movement of materials- Epithelia may be modified to aid movement of materials.

Adaptations of epithelia to move materials

i) Ciliated columnar epithelium lining the inside of the oviduct, ventricles of the brain,
spinal canal and respiratory passages bears numerous cilia at their free surface. These
are associated with mucus secreting goblet cells producing fluids in which the cilia
beat up rhythmically setting up currents which move materials from one location to
another.
ii) Pseudo stratified epithelium lining the trachea and bronchi possess longer ciliated
cells and shorter mucus secreting cells without cilia. The mucus traps bacteria, dust
and other small particles preventing them from reaching the lungs.
iii) Squamous epithelium lining hollow structures eg the heart chambers, blood vessels is
flat and smooth allowing friction free passage of materials through them.

CONNECTIVE TISSUES

Connective tissues are derived from the embryonic mesoderm.

Functions of connective tissues

a) It binds the various tissues together like the skin with muscles and muscles with bones.
b) It is a packing tissue forming sheath like bags around the body organs.
c) Areolar tissue protects the body against wounds and infections.
d) Adipose tissue stores fats, and insulates the body against heat loss.
e) Connective tissue is the major supportive tissue of the body, composed of bones and
cartilage which provides the body with a supportive framework.
f) Haemopoitic tissue produces blood.
g) Lymphatic tissue builds body immunity by producing antibodies.
h) Connective tissue separates the body organs, so that they do not interfere with each
other’s activities.
i) Protects blood vessels and organs where they enter or leave organs.

11
Connective tissues occur in different forms which are divided into two major classes

 Connective tissue proper which is further divided into loose and dense connective tissues
 Special connective tissues which include; cartilage, bone and blood. Cartilage and bone
form the skeletal tissue.

LOOSE CONNECTIVE TISSUES

They include areolar, adipose and recticular connective tissues

Areolar connective tissue

This is the most abundant type of connective tissue found all over the body beneath the skin, con
necting organs together and filling spaces between adjacent tissues.

Areolar connective tissue consists of a gelatinous glycoprotein matrix or ground substance contai
ning two types of protein fibres and four types of cells. The protein fibres include;

I) Collagen fibres

They are white fibres forming wavy bundles running parallel to each other and are not branched.
They are flexible but inelastic (non-stretchable)

II) Elastic fibres

They are thin yellow fibres which are highly branched forming a network in the matrix. They are
flexible and elastic. The main function of fibres is to give the areolar tissue its strength and toug
hness. It also allows the tissue to be flexible and elastic.

The cells within the matrix include;

a) Fibroblasts

These are flattened and spindle like shaped cells containing an oval nucleus. They are generally c
losely applied to fibres but migrate to the wounded tissue o secrete more fibres in that region that
effectively seal off the injured area. The function of fibroblasts is to secrete fibres

b) Macrophages/histocytes

They are large cells capable of amoeboid movement for which reason; they are referred to as am
oeboid cells. Their function is to engulf/ingest bacteria and other foreign particles. They are gene
rally mobile but at times, they wonder to areas of bacterial invasion. Therefore, they serve to def
end the body against diseases.

c) Mast cells

12
They are amoeboid cells which are oval shaped and contain granular cytoplasm. They are found i
n abundance close to blood vessels. They have the following functions

 Secretion of the matrix.

 Secretion of an anticoagulant, heparin.
 They secrete histamine, a substance attributed to the effects of allergy.
d) Fat cells

They are mainly filled by lipid droplets. They cytoplasm and nucleus of a fat cell are confined to
the margins of the periphery.

Diagram of an areolar tissue (FA page 36)

Areolar connective tissue is found in the skin and in most internal organs of vertebrates where it
allows the organs to expand. It also forms the protective covering of muscles, blood vessels and
verves.

Functions of the areolar connective tissue

1. Binds tissues and organs together.

2. Serves as a packing tissue filling spaces between adjacent tissues.
3. Support various tissues.
4. Provide tissues which resist strain and displacement.
5. Provides protection against wounds and infections.

Adipose tissue

13
It is a type of connective tissue with reduced matrix material and contains enlarged fat cells that
are numerous in number. Adipose tissue functions to store energy, insulate the body and provides
shock absorption to delicate mammalian organs eg the kidney. It also occurs beneath the skin, th
e buttocks. Adipose tissue occurs in two forms ie the white and brown adipose tissue.

Recticular connective tissue

It contains an abundance of recticular fibres. It provides a supporting framework for organs such
as those of the lymph nodes, spleen and the liver.

DENSE (FIBROUS) CONNECTIVE TISSUE (DCT)

DCT contains tightly packed collagen fibres making it stronger than the lose connective tissue. It
consists of two types, regular and irregular

The collagen fibres in the dense regular connective tissue are oriented in one direction to provide
strength in that direction. It is found in tendons and ligaments. Tendons connect muscles to bone
s, ligaments connect bone to bone.

Irregular dense connective tissue contains collagen fibres oriented in many different directions. It
is found in deep layers of the skin and the tough capsules that surround many of the organs eg th
e kidney, adrenal glands, nerves, bones and covering of the muscles as epimysium and periosteu
m. It provides support and strength.

SPECIAL CONNECTIVE TISSUES

SKELETAL TISSUES

CARTILAGE/GRISTLE

This is a connective tissue consisting of cells embedded in a matrix called chondrin. The matrix
is deposited by cells called chondroblasts and possess many fine fibres mostly collagen. Eventu
ally, the chondroblasts become enclosed in spaces called lacunae. In this state, they are termed a
s chondrocytes.

14
The margin of a piece of cartilage is enclosed by a dense layer of cells and are fibrils called peric
hondrium in which new chondroblasts are produced and constantly added to the internal matrix
of the cartilage.

Cartilage is highly adapted to resist any strains that are placed on it. The matrix of cartilage is co
mpressible and elastic. The collagen fibres resist any tensions which may be imposed on the tissu
e.

The cartilage in adults is restricted to the articular (joints) surfaces of bones that form freely mov
eable joints and other specific locations eg the nose, pinna, intervertebral discs, larynx, etc.

Types of cartilage

There are three types of cartilage each with the organic components of the matrix quite distinct.
They include; hyaline cartilage, yellow elastic cartilage and white fibrous cartilage.

Hyaline cartilage

It is the simplest form of cartilage which is elastic and compressible. It mainly comprises a semit
ransparent matrix and chondrocytes. It frequently contains fine collagen fibrils. It has no process
es extending from the lacunae into the matrix and neither are there blood vessels. Therefore, exc
hange of materials between the chondoblasts and the matrix is by diffusion.

It is located in the ends of the bones, in the nose and the wall of the trachea and bronchi. It also f
orms the embryonic skeleton in many vertebrates.

15
 Yellow elastic cartilage

It has a semi opaque matrix containing a network of yellow elastic fibres. The fibres confer great
er elasticity than found in the hyaline cartilage. Due to high elasticity and flexibility, the tissue q
uickly returns to its shape after distortion.

It is located in the external ear, in the epiglottis and cartilages of the pharynx

White fibrous cartilage

In addition to chondrocytes in the matrix, there are large bundles of densely packed collagen fibr
es. This gives the tissue a greater tensile strength than hyaline cartilage as well as a small degree
of flexibility.

It is located in the discs between adjacent vertebrae which provide a cushioning effect and the lig
amentous capsules surrounding the joints.

Adaptations of the collagen tissue to its function

 It is flexible but very strong and non stretchable allowing it to withstand the stresses of
movement of connective tissues.
 It has a fibrous structure which allows it to bind structures together in connective tissues.

BONE

This is very abundant providing support, protection and some metabolic functions. The bone has
an organic matrix containing collagen fibres, and is impregnated with small needle shaped crysta

16
ls of calcium phosphate in form of hydroxyapatite which is brittle but rigid giving bone great stre
ngth. Calcium carbonate is also contained within the matrix.

A bone is a dynamic living tissue that is constantly reconstructed through the life of an individual
by bone cells called osteoblasts. Osteoblasts secrete the matrix in which calcium phosphate is la
ter deposited. After calcium phosphate has been deposited, the osteoblasts become less active an
d are now called osteocytes and are energized in spaces called lacunae.

Another type of bone cells called osteoclasts exist in thematrix which play a role in dissolving th
e bone matrix to enable further reconstruction of a bone during growth. A bone is constructed in t
hin concentric layers called lamellae which are drawn around narrow channels called haversian
canalsthat run parallel to the bone length.

Haversian canals contain nerve fibres and blood vessels which keep the osteocytes alive. The con
centric lamellae and the encircled canal are termed as the haversian system/osteon.

The lacunae have very many fine channels called canaliculi containing cytoplasm which link up
with the central haversian canal, with other or press from one lamella to another. An artery and a
vein run through the haversian canal and capillaries branch from here through the canaliculi. A h
aversian canal also contains lymph vessels and nerve fibres.Covering the bone is a layer of dense
connective tissue called periosteum. The inner region of the periosteum has blood vessels and c
ontains cells that can develop into osteoblasts and osteoclasts.

Part of the transverse section of a haversian system

Types of bone

There are two types of bone ie compact/dense bone and spongy bone

Compact bone/dense bone

They are used in the growth of long bones and form the long shape of the bone between two swo
llen ends. The matrix of compact bones is composed of collagen and calcium phosphate, with qu
antities of magnesium, sodium, carbonate, nitrate ions. The combination of organic with inorgani
c materials produces a structure of great strength.

17
The lamellae are laid down in a manner that is suited to the force acting upon the bone and the lo
ad that has to be carried.

A transverse section of a compact bone

Longitudinal section of a compact bone

Spongy/cancellous bone

Spongy bone occurs within longer bones and is always surrounded by compact bone. Spongy bo
ne consists of thin bars or sheets of bone called trabeculae, interspersed with large spaces occupi
ed by the bone marrow.

The trabeculae contain osteocytes which are more or less irregularly dispersed in the matrix. The
matrix contains rather a smaller proportion of inorganic material than does the matrix of the com
pact bone. The trabeculae develop along the lines of stress within the bone.

The spaces within the spongy bone at the head (epiphysis) of the long bones contain red bone ma
rrow tissue. This very soft tissue is less dense than the bone, and is the site of red blood cell form
ation. Yellow marrow tissue, consisting principally of fat fills the spaces within the spongy bone
of the shaft.

Spongy bone in section

18
 Long bone in section

The development of bone (ossification)

Ossification is the process of formation and development of bone. A bone originates in two ways
; intramembranous ossification and endochodral ossification.

Intramembranous ossification

The thin bony plates of the skull and parts of some other bones eg clavicles are formed directly b
y clusters of ossification which appears inside fibrous membranes. The strands of bone formed b
y different clusters are called trabecullae and become linked to form a loose network described a
s spongy bone. Thus spongy bone is formed by intramembranous ossification.

As development continues, remodeling of the skull plates converts some of the spongy bone to c
ompact bone and allows the plates to reach their adult shape and size.

Endochondral ossification

19
Endochondral ossification is the process of replacement of cartilage by bone. The skeleton of the
vertebral embryo consists of mainly hyaline cartilage. Each cartilage element is surrounded by a
layer of dense connective tissue, the perichondrium

Ossification of bones (such the long bones of the arms and legs) begins when blood vessels penet
rate the perichondrium of the cartilage midway along the shaft/diaphysis of the cartilage model.

This stimulates some of the cells of the cartilage perichondrium to become osteoblasts which pro
duce a collar of compact bone in the shaft region. The layer of dense connective tissue now surro
unding the developing embryo is called the periosteum.

A primary ossification centre appears inside the shaft and is progressively invaded by a proliferat
ing number of blood vessels and osteoblasts. The matrix of the cartilage tends to become calcifie
d by deposition of calcium and phosphate, but is eroded by the osteoclasts leaving spaces which
eventually fuse to form the marrow cavity.

Working in small groups, osteoclasts tunnel through the bone leaving curvities which are invade
d by blood capillaries and new bone forming osteoblasts. Within the tunnel, osteoblasts lay down
a new bone matrix in concentric rings forming an arrangement called haversian system.

Cartilage continues to grow at either end producing an increase in length. Most of the cartilage is
later replaced by a spongy bone.

In mammals, secondary ossification centres develop in the swollen ends/epiphyses of the cartilag
e models of long bones. The epiphyses ossify more or less completely except for a thin layer of c
artilage called an epiphyseal plate, separating each epiphysis from the main shaft.

Increase in diameter of the bone shaft is achieved by continual remodeling and deposition of new
bone by osteoblasts of the periosteum. As maturity approaches, the thickness of the epiphyseal p

20
lates is reduced and finally the epiphyses and the bone shaft fuse completely leaving a faint epip
hyseal line. Ossification of all bones in human skeleton is normally completed by the age of 25.

Adaptations of bone to its function

a) A tough fibrous layer of dense connective tissue called periosteum provides a tough and
hard covering that surrounds the bone and protects the inner cells.
b) Bundles of collagen fibres from the periosteum penetrate the bone giving more
mechanical strength, providing an intimate connection between the underlying bone and
the periosteum and acting as a firm base for insertion of tendons, which contribute to
movement and locomotion.
c) Osteoblasts are arranged in concentric rings around a series of haversian canals in
compact bone thus lay down the matrix in a similar rigid and dense regular pattern to
provide uniform mechanical strength.
d) Bone lamellae contain numerous lacunae containing living bone cells called osteoblasts
which secrete the matrix of the bone.
e) Mature less active osteoblasts called osteocytes can be reactivated quickly regaining the
structure of active osteoblasts and depositing bone matrix, when structural changes in
bone are required.
f) Bone cells are embedded in a firm bone matrix which is rendered hard by deposition of
calcium salts and other inorganic ions.
g) Bone cells called osteoclasts responsible for dissolving of the matrix as it is laid down
enable reconstruction and remodeling of the bone during endochondral ossification.
h) An artery, a vein and a lymph vessel pass through a haversian canal of a compact one
allowing the passage of nutrients, respiratory gases and metabolic wastes towards and
away from the bone cells.
i) Each lacuna has fine cytoplasmic extensions called canaliculi which pass through
lamellae and make connections with other lacunae and with the central haversian canal,
allowing communication between the lacunae in different lamellae, and with the central
haversian canal.
j) Presence of numerous nerve fibres in the haversian canal allows co-ordination of bone
reconstruction enabling each bone to adapt its structure to meet any change in mechanical
requirement of an animal during its development.
k) Bone releases calcium and phosphate into the bloodstream as required by the body under
the control of the hormones parathormone and calcitonin.
l) Spongy bone has spaces between the trabeculae, reducing the weight of the bone,
allowing less restricted movement and locomotion.

Differences between a cartilage and bone

Cartilage Bone
No process extended from each lacuna into the Lacuna possesses canaliculi that extend into th

21
matrix. e matrix.
No blood vessels and nerves in the tissue. Blood vessels and nerves run through the have
rsian canal.
Exchange of material between chondrocytes oc Osteoblasts exchange materials by help of bloo
curs by diffusion. d capillaries passing through the canaliculi into
the lacunae.
Elastic and compressible Relatively incompressible as the matrix is high
ly composed of minerals eg calcium ions, mag
nesium ions.
Matrix is relatively semitransparent with hyali Matrix is opaque
ne cartilage and semi opaque in yellow elastic
cartilage.
The matrix is not calcified. Matrix is calcified with greater quantities of M
g2+, Na+, Ca2+ etc.
No concentric layers of lamellae and no havers Consist of concentric layers of lamellae surrou
ian canals present. nding the haversian canal.
Rather inactive. An active tissue with metabolic activity.
Matrix secreting cells are called chondroblasts. Matrix secreting cells are called osteoblasts.
It is differentiated into hyaline, white fibrous a It is structurally differentiated into compact an
nd yellow elastic cartilage. d spongy bones
It is flexible due to relatively soft and flexible It is rigid due to solid matrix called osteon.
matrix called chondrin.
It is less strong. It is stronger.
Chondroblasts are randomly scattered in the m Osteoblasts are in concentric layers around the
atrix and occur in singles, pairs or fours. haversian canal.
It is mostly found in areas where cushioning is It is located in areas where maximum support i
required. s needed.

MUSCLE TISSUE

There are three types of muscular tissue which include: voluntary muscle, involuntary muscle
and cardiac muscle.

Basic structure of a muscular tissue

a) All muscle fibres are made up of elongated and thin cells called muscle cells or muscle
fibres.
b) The muscle fibres contain a specialized cytoplasm called sarcoplasm that contains a
network of membranes called sarcoplasmic reticulum.
c) Muscle fibres may be bound by a cell membrane called sarcolemma.
d) Each muscle may contain numerous thin myofibrils.

SKELETAL/VOLUNTARY/STRIATED OR STRIPED MUSCLE

22
It is said to be striated because its muscle cells have transverse stripes when viewed in longitudin
al section.

Distribution of skeletal muscles

It is found attached to the skeleton in the head, trunk and limbs hence the name skeletal muscle.

Gross structure of a skeletal muscle

The skeletal muscle is attached to a bone in at least two places namely; the origin, a fixed non-m
oveable part of the skeleton and the insertion, a moveable part of the skeleton.

Attachment is by a means of tough relatively inelastic tendons made up of almost entirely collag
en fibres. At one end, the tendon is continuous with the outer covering of the muscle while the ot
her end is of the tendon combines with the outer layer of the bone called periosteum and forms a
very firm attachment.

The skeletal muscle is composed of bundles of muscle fibres each surrounded by a connective tis
sue, endomysium. Each bundle of muscle fibres is surrounded by perimysium, a connective tiss
ue, the various bundles are surrounded by an epimysium, a connective tissue sheath.

Skeletal muscle showing the layers of connective tissue

Histology of a striated muscle (skeletal muscle)

The muscle is made up of many hundreds of long muscle cells called muscle fibres. Each muscl
e fibre is filled with a cytoplasm called sarcoplasm in which about 100 nuclei are spaced out eve
nly just beneath the bounding membrane called sarcolemma.

In the sarcoplasm, there are many thin myofibrils which possess characteristic cross striations. T
he myofibrils line up perpendicular with the cross striations next to each other. The myofibrils ar
e composed of protein filaments called actin and myosin.

23
 Fine structure of a striated muscle

Each myofibril is divided into light and dark bands. The light band has a comparatively light regi
on in the middle called H- zone, and it has dark regions on either sides. In the middle of the H- z
one is a dark line called the M- line.

Running through the light bands in the middle is the Z- line. The dark and light bands are called
A and I bands respectively. I means isotropic, as it allows light to pass through and so appears lig
hter. A means anisotropic as it does not allow light to pass through, so it appears darker.

The region of a myofibril between two Z-lines is called a sarcomere and is described as the func
tional basic unit of a myofibril. Alternating light and dark bands are due to two types of protein fi
laments which run longitudinally. These are the thin actin and thick myosin protein filaments. Th
e thick myosin filaments are confined to the dark band and the thin actin protein filaments occur
in the light band but extend in between the thick myosin filaments within the dark band.

The segments on either side of the H- zone are due to both thick myosin and thin actin filaments
overlapping. The H-zone consists only thick myosin protein filaments. The thin actin filaments al
one are found in the light band.

Within each muscle fibre, there is an internal membrane system, the sarcoplasmic reticulum sur
rounding the myofibrils. The sarcoplasmic reticulum includes a system of transverse tubules (the
T- system) which run into the muscle fibre from the sarcolemma at positions corresponding to th
e Z- lines.

Connected with the T- system are vesicles containing calcium ions in high concentration. Ca2+hel
ps in hydrolysis of ATP. After muscle contraction, calcium ions are actively removed into the lo
ngitudinal tubules thereby lowering the concentration to a level below that at which ATP hydroly
sis can occur.

24
Diagram of a myofibril to show the transverse and longitudinal tubules of the sarcoplasmic
reticulum (FA page322)

There are four blood capillaries surrounding each muscle fibre. Each branch of the axon terminat
es at a plate like structure called neuromuscular junction. The motor end plate forms the neuro
ne-to-muscle synapse, the connection between the motor neurone and the muscle fibre.

All the muscles served by the same motor neurone are called motor unit because they work as a
unit contracting and relaxing at the same time. The motor unit is the basic functional unit of a ske
letal muscle.

Actin and myosin filaments

Actin filaments

This consists of a fibrous protein (F- actin) together with two accession proteins, tropomyosin a
nd troponin, both of which are globular.

Tropomyosin forms 2 helical strands around the F- actin in a longitudinal section. This arrangem
ent brings about relaxation and contraction of the muscle. Troponin binds to the tropomyosin and
calcium ions.

Myosin filament

It consists of two distinct regions, a long rod shaped region called a myosin rod, and a myosinhe
ad. When a muscle is at rest, the tropomyosin blocks the sites to which myosin attaches. When c
alcium ions are released from the sarcoplasmic reticulum, they bind to troponin causing the tropo
myosin to move away from the myosin binding site.

When excitation of muscles by nerve impulses stops, Ca2+ are pumped back into the sarcoplasmi
c reticulum and the muscle relaxes.

The sliding filament hypothesis of muscle contraction

The theory proposes that during muscle contraction, the thick and thin actin filaments slide past e
ach other leading to shortening of sarcomeres.

25
The dark bands/A- bands remain the same length, and the light bands (I- bands) and the H- zone
get shorter, then the Z- lines move closer together.

In the absence of calcium ions, tropomyosin blocks access of myosin heads to the binding site of
an actin filament. When a nerve impulse through a motor neurone arrives at the neuromuscular j
unction at the surface of the muscle fibre, the action potential at the motor end plate is propagate
d through the transverse tubules to the vesicles. This causes the release of calcium ions from the
vesicles down their concentration gradient in the sarcoplasm.

The calcium ions bind to troponin forming a calcium troponin complex. The positions of tropon
in and tropomyosin are altered such that the myosin head has access to its binding site on the fibr
ous actin (F- actin) forming actomyosin.

Myosin hydrolyses ATP in presence of ATPase enzyme on myosin head and undergoes conform
ational changes into a higher energy state. The myosin head binds into actin forming a cross brid
ge between the actin and myosin filaments.

Each bridge detouches itself from the thin filaments and retouches itself to another site further al
ong, and the cycle of cross bridge formation is repeated through a chain of ratchet mechanism lea
ding to muscle contraction.

When calcium ion level decreases, troponin blocks tropomyosin in the blocking position and acti
n filament slide back to their respective state.

Diagrams illustrating the mechanism by which actin filaments slide past the myosin filame
nts

26
After each bridge has completed its movement, it detouches itself from the thin filament and reatt
aches to another site further along the actin. The cycle is repeated at 50- 100 times per second.

The combined movement of many bridges has the effect of pulling the thin filaments past the thi
ck ones resulting in contraction of the muscle. Contraction of the muscle is said to be brought ab
out by the bridges going through a kind of ratchet mechanism as illustrated in the diagram below.

Diagram summarizing the proposed ratchet mechanism during muscle contraction

Diagrams from the electron micrographs confirming the sliding filament theory

Stretched/relaxed myofibril

Contracted myofibril

27
 On contraction

 Length of the dark band/A- band/anisotropic band remains the same.

 Length of the light band/I band/isotropic band shortens.
 Length of the H-zone shortens.
 Length of the dark ends of the dark band increases.
 Overall length of the myofibril shortens.

NB: When the muscle is severely contracted, the ends of the thin filaments may touch the M- lin
e and the H-zone disappears. In that condition, the filaments (thick and thin) uniformly overlap i
n the whole length of fibres.

Transverse section of the H-zone Transverse section of the end of the dark band

Transverse section of the light band

Q Describe the mechanism of muscle contraction

Arrival of the impulse at the neuromuscular junction depolarizes the presynaptic membrane and i
ncreases its permeability to towards calcium ions. This leads to diffusion of calcium ions from th
e synaptic cleft into the synaptic knob via the presynaptic membrane.

Ca2+ induces the synaptic vesicles to fuse with the presynaptic membrane and release the neurotr
ansmitter substance eg acetylcholine into the synaptic cleft by exocytosis. Acetylcholine settles o
n the receptor sites of the post synaptic membrane and induces the post synaptic membrane to op
en the sodium channels. Sodium ions rapidly diffuse through the synaptic cleft into the sarcoplas
m.

28
This depolarises the post synaptic membrane until a wave of depolarization moves across the sar
colemma to the T-system of the sarcoplasmic reticulum. The vesicles connected to the transverse
tubules are stimulated to secrete calcium ions into the sarcoplasm.

Ca2+ combines with troponin to form calcium troponin complex which leads to a conformational
change in the structure of tropomyosin such that it vacates/leaves the active sites of the actin fila
ment. Myosin heads bind on the sites to form actomyosin. This activates ATPase enzyme on the
myosin head to catalyse the hydrolysis of ATP into ADP and energy is released.

This energy is used by the myosin heads to bend away from 900C to 450C thereby pulling the acti
n filament inwards towards the centre of the sarcomeres while sliding past the stationary myosin
filament. This causes the following changes;

 Decrease in the length of the H- zone.

 Decrease in the length of the sarcomere.
 Decrease in the length of the I- band.
 Increase in the length of the dark ends of the A- band.

ATP molecules attach on the myosin heads to provide them with energy to detouch themselves fr
om the sites and reattach at another site and continue to pull the actin filaments until maximum c
ontraction.

Muscle relaxation

When the muscle is no longer stimulated by an impulse/wave of depolarization, calcium ions are
pumped back into the sarcoplasmic reticulum using the calcium pump and energy from hydrolysi
s of ATP until its concentration fall below the contractile threshold in the sarcoplasm. This result
s into detouchment of the myosin heads from the actin binding site using energy from the hydrol
ysis of ATP to ADP.

The actin filaments revert to their original position and tropomyosin reoccupies the actin binding
site. The length of the sarcomere, I- band and H-zone increase back to their original and that of t
he dark regions of the dark band decrease back to their original position.

Adaptation of skeletal muscle tissue for its function/relationship between structure and fun
ction of a striated muscle

 It consists of elongated fibres, allowing considerable contractile length.

 Its fibres are parallel to give it maximum contractile effect and to allow each fibre to be
controlled individually which gives ability to vary the length of the whole muscle
contraction necessary for proper control of skeletal movement.
 The ends of the muscle fibre are tapered and interwoven with each other to provide
adequate mechanical strength during muscle contraction.

29
 Its cells contain a large number of mitochondria to provide large amounts of ATP for
muscle contraction.
 In their arrangements, the actin and myosin filaments fit into each other to allow them
slide over each other to cause contraction.
 The cells have a rich blood supply to provide adequate supply of oxygen and nutrients.
 The muscle cells have myoglobin to store oxygen and release it for respiration when
blood oxygen levels are low.
 Has a specialized region called the motor end plate where the axon of a motor neurone
divides and forms fine non myelinated branches (dendrites) ending in synaptic knobs
running in shallow troughs on the sarcolemma allowing nervous stimulation and control
of the muscle.
 The sarcolemma folds inwards and forms a system of tubes called the T- system
(transverse tubules) which run parallel through the sarcoplasm to the Z- lines allowing a
nerve impulse arriving along a motor neurone at the neuromuscular junction at the
surface of a muscle fibre to be propagated as a wave of depolarization (action potential)
through the T- system causing release of calcium ions of the sarcoplasmic reticulum to
activate the process of muscle contraction.
 The specialized endoplasmic reticulum of the muscle fibre called the sarcoplasmic
reticulum forms the vesicles at the Z- line of the sarcomeres which contain calcium ions
used to activate the process of muscle contraction.
 Ability to generate ATP using phosphocreatine during anaerobic conditions for a constant
supply of ATP in the muscle.
 Ability to respire anaerobically for continued muscle contraction in anaerobic conditions.

VISCERAL/INVOLUNTARY/UNSTRIATED/UNSTRIPED/SMOOTH MUSCLE

 It consists of muscle cells called muscle fibres which are spindle shaped and tapering at
both ends and uninucleated.
 The nucleus is single, elongated in shape, centrally placed and surrounded by little
sarcoplasm.
 The muscle fibres lack a sarcolemma.
 Each muscle fibre consists of numerous inconspicuous, fine contractile myofibrils
arranged longitudinally.
 The actin and myosin filaments are evenly distributed hence there are no striations or
light and dark bands.

30
  Smooth muscle fibres are shorter than striated muscle fibres.
 Has sarcoplasmic reticulum but less extensive than in striated muscle.
 Has rings of smooth muscle fibres called sphincter muscle fibres for example; pyloric,
cardiac and anal sphincters.
 Has prominent mitochondria but less numerous than in striated muscle.

Innervations and activity of the smooth muscle

 Smooth muscle is involuntary in action, so cannot be moved by ones will.

 Innervated by two sets of nerves from the autonomic nervous system (sympathetic and
parasympathetic).
 Smooth muscle fibres undergo prolonged and slow, sustained rhythmical contractions and
relaxations as in peristalsis, hence fatigues slowly.

The smooth muscle is located in the tracts of the intestines, genitals, urinary and respiratory syste
ms and the walls of blood vessels

Functions of the smooth muscle

i. The anal sphincter controls the elimination of feaces from the body.
ii. The pyloric sphincter controls passage of food from the stomach to the duodenum.
iii. Small sphincter muscle surrounds some blood vessels to control the distribution of
blood and regulation of blood pressure.
iv. Control movement of materials with the body visceral organs.

CARDIAC MUSCLE

It is found only in the heart.

The structure of a cardiac muscle

A cardiac muscle consists of a network of interconnected cells called cardiac muscle fibres. Each
muscle fibre is short, cylindrical and branched. Each muscle fibre possesses one large mitochon
drion, with one nucleus or two nuclei, abundant cytoplasm, glycogen granules, well developed T
system and poorly developed endoplasmic reticulum consisting of a network of tubules.

Cardiac muscle fibres are terminally branched and connected to each other by intercalated discs.
Actin and myosin filaments are regularly arranged to give faint but regular cross striations. Musc
le fibres branch and cross connect with each other to form a complex netlike arrangement.

31
 Illustrations of the structure of a cardiac muscle (FA page 171)

Innervations and functioning of the cardiac muscle

 Cardiac muscle is myogenic meaning that the contractions are developed within the
muscle.
 The rate of contraction can be influenced by the autonomic nervous system.
 Interconnections between the fibres (intercalated discs) ensure a rapid and uniform spread
of the excitation.
 Have rhythmic rapid contractions and relaxation with a long refractory period and so do
not fatigue as contraction is not sustained.
 Need a constant supply of large amounts of energy.
 A small number of cardiac muscle fibres and a few nerve endings form the Sino atrial
node (SAN) located near the opening of the vena cava which stimulates heart beat on
their own.

Adaptations of the cardiac muscles to their function

a) Cardiac muscle cells are highly branched terminally and connected to each other by
intercalated discs to form a network that allows rapid spread of waves of electrical
excitation from cell to cell, so that linked muscle cells rapidly contract rhythmically and
simultaneously for fast heartbeat.
b) Dense network of blood capillaries ensures adequate supply of oxygen and food nutrients,
for fast production of adequate ATP, for continuous rapid muscle contraction and rapid
excretion of carbon dioxide and other metabolic wastes.
c) Numerous large mitochondria and glycogen granules rapidly provide adequate amounts
of energy in form of adenosine triphosphate (ATP) by aerobic respiration for rapid

32
 contraction without fatigue.
d) Has the Sino atrial node (SAN) which emits waves of electrical excitation that initiate
continuous and rhythmic contraction without fatigue, for continuous heartbeat.
e) Have striations for mechanical strength to support its fast and continuous contractions.
f) Undergoes rapid rhythmic contractions and relaxations with long refractory periods and
thus does not fatigue as contraction is not sustained.
g) Well-developed T-system for rapid transmission of impulses thus rapid contraction and
relaxation.
h) Branched muscle fibres offer a large surface area for fast spread of waves of electrical
excitation for continuous contraction hence continuous heartbeat.

A comparison of voluntary, involuntary and cardiac muscles (similarities and differences)

Feature Striated muscle Smooth muscle Cardiac muscle

Names Striated, striped, voluntar Unstriated, unstriped, s Heart muscle, cardiac mu
y muscle mooth, involuntary, no scle
n-striated muscle
Specialization Most highly specialised Least specialised More specialised that sm
ooth muscle
Shape Elongated, cylindrical an Spindle shaped muscle Elongated cylindrical and
d unbranched muscle fib fibres with tapered end branched muscle fibres.
res. s.
Arrangement Arranged in bundles. Arranged in bundles, s Interconnected to form a
heets or rings. network.
Nucleus Multinucleated myofibril Uninucleated myofibril One or two nuclei in bet
s with peripherally locate s with centrally located ween intercalated discs.
d nucleus. elongated nucleus.
Cytoplasmic c Numerous mitochondria Prominent but less mit Numerous large mitocho
ontents in rows at the periphery ochondria. ndria
and between fibres.
Prominent smooth endop Individual tubules of s Poorly developed SER c
lasmic reticulum forming mooth endoplasmic reti onsisting of a network of
a network of tubules. T- culum. tubules.
System well developed.

Glycogen granules and s Glycogen granules pres Glycogen granules prese

ome lipid droplets. ent. nt.
Blood supply Rich blood supply. Poor blood supply. Rich blood supply.
Striations or ba Striations of light and da No striations or bands. Faint regular striations pr
nds rk bands esent.
Intercalated di Absent Absent Present
scs
Myofilaments/ Very conspicuous Inconspicuous Conspicuous
Myofibrils

33
Innervations Under control of the volu Under control of the au Myogenic, but rate of
ntary nervous system. tonomic nervous syste contraction can be
m. influenced by the
autonomic nervous syste
m.
Contractions Powerful rapid contracti Shows sustained slow Continuous rapid rhythm
on with short refractory and rhythmic contracti ical contraction and relax
period (rest period). ons and relaxation with ation with long refractory
a long refractory perio period hence contraction
d. not sustained.
Fatigue Fatigues quickly and easi Does not fatigue easily. Does not fatigue.
ly.
Location Attached to the skeleton In the walls of intestine Only in the walls of the h
in the head, trunk and li s of genital, urinary an eart.
mbs. d respiratory tracts, in
walls of blood vessels.

Energy Large amount of energy Much energy needed b Needs a constant and goo
needed at once. ut constant supply requ d energy supply.
ired.
Sarcolemma Present Absent Present
Mode of worki Voluntary Involuntary Involuntary
ng

NERVOUS TISSUE

It contains densely packed nerve cells called neurons which are specialized for conduction of ner
ve impulses. The neurons are the basic functional units of the nervous system.

Types of neurons/nerve cells

 Sensory/afferent neurons:- These conduct impulses from the receptors to the central
nervous system.
 Motor /efferent neurones:- These conduct impulses from the central nervous system to
the effectors.
 Relay/intermediate neurons:- These transmit impulses from the sensory neuron to the
motor neurone. They are only found in the central nervous system.

Diagram of a motor neuron

34
 Diagram of a sensory neuron

Diagram of a relay neuron

NB: The process which brings impulses towards the cell body is called a Dendron and the one w
hich conducts impulses from the cell body is called the axon

Assignment Compare the structure of a motor neurone and a sensory neurone

Structure of a neuron

Each neuron posses a cell body and cytoplasmic extensions (nerve fibers’). Each cell body contai
ns a nucleus and abundant granular cytoplasm. The cytoplasm also contains prominent conical gr
anules called Nissl’s granules which are groups of ribosomes and rough endoplasmic reticulum r
ich in RNA and associated with protein synthesis.

From the cell body, extends out two types of cytoplasmic extensions; a Dendron and axon. Depe
nding on the number and arrangement of these processes, the neurons are said to be unipolar, pse
udounipolar and multipolar.

Unipolar:- It is a neuron with the axon as the only large branch from the cell body eg arthropod
motor neuron

Bipolar neuron:- It is one where two processes, an axon and a Dendron project from the cell bo
dy. Examples of bipolar neurons are found in the retina of the mammalian eye.

35
Multipolar neuron:- It is one which has one axon and several dendrons from the cell body. An e
xample is the motor neuron

Pseudounipolar:- It is where the cell body is not found along the axis/end of the axon. Instead, it
is connected by a short side branch of the axon eg sensory neuron

PLANT TISSUES

1) MERISTEMATIC TISSUE

It is a plant tissue consisting of actively dividing cells which give rise to cells that differentiate in
to new tissues of the plant.

Meristem

A meristem is a group of cells which remain with the ability to divide by mitosis producing daug
hter cells which grow to form the rest of the plant body.

Types of meristems

Apical meristems:- They are found at the shoot tip and root tip. They divide continuously by mit
osis leading to primary growth of the plant body that is increase in length of the shoot or root.

Lateral meristems (cambium):- They are responsible for secondary growth of the shoot and root i
e increase in girth. They include the vascular cambium which gives rise to secondary vascular ti
ssue including secondary xylem and phloem. They also include the cork cambium (phellogen)
which gives rise to cork (phellem) which replaces the epidermis.

Intercalary meristems:-

 They allow growth in length in regions other than the root and shoot tips.
 Ensures continued growth where tissues are damaged such as when eaten by herbivores
in grasses.

SIMPLE PLANT TISSUES

36
They are tissues consisting of one type of cell. They include the parenchyma, collenchyma and s
clerenchyma

2) PARENCHYMA

It consists of living cells which are relatively undifferentiated. The cells are either roughly spheri
cal or elongated. The cells have thin cell walls made up of cellulose, pectins and hemicelluloses.
The cells also have large sap vacuoles with dense but peripheral cytoplasm.

Parenchyma tissue is located in the cortex, pith and medullary rays of wood. It also serves as a pa
cking tissue in xylem and phloem

Functions of the parenchyma tissue

a) Acts as a packing tissue ie cells of the parenchyma fill spaces between other specialized
tissues eg in the cortex, pith, between the xylem vessels and phloem.
b) It contains intercellular air spaces which allow gaseous exchange.
c) When they are turgid, parenchyma cells become closely packed thus provide support for
the organs in which they occur. For example in the leaves and in stems of herbaceous
plants.
d) It is a storage tissue due to possession of starch granules and large food vacuoles.
Therefore, the tissue is abundant in storage organs eg the Irish potato.
e) It allows transportation of materials through cells by symplast pathway or apoplast
pathway.
f) The parenchyma tissue is metabolically active as it is composed living cells for example
some parenchyma are photosynthetic.
g) Growth of the pericycle in the roots where it retains the meristematic activity producing
lateral roots and contributing to secondary growth.
h) In the endodermis, cells are covered by a fatty substance (suberin) that forms the
casparian strip that prevents apoplast transportation of water through the root.

Transverse section of a parenchyma tissue

Relationship between structure and function of the parenchyma tissue

i) The cells are unspecialized to perform a variety of functions.

37
 j) Many intercellular spaces to allow diffusion and exchange of gases.
k) Thin cellulose cell walls to allow passage of materials for transport.
l) Transparent cell walls to allow light penetration for photosynthesis.
m) The cells are large and contain large vacuoles with a thin layer of cytoplasm to provide
storage space for materials of the plant.
n) Have isodiametric, roughly spherical or elongated cells to serve as a packing material
between specialized cells.
o) Cells have permeable walls to allow entry of light for photosynthesis.
p) Cells have leucoplasts such as amyloplasts to store food such as starch.
q) Cells have chloroplasts to allow photosynthesis.
r) Cell walls contain cellulose, pectins and hemicelluloses for support.
s) The cells have chromoplasts such as in petals to provide bright colour to attract insects
for pollination.

Modified parenchyma

They include; epidermis, mesophyll, endodermis, pericycle, companion cells and transfer cells.

a) Epidermis/epidermal cells

It is a layer of one cell thick that covers the whole primary plant body.

Functions

 The basic function is to protect the plant body from desiccation and infection. This is
achieved by secreting cutin and forms the cuticle that is impervious to water.
 Specialized epidermal cells (the guard cells) bound/guard the stomata and are important
in opening and closing of stomata.
 Hair like structures on cuticle (epidermis) serve various purposes./ for example, root hairs
increase on the surface area for absorption of water and mineral salts by the roots.
 Hooked hairs of climbing stems prevent them from slipping off their supports.
 Glandular cells on the cuticle secrete sticky substance that traps and kills insects and they
may also secrete scent.
 The epidermal hairs of leaves reduce water loss from the plant as well as reflecting the
sun’s radiations.
 Being transparent, the epidermis allows passage of light in the mesophyll cells for
photosynthesis.
 The epidermis may develop hairs which form a barrier around the nectories of flowers
preventing access to crowling insects and promoting cross pollination.

Q State the various modifications of the epidermis to serve different functions

b) Mesophyll cells (chlorenchyma)

38
Mesophyll is a packing tissue located between the upper and lower epidermis of leaves. There ar
e two types of mesophyll cells.

 Palisade mesophyll cells- They are located in the upper layer called the palisade
mesophyll layer. Cells are elongated and columnar in shape. They contain a large number
of chloroplasts. The cells are tightly packed with very few and narrow air spaces.
 Spongy mesophyll cells- They are located in the lower layer called the spongy mesophyll
layer. Cells are spherical and irregularly shaped with fewer chloroplasts. They possess
large intercellular air spaces between the cells.

The functions of the mesophyll include: photosynthesis, gaseous exchange and Storage of star
ch

Adaptations of the mesophyll to its function

i. Palisade mesophyll cells are column shaped with numerous chloroplasts in a thin
layer of cytoplasm to carry out photosynthesis.
ii. Palisade mesophyll cells are tightly packed together forming a continuous layer that
traps incoming light.
iii. The chloroplasts within the mesophyll cells can move towards light allowing them to
be in the best positions to receive light.
iv. Chlorophyll within the chloroplasts is contained within the grana, where it is arranged
on the sides of a series of unit membrane allowing chlorophyll to receive maximum
light.
v. The structural arrangement of chlorophyll on the photosynthetic membrane brings
chlorophyll in close proximity to other pigments such as carotenoids and enzymes
necessary for its functioning in light harvesting.
vi. Spongy mesophyll cells are irregularly shaped hence fit together loosely leaving large
air spaces to allow efficient gaseous exchange via the stomata.
vii. The mesophyll cells contain numerous amyloplasts for storing starch.
c) Endodermis

It is the inner most layer of the cortex surrounding the vascular tissue of the roots and stems. It c
onsists of living, elongated and flattened cells. The cell wall of endodermal cells comprises cellul
ose, pectins, hemicelluloses and deposits of suberin

Functions of the endodermis

 Acts as a selective barrier to movement of water and mineral salts between the cortex and

39
 xylem in roots.
 In dicot stems, it stores starch forming a starch sheath with a possible role in the gravity
response of stems.

Adaptations of the endodermis to its functions

 The endodermis of roots has the casparian strip (made up of suberin) which is
impermeable to water and prevents water and solutes from flowing through the air spaces
of the cell walls of the endodermal cells (apoplast pathway). This forces water through
the cell surface membrane into the cytoplasm of the endodermal cells, hence allowing the
endodermal cells to regulate the movement of solutes through the xylem.
 Active pumping of salts by endodermal cells into the xylem allows rapid movement of
water by osmosis into the xylem leading to a buildup of root pressure.
 Control of movement of water and solutes by endodermal cells acts as a protective
measure against the entry of pathogens and toxic substances into the xylem.
 In dicots, the endodemal cells contain amyloplasts for storing starch grains forming a
starch sheath.
d) Pericycle

It is a layer of modified parenchyma, one to several cells thick, located in roots between the centr
al vascular tissue and the endodermis. It consists of one to several layers of living, roughly spheri
cal and elongated cells. Their cell walls are composed of cellulose, pectins and hemicelluloses.

Functions of the pericycle

 Produces lateral roots.

 Contributes to secondary growth

Adaptations of the pericycle to its function

 It retains its capacity for cell division (meristematic activity) to produce lateral roots.
 Due to its meristematic activity, it contributes to secondary thickening of the roots.
e) Companion cells

They are specialized parenchyma cells found adjacent to the sieve tubes.

They have a prominent nucleus, dense cytoplasm with numerous small vacuoles, plastids and the
usual cell organelles. They are metabolically very active with numerous mitochondria and ribos
omes. Each companion cell is connected to a sieve element by plasmodesmata.

Functions of the companion cells

 Control of the activity of the adjacent metabolically inactive sieve tube elements.

40
  Provide energy needed for the active processes which occur during translocation of
organic solutes in the sieve tubes

Adaptations of the companion cells to their function

 Plasmodesmata connect sieve elements with companion cells allowing communication

and exchange of materials between companion cells and sieve tube elements.
 Companion cells have large nucleus to effect metabolic activity over both companion
cells and sieve tubes.
 Companion cells contain numerous mitochondria to produce energy for active transport
of materials in the sieve elements.
f) Transfer cells

They are modified form of parenchyma companion cells found next to the sieve tube. They have
numerous internal projections of the cell wall formed by extra thickening of the cell wall. They p
osses numerous mitochondria in the dense cytoplasm

The function of transfer cells is active uptake of salts from neighbouring cells.6cells.6e

Adaptations of transfer cells to their function

 Numerous internal projections of the cell wall increase the surface area of the cell wall
and bring it to closer association with the cytoplasm.
 Numerous mitochondria in thee cytoplasm provide energy for active transport of organic
solutes such as sugars from neighbouring cells.
 Have a large amount of starch granules which are broken down to glucose for aerobic
respiration.
3) COLLENCHYMAu

Collenchyma consists of living cells modified to give support and mechanical strength. The colle
nchyma is the first mechanical tissue to develop in the primary plant body.

Structure of collenchyma tissue

It consists of living cells. The cell walls consist of cellulose, pectins and hemicelluloses. The cell
s are closely packed without air spaces between them. The cells are elongated and polygonal wit
h tapering ends.

The cells have extra deposits of cellulose at the corners of the cells causing uneven thickening of
the cell walls. Cells are elongated, parallel to the longitudinal axis of the in which they are found.

Collenchyma tissue cells in transverse

41
 Distribution and functions

Collenchyma cells which are relatively flexible provide support for plant organs allowing them t
o bend without breaking.

It is mainly found in young plants, herbaceous plants and in organs such as the leaves in which s
econdary growth does not occur.

In the leaves, they are mainly found in the midribs of dicotyledonous leaves.

They are also located at the periphery of the organs usually under the epidermis

Adaptations of collenchyma to its function

Deposition of extra cellulose at the corner of the cells leads to development of unevenly thickene
d cell walls to provide support and mechanical strength.

Cells are living and can grow and stretch, thus provide mechanical strength without imposing lim
itations on the growth of the other cells around it, allowing continued growth in young stems and
leaves.

Cells are located towards the periphery of the organ just below the epidermis in the outer regions
of the cortex to increase its support value in stems andm peandmitiole.

4) SCLERENCHYMA

They are living and can only elongate when they are young. The mature cells are dead, incapable
of elongation and contain no cytoplasm. The primary cell wall is composed of cellulose, pectins,
hemicelluloses and thickened with deposits of lignin. Its thick cell walls contain simple pits, are
as where lignin is not deposited on primary wall due to presence of a group of plasmodesmata.

There are two types of sclerenchyma cells that is fibres and sclereids

42
 Fibres

The cells are elongated and hollow with narrow lumens. The cells are polygonal in shape with ta
pering interlocking ends. The fibres are found in the outer regions of the cortex, pericycle of ste
ms, xylem and phloem. Its structure is as illustrated below

Sclereids (stone cells) which are roughly spherical or irregular in shape they are found in the cor
tex, pith, phloem, shells and stones of fruits, seed coats. Its structure is as illustrated below

Adaptations of sclerenchyma to its function

 Have elongated fibres and spherical sclereids closely packed together to provide
mechanical support.
 The primary cell wall is heavily thickened and lignified with heavy deposits of lignin,
with great tensile strength and compression strength for support and mechanical
protection.
 High tensile strength of lignified walls prevents breakage on stretching.
 High compression strength of the lignified walls prevents buckling or crushing under
pressure.
 Fibres are arranged into strands or sheets of tissue that extends longitudinally to provide
combined collective strength.
 Ends of cells of fibres interlock with the tapering ends of one another increasing
combined supportive strength.

43
44
 XYLEM

It is a vascular tissue with two main functions; conductance of water and mineral salts and provid
ing mechanical support to the plant. The xylem consists of four types of cells; tracheids, vessels e
lements, parenchyma and fibres.

Tracheids

They are single cells with thick walls extensively lignified by heavy deposits of lignin. Some part
s are not lignified forming bordered pits. They have tapering end walls that overlap with adjacent
tracheids. They are dead with empty lumens when mature. The cells are polygonal in cross secti
on with 5 or 6 sides.

The tracheids represent the original primitive water conducting cells of vascular plants. They are
the only cells found in the xylem of the more ancestral vascular plants and are the only conductin
g elements in conifers.

Structure of a tracheid

Vessels

They are very long tubular structures formed by fusion of several elongated cells (vessel element
s) end to end in a row. Vessel elements are shorter and wider than tracheids. A vessel is formed
when neighbouring vessel elements of a given row fuse as a result of their end walls breaking do
wn. The walls of the vessels are heavily lignified, but with unlignified portions called bordered p
its.

Differences between vessels and tracheids

Vessels Tracheids
They are cylindrical in shape. They are 5 or 6 sided in cross section

45
Have open ended walls on either sides. Have perforated closed end walls.
Have no tapering ends. Have tapering ends.
Offer less resistance towards water passage. Offer a significant or more resistance to water
passage.
Fast conduction of large volume of water. Conduction of less volume of water.

Protoxylem and metaxylem

The first vessels of the xylem to form are the protoxylem. They have the following characteristic
s.

 They are small in cross section.

 They are located in the inner most region of the of the xylem.
 They are found on the part of the apex just below the apical meristem where elongation
of surrounding cells is still taking place.
 They are still young.
 They are not fully lignified.
 They undergo stretching.
 They collapse as they mature.

Metaxylem

 They are the recently formed vessels of the xylem

 They have larger cross section.
 They are located further out in the stems.
 They undergo more extensive lignifications.
 Mature metaxylem vessels cannot stretch or grow because they are dead, rigid and fully
lignified tubules.
 They become part of the permanent tissues of the plant.

Illustration

NB: 1.The xylem fibres provide additional mechanical strength to the xylem.

2. The xylem parenchyma serve functions such as food storage, deposition of tannins, crystals an
d other chemical compounds, radial transport of food and water and gaseous exchange through in
tercellular spaces.

Adaptations of xylem to its functions

1. Xylem vessels and tracheids consist of long cylindrical cells joined end to end, hence are
continuous with each other ensuring continuous flow of water in a continuous unbroken
column.
2. End walls of the xylem vessels are completely broken down to form continuous tubes that

46
 allow uninterrupted flow of water.
3. Tracheids have tapering, elongated, sloping end walls that overlap with adjacent tracheids
perforated with large cellulose lined bordered pits that allow water to pass from one cell
to another.
4. During development, the protoplasmic contents of vessels and tracheids die leaving
empty hollow lumens, permitting uninterrupted flow of water without obstruction by
living content.
5. Side walls of the vessels and tracheids are perforated with numerous pits permitting
lateral/sidewise flow of water and salts in and out of the lumen where necessary.
6. Impregnation of cellulose walls with lignin increases adhesion of water molecules to
walls thereby facilitating the rise of water by capillarity.
7. Lignifications of walls confer rigidity preventing walls from collapsing under large
tension forces set up by the transpiration pull.
8. Narrowness of lumens of vessels and tracheids increases the rise of water by capillarity.
9. Extreme narrowness of lumens of tracheids and vessels offer a high tensile strength
preventing xylem from collapsing during transport of water through them.
10. Vessels, tracheids and fibres are dead at maturity, and thus provide mechanical strength
and support to the plant.
11. Xylem fibres have extremely thick walls which are heavily lignified and with narrow
lumens to provide additional mechanical strength and support to the xylem.

5) PHLOEM

It is a vascular tissue modified for translocation of manufactured food. It is composed of five

types of cells ie sieve tube elements, companion cells, parenchyma, fibres and sclereids.

Sieve tubes

They are long tube like structures formed by end to end fusion of cells called sieve tube elem
ents/sieve elements. The sieve elements have walls made of cellulose and pectins, but their n
uclei degenerate and are lost as they mature.

The cytoplasm is confined to a thin layer around the periphery of the cell. Sieve elements are
living but metabolically depend on adjacent companion cells. In between the sieve tube elem
ents are sieve plates, formed from the two adjoining end walls of neighbouring sieve element
s.

The sieve plates are perforated by sieve pores formed by enlargement of plasmodesmata. The
sieve plates are made up of a polysaccharide called callose.

Companion cells

47
They have a thin cell wall and dense cytoplasm with a prominent large nucleus, numerous mi
tochondria, plastids and small vacuoles and extensive endoplasmic reticulum. Companion cel
ls are metabolically active and essential for the survival of sieve elements.

Structure

Protophloem and metaphloem

Protophloem is the first phloem formed in the zone of elongation of the growing root or stem.
Protophloem becomes stretched and eventually collapses becoming non functional as the tiss
ue around it grows. As more phloem is produced, the protophloem matures after elongation h
as stopped to produce the metaphloem.

Adaptations of the phloem to its function

 Sieve tube elements are joined end to end, their walls are perforated with sieve pores in
sieve plates allowing passage of materials unimpeded from one cell to another.
 Sieve elements lack nuclei and possess a thin cytoplasm pushed to the sides of the cell,
creating room for passage of organic materials in solution with minimal obstruction.
 Plasmodesmata connect sieve elements to companion cells which are metabolically active
allowing communication and exchange of materials between sieve elements and
companion cells.
 Sieve elements contain cytoplasmic filaments continuous with similar filaments in other
sieve elements via sieve pores in the sieve plate, which consist of a contractile phloem
protein capable of streaming and sliding organic materials from one sieve element to
another by wave like movements of the filaments.
 Companion cells contain dense cytoplasm with numerous mitochondria, a large nucleus,
plastids, small vacuoles and extensive endoplasmic reticulum making them metabolically
active to meet the metabolic needs of the sieve elements.

48
  Companion cells possess numerous mitochondria to provide energy in form of ATP for
active transport of materials.
 Modified parenchyma companion cells called transfer cells found next to sieve tubes bear
numerous internal projections increasing surface area of the cell membrane.
 Transfer cells also contain numerous mitochondria for active uptake of solutes from
neighbouring cells during loading of sieve tubes.
 Phloem consist of living cells allowing live active transport of materials since the
mechanism of loading sieve tubes and transport of solutes requires energy.
 Sclereids are lignified to provide support to the vascular tissue of the phloem.
 The companion cells are elongated and thin walled to provide a large surface area for
diffusion of materials to the neighbouring cells.
 The presence of meristematic parenchyma for development of new cells.

LEVELS OF ORGANISATION

1. Unicellular level of organization

It is one where the physiological functions/processes of an organism are mainly performed by cel
l organelles. It is represented by single celled organisms such as paramecium.

Paramecium (structure FA page 43)

It has the following main parts and their functions.

 Macronucleus- It controls metabolic functions including growth.

 Micronucleus- It is responsible for sexual reproduction.
 Cilia- It is responsible for locomotion and creation of food currents.
 Contractile vacuoles (anterior and posterior vacuoles)- Are for osmoregulation. They
eliminate excess water from the cell to the exterior.
 Trichocysts- They are tiny explosive sacs containing needle shaped thread. They are used
for defence in some species. They are used for paralyzing the prey.
 Food vacuoles- They contain food particles at the base of cytoproct and it is where
digestion takes place.
 Cytoproct- They are sites through which undigested food is expelled to the exterior.

Advantages of unicellular organization

i. The distance from the surface of the organism to its centre is very short, that allows
quick and efficient diffusion of materials in and out of the cell.
ii. It exposes a large surface area to volume ratio which also makes diffusion of
materials easy.
iii. Easy reproduction through fission.

49
 Disadvantages

i. There is inefficiency in the processes of an organism due to lack of specialization.

ii. It is not possible for an organism to grow to a large size.

Advantages of small size

i. It allows fast locomotion.

ii. It presents a larger surface area to volume ratio hence allowing faster exchange of
gases with the surrounding medium.
iii. Organisms can easily pass through limited spaces.
iv. The organism can occupy a variety of habitats.
v. It has a high reproductive rate with a high survival chance.
vi. Organisms cannot easily be noticed by the predator/enemy.

Disadvantages of small size

i. The organisms usually have a high reproductive rate hence eat a lot.
ii. The organisms lose a lot of heat because of having a large surface area to volume
ratio.
iii. The organism cannot intimidate/scare the predator/enemy.
iv. The organism can easily be consumed by the predator wholly.

Advantages of big size

i. It allows the organism to have a low metabolic rate hence eats less.
ii. The organism loses less heat because it has a small surface area to volume ratio.
iii. The organism can easily intimidate/scare the predator/enemy.
iv. A large organism cannot easily be consumed, only parts of it can be consumed.

Disadvantages of big size

i. The locomotion is usually slow.

ii. It presents a small surface area hence slower exchange of materials with the
surrounding medium.
iii. There is a problem of passing through limited space hence can occupy limited
habitats.
iv. It usually allows a low reproductive rate with low survival chances.
v. A large organism can easily be noticed by the predator/enemy.
2. Tissue level of organization

It is the level of organization where the physiological processes are only carried out by isolated c
ells and tissues. Organisms in this level are mainly primitive multicellular animals. Such animals
have very few organs. Example is the hydra.

50
 HYDRA

It has a simple sac like body. Its cells are interdependent and to some extent integrated by the ner
vous system.

Structure of the wall of the hydra (FA page 44)

The wall of the hydra

It is composed of seven different types of cells arranged in two sheet like layers called ectoderm
and endoderm. The ectoderm lines the external surface whereas the endoderm lines the body cavi
ty/enteron.

Between the ectoderm and endoderm is an intersurface called the mesogloea in which there are n
erve cells that are interconnected to form a nerve net. Each of the different cells performs a speci
fic function as stated below.

 Musculo epithelial cells- These are cells lining the upper surface in the ectoderm and are
mainly for protection.
 Nematoblasts- (stinging cells). It contains a thread which contains a toxic fluid. It is used
in the piercing and poisoning of prey.
 Sensory cells- They are for detecting changes in the surrounding environment and pass
impulses to the nerve cells.

51
  Nerve cells- They are interconnected to form a nerve net. They are connected to the
contractile muscle tail of the musculo epithelial cell. Therefore, their role is to pass on the
information ie conducting impulses to those muscle tails of the musculo epithelial cells.
 Muscle tails- They are contractile muscles at the base of each musculo epithelial cell.
They are concerned with bringing about the movement or beating of the tentacles and
flagella when they contract.
 Glandular cells- They secrete enzymes that carry out digestion in the enteron.
 Phagocytic musculo epithelial cells- For taking in small particles from the enteron so that
digestion is completed intracellularly.
 Flagellated musculo epithelial cells- Are important for effective movement of materials
eg food materials and stirring it up in the enteron.
 Interstitial cells- For generation or production of new cells.

Feeding mechanism in a hydra

Food of a hydra is usually water fleas. The water fleas are immobilized by the stinging cells and
pulled to the mouth by the by the tentacles under the influence of the contraction of the muscle ta
ils which are controlled by the nerve net in the mesogloea.

When the food reaches in the enteron, digestion starts by the enzymes secreted from the glandula
r cells. Digestion is completed inside the musculo epithelial cells which take up food particles fro
m the enteron by phagocytosis.

3. Colonial organization

In colonial organization, cells are functionally isolated ie their activities are not co-ordinated. Ea
ch cell when isolated can exist on its own for example in a sponge. (structure FA page 45)

52
The various cells and their functions include:

a) Mesenchyme cells- These secrete siliceous spicule for strengthening the wall of the
sponge.
b) Collar cells- These maintain flow of water by beating of their flagella.
c) Wandering amoebocytes- They move around between the other cells.
d) Hollow pore cell- It forms lining of pore through which water containing food particles
is drawn into sponge.
e) Epithelial cells- They line the surface of the body
4. Organ level of organization

It is a group of organization where the physiological functions are mainly performed by o

rgans and organ systems. An example is mammals. This form of organization occurs mai
nly in the higher multicellular animals.

Advantages of being multicellular

a) It allows increase in size.

53
b) It brings about specialization of specific cells for a particular function, thus improving
efficiency of an organism as a whole. It permits exploitation of environment in which
single celled organisms cannot live.
c) It allows development of better tissues for example muscles for quick movement,
skeleton for support and quick movement.
d) There is also development of sophiscated physiological mechanisms eg maintenance
of body temperature.

54