Advance Plant anatomy / Master student Introduction ,Plant cell structure/protoplast

Dr. HADEEL AHMED AL-AMERI

INTRODUCTION TO PLANT ANATOMY

Plant anatomy or phytotomy is the general study of the
internal structure of plants (shape, structure, and size ). Originally it
included plant morphology, the description of the physical form and
external structure of plants, but since the mid-20th century plant anatomy
has been considered a separate field referring only to internal plant
structure. Plant anatomy is now frequently investigated at the cellular
level, and often involves the sectioning of tissues and microscopy.
Some studies of plant anatomy use a systems approach, organized
on the basis of the plant's activities, such as nutrient transport, flowering,
pollination, embryogenesis or seed development. Others are more
classically divided into the following structural categories:
A- Flower anatomy : including study of the Calyx, Corolla, Androecium,
and Gynoecium.
B- Leaf anatomy : including study of the Epidermis, stomata and Palisade
cells.
C- Stem anatomy : including stem structure and vascular tissues buds
and shoot apex.
D- Fruit/Seed anatomy : including structure of the ovule, seed, Pericarp
and Accessory fruit.
E- Wood anatomy : including structure of the Bark, Cork, Xylem,
Phloem, Vascular cambium, Heartwood and sapwood and branch
collar,
F- Root anatomy : including Structure of the root, Root tip, Endodermis.
Because of plant anatomy as a part of botany (the study of plants),
and It is focuses on the structural or body parts and systems that make up
a plant, thus a typical plant body consists of three major vegetative
organs: the root, the stem, and the leaf, as well as a set of reproductive
parts of that include flowers, fruits, and seeds.

1- THE ROOTS
A plant's roots, like the foundation of a skyscraper, help it to stay
upright. They also absorb water and dissolved minerals from the ground
and give the plant what it needs to make its own food. Most roots grow
underground and move downward because of the influence of gravity,
although the roots of some water plants float. Other root systems, like that
of the English ivy, actually attach themselves to a vertical surface and
allow the plant to climb. There are two main types of root systems:
taproot and fibrous. Plants that have taproots grow a single, long root that
penetrates straight down and firmly anchors the plant.
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Advance Plant anatomy / Master student Introduction ,Plant cell structure/protoplast
Dr. HADEEL AHMED AL-AMERI

Trees and dandelions have

taproots that serve this function.
Fibrous roots are shorter and more
shallow and form a branching
network. Grass has a fibrous root
system that grows at a shallow
level and in all directions. Inside a
root are pipelines or veins that
carry water and minerals to the rest
of the plant. These pipes are
concentrated in the center of the
root, like the lead in the center of a
pencil. At the end of each root is a
cap that protects it as it pushes
farther into the soil. Extending
from the sides of the root, but
further back from the root cap are
root hairs. These hairs are the main
water and oxygen absorbing parts
of a plant. Materials enter and
leave roots by two main processes:
diffusion and osmosis. When
molecules are distributed
unequally, nature always seeks a
balance and molecules will move
from an area of high concentration
to one of low concentration. When
the cells of a root hair have little
oxygen and the soil around the root
hair has a lot, oxygen will move
from the soil to the root
automatically without the plant
having to expend any energy.
Osmosis is a similar situation (from
high to low concentration), but it
occurs when molecules, like those
of water, move across a membrane
that will not allow other materials
to pass. Like diffusion, osmosis
does not require the plant to use
any energy.

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Advance Plant anatomy / Master student Introduction ,Plant cell structure/protoplast
Dr. HADEEL AHMED AL-AMERI
2- THE STEMS
Plant stems perform two functions. They support the parts of the
plant aboveground (usually the buds, leaves, and flowers), and they carry
water and food from place to place within the plant itself. A stem is made
up of an outer layer, the epidermis; an inner layer, the cortex; and a
central zone called the pith. The stem of a green plant holds itself up by
having thousands of cells lined up next to and on top of each other. As the
cells take in water, they expand like a full balloon, and since their walls
are elastic, they stretch very tight against each other and against the stem
wall. It is their pressure that holds the stem up. A plant droops when its
cells lack water and have begun to shrink. Woody plants, like trees, also
contain a material called lignin that strengthen cell walls and make them
more rigid. A plant's stem also functions as its circulatory system and
uses what is called vascular tissue to form long tubes through which
materials move from the roots to the leaves and from the leaves to the
roots.
3- THE LEAVES
A leaf (plural leaves) is the principal lateral appendage of
the vascular plant stem usually borne above ground and specialized
for photosynthesis. The leaves and stem together form the shoot. Leaves
are collectively referred to as foliage, as in "autumn foliage". In most
leaves, the primary photosynthetic tissue, the palisade mesophyll, is
located on the upper side of the blade or lamina of the leaf but in some
species, including the mature foliage of Eucalyptus, palisade mesophyll is
present on both sides and the leaves are said to be isobilateral. Most
leaves are flattened and have distinct upper (adaxial) and lower (abaxial)
surfaces that differ in color, hairiness, the number of stomata (pores that
intake and output gases), the amount and structure of epicuticular
wax and other features. Leaves are mostly green in color due to the
presence of a compound called chlorophyll that is essential for
photosynthesis as it absorbs light energy from the sun. A leaf with white
patches or edges is called a variegated leaf.
Leaves can have many different shapes, sizes, and textures. The
broad, flat leaves with complex venation of flowering plants are known
as megaphylls and the species that bear them, the majority, as broad-
leaved or megaphyllous plants. In the clubmosses, with different
evolutionary origins, the leaves are simple (with only a single vein) and
are known as microphylls. Some leaves, such as bulb scales, are not
above ground. In many aquatic species, the leaves are submerged in
water. Succulent plants often have thick juicy leaves, but some leaves are
without major photosynthetic function and may be dead at maturity, as in

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Advance Plant anatomy / Master student Introduction ,Plant cell structure/protoplast
Dr. HADEEL AHMED AL-AMERI
some cataphylls and spines. Furthermore, several kinds of leaf-like
structures found in vascular plants are not totally homologous with them.
Examples include flattened plant stems called phylloclades and cladodes,
and flattened leaf stems called phyllodes which differ from leaves both in
their structure and origin. Some structures of non-vascular plants look
and function much like leaves. Examples include
the phyllids of mosses and liverworts.
4- FLOWERS AND SEEDS
The reproductive part of a seed-producing plant is called the
flower. Flowers have male and female cells that produce a seed when
they unite. The stamen is the male reproductive organ in a flower and
contains the male cells (pollen) in its anther that grows at the tip of its
long, narrow stalk. The pistil is the female reproductive organ and looks
like a long-necked bottle. It has a round base containing the ovary, a
slender tube or long neck called the style, and a flattened, sticky top
called the stigma. Once a flower opens, its petals (which are a type of
leaf) protect the sex organs and serve to help pollination (the transfer of
pollen to the female parts) by attracting animals like bees and birds.
When this happens, fertilization occurs and the ovaries become seeds.
Seeds have three main parts: the coat, the embryo, and the food
storage tissue. The coat protects the embryo, which is the beginning of a
plant and grows by using food stored in the seed. Most seeds are enclosed
in fruit that can be dry like a ripe bean pod, or fleshy like an apple or a
peach. Other plants, like fir trees, have naked or uncovered seeds that
form on the upper side of the scales that make up a pine cone. All are
designed to be scattered as far as possible from the parent plant to ensure
the further survival of the species.

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Advance Plant anatomy / Master student Introduction ,Plant cell structure/protoplast
Dr. HADEEL AHMED AL-AMERI
Plant life cycle
Germination Depending on
the type of seed, it may or may
not require soil or light to
germinate. However, most all
plants need water in order for this
process to occur. As water is
absorbed by the seed, it begins to
expand or swell, eventually
cracking or splitting the seed
coat. Once germination occurs,
the new plant will gradually
begin to emerge. The root, which
anchors the plant to the soil,
grows downward. This also
enables the plant to take up water
and nutrients required for growth.
The shoot then grows upward as
it reaches for light. Once the
shoot reaches the surface, it
becomes a sprout. The sprout will
eventually take on a green color
(chlorophyll) upon developing its
first leaves, at which time the
plant becomes a seedling.

Basic Plant Life Cycle: Seedlings, Flowers, & Pollination Once the
seedling develops these first leaves, it is able to make its own food
through photosynthesis. Light is important for this process to occur, as
this is where the plant gets its energy. As it grows and becomes stronger,
the seedling changes into a young adult plant, with many leaves. Over
time, the young plant will begin to produce buds at the growing tips.
These will eventually open up into flowers. Pollination must occur in
order for fertilization to happen, which creates new seeds.
The flowers transform into fruiting bodies, which protect the
numerous seeds that are inside. As the seeds mature or ripen, the flowers
will eventually fade away or drop. Once the seeds have dried, they are
ready to be planted (or stored), repeating the life cycle of a flowering
plant all over again.

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Advance Plant anatomy / Master student Introduction ,Plant cell structure/protoplast
Dr. HADEEL AHMED AL-AMERI

PLANT CELL STRUCTURE

Plant cell is the structural and functional unit in plant. The cell consist of
two main parts :
1- Cell wall
2- Protoplast

Protoplasts are spherical naked plant cells produced by the removal of the
cell wall with digestive enzymes. They are usually derived from either
leaf tissue or from cell suspension cultures and have been isolated from a
wide variety of plant species.
The cytoplasm along with the cellular organelles is
called protoplasm i.e., protoplasm = Cytoplasm + nucleus. It does not
include cell membrane or cell wall. Protoplast is the living part of cell and
does not include the cell wall. Protoplast = cell membrane + cytoplasm +
nucleus.

I- Cell Membrane Function and Structure

The cell membrane (plasma membrane) is a thin semi-permeable
membrane that surrounds the cytoplasm of a cell. Its function is to protect
the integrity of the interior of the cell by allowing certain substances into
the cell while keeping other substances out. It also serves as a base of
attachment for the cytoskeleton in some organisms and the cell wall in

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Advance Plant anatomy / Master student Introduction ,Plant cell structure/protoplast
Dr. HADEEL AHMED AL-AMERI
others. Thus the cell membrane also serves to help support the cell and
help maintain its shape.
Another function of the membrane is to regulate cell growth
through the balance of endocytosis and exocytosis. In endocytosis, lipids
and proteins are removed from the cell membrane as substances are
internalized. In exocytosis, vesicles containing lipids and proteins fuse
with the cell membrane increasing cell size. Animal cells, plant
cells, prokaryotic cells, and fungal cells have plasma membranes. Internal
organelles are also encased by membranes.
The cell membrane is primarily composed of a mix
of proteins and lipids. Depending on the membrane’s location and role in
the body, lipids can make up anywhere from 20 to 80 percent of the
membrane, with the remainder being proteins. While lipids help to give
membranes their flexibility, proteins monitor and maintain the cell's
chemical climate and assist in the transfer of molecules across the
membrane.

II- Living Components of plant cell

The substance except nucleus surrounded by the plasma lemma of cell is
known as cytoplasm. Its contains :

1- Endoplasmic reticulum
Cytoplasm contains an extensive network of membrane enclosed
spaces; these spaces along with the membranes enclosing them are known
as endoplasmic reticulum (ER).

2- Ribosomes
Ribosomes are particles of about 200 A° diameter; they are
composed of RNA and protein. Generally ribosomes are attached to the
outer surfaces of ER membranes. This converts smooth ER elements into
rough ER.

3- Golgi body
It consists of 2-7 flat cisternae stacked close to each other. Golgi
bodies originate from ER elements.

4- Lysosomes
Lysosomes are vesicles of 400-800 μm formed by budding of
Golgi bodies and they contain hydrolytic enzymes.

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Advance Plant anatomy / Master student Introduction ,Plant cell structure/protoplast
Dr. HADEEL AHMED AL-AMERI

The main enzyme present in lysosomes is acid phosphatase, other

enzymes are acid DNAase, acid RNAase and β galactosidase etc.

5- Mitochondria
These are cylindrical bodies with an average diameter of 0.2 to 1 μ
and ordinarily 3-10 μ in length.

6- Nucleus:
It is a denser, rounded or spherical protoplasmic body enclosed in
the protoplasm. Its shape and size differs greatly according to size of cell.
It is composed of following organelles:
a- Nuclear Membrane: Nucleus is bounded by a membrane on the
outside called nuclear membrane, which is double walled and having
numerous minute pores.
b- Nucleoplasm: It is viscous, non-staining, granular, colorless fluid
inside the nuclear membrane. It is also known as nuclear sap or
karyolymph.
c- Chromatin network: The threads like bodies forming a reticulum are
suspended in nucleoplasm, which are network of chromosomes.
d- Nucleolus: A spherical round body usually single but may be double.
It plays important role in protein synthesis. It is associated with a
particular nuclear organizing chromosome.

7- Plastids
Plastids are the organelles which are peculiar to plant cells. Plastids
that contain high concentration of carotenoid pigments are called
‘chromoplasts’. They give yellow, orange and red colors to many fruits
(tomato), roots (carrot) and flower petals.
Nonpigmented plastids are called ‘leucoplasts’. An important type
of leucoplast is ‘amyoplast’ which is a starch-storing plastid.
Chloroplasts are the plastids that contain green pigment,
chlorophyll. They are found in green tissues of plant, especially leaf.
They are absent in roots. The chloroplast is surrounded by the inner and
outer membranes.
Chloroplasts also contain third system of membrane called
thylakoid. All the chlorophyll is contained within this membrane, which
is the site of light reactions of photosynthesis. Thylakoid membranes are
highly folded and appear like stacked coins. These stacked membranes
are known as grana lamellae (or grana thylakoid). The membranes

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Advance Plant anatomy / Master student Introduction ,Plant cell structure/protoplast
Dr. HADEEL AHMED AL-AMERI
without stacking are known as stroma lamellae (or stroma thylakoid).
Each stack is called granum, the inner space within a thylakoid is known
as lumen, the region of the chloroplast that is inside the inner membrane
and surrounds thylakoids is known as stroma. The carbon reactions take
place in stroma. Chloroplasts contain their own DNA and protein-
synthesizing machinery. Chloroplast genome is smaller (145 kb) than
mitochondrial genome (200 kb). Chloroplast DNA is circular and histone-
free. Ribosomes occur free in stroma or bound to the outer surface of
thylakoid membrane. As is mitochondria, most of the chloroplast’s
proteins are encoded by nuclear genes, synthesized in cytosol and
transported to organelle.

III- Non-living components of plant cell

1-Vacuoles
Vacuoles are organelles bounded by a single membrane, the
tonoplast, or vacuolar membrane. They are multifunctional organelles and
are widely diverse in form, size, content, and functional dynamics. A
single cell may contain more than one kind of vacuole. Some vacuoles
function primarily as storage organelles, others as lytic compartments,
vacuoles are involved with the breakdown of macromolecules and the
recycling of components within the cell. Entire organelles, such as
senescent plastids and mitochondria, may be engulfed and subsequently
degraded by Vacuoles are organelles bounded by a single membrane
vacuoles containing large numbers of hydrolytic and oxidizing enzymes.
Many meristematic plant cells contain numerous small vacuoles. In the
mature cell as much as 90% of the volume may be taken up by the
vacuole, with the rest of the cytoplasm consisting of a thin peripheral
layer closely pressed against the cell wall. Being a selectively permeable
membrane, the tonoplast is involved with the regulation of osmotic
phenomena associated with the vacuoles.

The principal component of the non–protein-storing vacuoles is

water, with other components varying according to the type of plant,
organ, and cell and their developmental and physiological state. In
addition to inorganic ions such as Ca2+, Cl−, K+, Na+, NO3, such
vacuoles commonly contain sugars, organic acids, and amino acids, and
the aqueous solution commonly is called cell sap.

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Advance Plant anatomy / Master student Introduction ,Plant cell structure/protoplast
Dr. HADEEL AHMED AL-AMERI
2- Starch grains
A starch is the most
abundant carbohydrate in the plant
world. Moreover it is the principal
storage polysaccharide in plants.
During photosynthesis assimilatory
starch is formed in chloroplasts.
Later it is broken down into
sugars, transported to storage cells,
a re-synthesized as storage starch
in amyloplasts. As mentioned
previously, an amyloplast may
contain one (simple) or more
(compound) starch grains. If
several starch grains develop
together, they may become
enclosed in common outer layers,
forming a complex starch grain.
Starch grains, or granules, are
varied in shape and size and
commonly show layering around a
point, the hilum, which may be the
center of the grain or to one side.
The layering of starch grains is
attributed to an alternation of these
two polysaccharide molecules.

The layering is accentuated when the starch grain is placed in water

because of differential swelling of the two substances: amylose is soluble
in water, and amylopectin is not. Storage starch occurs widely in the plant
body. It is found in parenchyma cells of the cortex, pith, and vascular
tissues of roots and stems; in parenchyma cells of fleshy leaves (bulb
scales), rhizomes, tubers, corms, fruits, and cotyledons; and in the
endosperm of seeds.

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Advance Plant anatomy / Master student Introduction ,Plant cell structure/protoplast
Dr. HADEEL AHMED AL-AMERI
3- Aleurone grains

It is storage proteins that found in

wheat, which considerable part of
the prolamins aggregate directly
into it within the rough ER and
then are transported in distinct
vesicles to the vacuoles without
Golgi involvement. Structurally
consist of an amorphous
proteinaceous matrix surrounded
by a bounding membrane. Other
protein bodies may contain one
or more non-proteinaceous
globoids or one or more globoids
and one or more protein
crystalloids, in addition to the
proteinaceous matrix.

3- Oil bodies
Are more or less spherical structures that impart a granular
appearance to the cytoplasm of a plant cell when viewed with the light
microscope. Oil bodies are widely distributed throughout the plant body
but are most abundant in fruits and seeds. Approximately 45% of the
weight of sun flower, peanut, and sesame seed is composed of oil. The oil
provides energy and a source of carbon to the developing seedling.

4- Crystals
Inorganic deposits in plants consist mostly of calcium salts and
anhydrides of silica. Among the calcium salts, the most common is
calcium oxalate.

a- Calcium oxalate:
It occurs in the majority of plant families, notable exceptions being
the Cucurbitaceae and some families of Liliales, Poales, and all
Alismatidae . Calcium oxalate occurs as mono- and dihydrate salts in
many crystalline forms. The monohydrate is the more stable and is more
commonly found in plants than is the dihydrate. The most common forms
of calcium oxalate crystals are:

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Advance Plant anatomy / Master student Introduction ,Plant cell structure/protoplast
Dr. HADEEL AHMED AL-AMERI

(1) Prismatic crystals, variously

shaped prisms, usually one per
cell. Occur Prismatic crystals in
Allium cepa.

(2) Raphide, needle-shaped

crystals that occur in bundles
of Lemna sp.

(3) Druses, spherical aggregates

of prismatic crystals.

(4) Styloids, elongated crystals with pointed or ridged ends, one or two to
a cell.

(5) Crystal sand

Very small crystals, usually in masses. In some tissues calcium

oxalate crystals arise in cells that resemble adjacent, crystal-free cells. In
others, the crystals are formed in cells called crystal idioblasts specialized
to produce crystals. Crystal idioblasts contain an abundance of ER and
Golgi bodies. Most crystal cells are probably alive at maturity. The
location and type of calcium oxalate crystals within a given taxon may be

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Advance Plant anatomy / Master student Introduction ,Plant cell structure/protoplast
Dr. HADEEL AHMED AL-AMERI
very consistent and, hence, useful in taxonomic classification. Calcium
oxalate crystals usually develop in vacuoles.

2- Calcium carbonate crystals

They are not common in
seed plants. The best known
calcium carbonate formations are
cystoliths, which are formed in
specialized enlarged cells called
lithocysts of the ground
parenchyma and epidermis. The
cystolith develops outside the
plasma membrane in association
with the cell wall of the lithocyst.
Callose, cellulose, silica, and
pectic substances also enter into
the composition of cystoliths,
which are confined to a limited
number (14) of plant families.
Found in Fucus elastic.

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Advance Plant anatomy / Master student Plant cell structure/cell wall
Dr. HADEEL AHMED AL-AMERI
Plant Cell Structure
CELL WALL
One of the most distinctive characteristics of plant cells with
respect to those of animals is the cell wall, which some authors
consider as the extracellular matrix of plants. It is the structure that R.
Hook saw and drew when he gave name to the cell. The cell wall is
located externally to the plasma membrane, is synthetized by the cell
itself and consists mainly of cellulose, in addition to other
polysaccharides and glycoproteins. Different cell types present
differences in composition and organization in their cell wall,
depending on the function they are developing. In fact, the different
cell types of plants can be identified by the characteristics of their cell
wall.

Diagram of a typical parenchymatic cell with its primary cell wall.

The cell wall has a mechanical function. It is responsible for the

shape and size of plant cells and provides them with both structural
support and protection as an exoskeleton. As a result, it is also
responsible for the rigidity of the plant and for keeping its aerial
structures and the organs that form it upright. Another of its main
functions is to be a means of communication and transport of
molecules and water between cells, both between nearby and distant
cells. It also participates in the fight against pathogens being able to
trigger defense responses, or give texture to the tissues of the fruits.
Morphologically, the cell wall is formed by layers or sheets. All cells
have at least two: middle lamella and primary wall. The middle lamella
is synthetized and shared by cells that are contiguous with each other,
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Advance Plant anatomy / Master student Plant cell structure/cell wall
Dr. HADEEL AHMED AL-AMERI
while the primary cell wall is synthetized and belongs to each cell. In
some plant cells, a third thicker layer called secondary cell wall is
deposited. Most of the wood in the trees correspond to the secondary
cell wall.

Images of cells with primary cell wall and with secondary cell wall (the
latter also have primary cell wall, although it is not visible). All of them
have middle lamella, which is not distinguished in the images. They
correspond to conductive vessels: the one on the left stained with
toluidine blue and the one on the right with safranin - fast green.

Diagram of the primary and secondary cell walls. S1, S2, and S3 are
layers of the secondary cell wall.

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Advance Plant anatomy / Master student Plant cell structure/cell wall
Dr. HADEEL AHMED AL-AMERI
MIDDLE LAMELLA
The outermost layer of the cell wall and the first to form is the
middle lamella. It acts as a glue that binds neighboring cells. It is the
only layer synthetized and shared by two contiguous cells. It has an
amorphous appearance and is very thin; its thickness is close to the
resolution limit of the light microscope. Nevertheless, it does not
appear as a well-defined layer even with the electron microscope. In
some tissues, there are intercellular spaces and, therefore, the middle
lamella has one of their surfaces free. The middle lamella consists
mainly of pectins, although it can be lignified in those cells that have a
secondary cell wall.

PRIMARY CELL WALL

The primary cell wall is the first clearly visible layer of the cell
wall and it is located between the plasma membrane and the middle
lamella or, in some cells, between the secondary wall and the middle
lamella. It is responsible for the initial shape and size of the plant cell,
and determines its subsequent changes in shape and size. It appears
in all plant cells and originates during cell division, but the primary
cell wall is also synthetized during growth in cell size. In addition,
through the life of the cell there is a recycling with degradation and
new synthesis of the components of the primary cell wall. In cells that
are metabolically active, secretory, or capable of dividing, it remains
the only component of the cell wall, in addition to the middle lamella.
Plant cells stop growing when the primary cell wall become rigid,
which may be due to a change in their composition.

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Advance Plant anatomy / Master student Plant cell structure/cell wall
Dr. HADEEL AHMED AL-AMERI

Primary cell walls (arrows) in different tissues. The shape and size of the
cells are conditioned by their cell wall.

The cell wall is a barrier to the free diffusion of molecules between

cells; of course, it is much less permissive than the extracellular matrix
of animals. However, cells need to communicate with each other. To
do this, plant cells create channels that cross the cell walls and allow
direct communication between adjacent cytoplasms. These channels
are called plasmodesmata. With very few exceptions, all cells in a
plant are connected with their neighbors by plasmodesmata. This
connection is literal, that is, the plasma membranes of neighboring
cells are continuous with each other, so we could say that a plant is a
large syncytium. Plasmodesmata can appear concentrated in certain
areas of the cell wall forming what are called primary pore fields,
which form depressions in the cell wall since its thickness is smaller.

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Advance Plant anatomy / Master student Plant cell structure/cell wall
Dr. HADEEL AHMED AL-AMERI

Plasmodesmata in the primary cell wall. In the lower part of the figure,
there is a drawing of an electron microphotography.
In the primary cell wall there are also areas with a greater thinness,
although they do not produce interruptions in it, called primary pits;
these structures may appear isolated or grouped in areas called primary
pit fields. Plasmodesmata are usually concentrated in primary pit
fields.

Composition
The primary cell wall is composed, considering the dry weight, by
25-30% of cellulose, 30% of hemicellulose, 35% of pectins and 1-5%
of glycoproteins. The proportion and types of components that
constitute the primary cell walls varies between cell types. It has been
estimated that more than 2000 genes are required for the synthesis and
remodeling of the primary cell wall.
Cells with primary cell wall are usually metabolically active.
Usually the primary cell wall is thin, around 0.1 µm thick. In addition,
the cells that develop secondary cell wall usually have thin primary
cell wall. Only a few cells achieve thick primary cell walls, such as
some endosperm and collenchyma cells. In any case, the thickness can
change according to the conditions in which the cell is located. In the
primary cell wall there are pores (not to be confuse with
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Advance Plant anatomy / Master student Plant cell structure/cell wall
Dr. HADEEL AHMED AL-AMERI
plasmodesmata) with a diameter ranging from 4 to 8 nm, which allow
the passage of water, ions and small molecules such as sugars and
amino acids, and hormones.
Cellulose. Cellulose is the main component of plant walls. It is a
linear polysaccharide formed by glucose. The formula is (C6H10O5)n,
where n can be greater to 500 per polysaccharide chain. The long
cellulose molecules associate with each other through hydrogen bonds
and Van der Waals forces to form structures called cellulose
microfibrills, formed by 50 cellulose molecules oriented with the same
polarity. Microfibrills are associated with each other through bonds
formed between them and other carbohydrates, mainly hemicellulose
and pectins, which result in fibrils and cellulose fibers, visible under
the light microscope. The cellulose fibers can measure about 0.5 µm in
diameter and 4 to 7 µm in length. The strength of cellulose fibers is
similar to that of steel and the bonds between cellulose molecules by
hydrogen bonds make the cellulose microfibrills have
crystalline properties in some regions, while the rest acquires
paracrystalline properties.

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Advance Plant anatomy / Master student Plant cell structure/cell wall
Dr. HADEEL AHMED AL-AMERI

Organization of cellulose molecules. The glycoproteins have not been

represented. The detailed organization of the cellulose microfibrills has
not been solved yet and two models were proposed. In one of them
(model 2) pectins play a predominant role.

As with hyaluronate (hyaluronic acid), cellulose is synthetized in

the plasma membrane thanks to the action of cellulose synthase, a

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Advance Plant anatomy / Master student Plant cell structure/cell wall
Dr. HADEEL AHMED AL-AMERI
transmembrane protein with an amino acid sequence that crosses 8
times the plasma membrane. Up to 36 cellulose synthase enzymes bind
at a point on the plasma membrane to form the so-called cellulose
synthase complex that is rosette-shaped, and is so large that it can be
observed with the electron microscope. Each rosette can synthetize up
to 36 molecules of glucose simultaneously. The cellulose molecules
that polymerize nearby are joined laterally by hydrogen bonds. These
new cellulose molecules are also associated with the microfibrills that
had previously formed piles of these microfibrills, fibrils and cellulose
fibers.

Molecular composition of the xyloglucan, the most common

hemicellulose.
Hemicellulose is actually a family of polysaccharides of 200 to
500 monosaccharides. The type of hemicellulose that appears in the cell
wall varies greatly between tissues and cell types. It is synthetized in the
Golgi apparatus and is transported to the plasma membrane in vesicles,
where it is released by exocytosis. Xyloglucan is the most frequent
molecule of hemicellulose. Structurally, hemicellulose is similar to
cellulose so it can establish hydrogen bonds with it. As the
hemicellulose molecules are synthetized, they coat the cellulose
microfibrills, helping to the cohesion to form cellulose fibers.
Pectins : form a very diverse group of acidic
polysaccharides synthetized in the Golgi apparatus and secreted into the
cell wall. Together they form a gel-like structure that is located between
the cellulose microfibrills. They seem the main responsible for the
formation of pores that allow the diffusion of small molecules through
the primary wall. Pectins are more abundant in the dicotyledons that in

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the monocotyledons. For example, grasses contain only traces of
pectins. Pectins appear absent in the secondary cell wall (see below).
Glycoproteins of the cell wall are usually rich in proline,
hydroxyproline and glycine, amino acids that are found in much
repeated sequences. The type of glycoprotein usually varies a lot
between cell types. One of the most common family of glycoproteins
are extensins, which are rich in prolines. The glycoproteins have an
apparent structural function, although there are also enzymes such as
peroxidases, laccases, phosphatases, cellulases, pectidases, among
others.
Callose is a substance that is deposited between the plasma
membrane and the cell wall; then, it cannot be considered strictly as a
component of the primary cell wall. It is mainly located around
the openings of the plasmodesmata . The callose is synthetized,
released and deposited in response to cellular stress, either by wounds
or by pathogens, and its mission is to obliterate the plasmodesma
channel and cut or decrease the communication between neighboring
cells. It also appears in other places with less clear functions, such as in
the pollen tubes or in the cell plate during cytokinesis.
Some specialized cells have other particular molecules. For
example, cutine and other waxy polymers are deposited in the free
surface of the epidermal cells forming a structure called cuticle. This
layer, which can be very thick, prevents the loss of water and protects
against pathogens. The suberin is deposited in the cell wall of other
cells such as those of the suber or cork of the periderm, in the root
endodermis and in cells that surround some nerves of the leaves.
Suberin has two domains, one is inserted into the primary wall and the
other is between the primary wall and the plasma membrane.
It can be said that the primary cell wall is a framework of cellulose
microfibrills, connected by hemicellulose molecules and embedded in
a pectin matrix. The three-dimensional organization of cellulose,
hemicellulose and pectins is not clear yet. Several models have been
proposed and the most cited assumes that hemicellulose molecules
bind tightly to cellulose molecules by hydrogen bonds. However,
recently it has been seen that hemicellulose does not have as many
connections with the cellulose as was thought and that pectins seem to
play a more important role in the compaction of the cell wall. For
example, it seems that pectins have more links with cellulose than
hemicellulose with cellulose. Pectins help to hydrate the primary cell
wall.

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Cell growth
When the cell wall grows, it is necessary to distinguish between
growth in thickness (deposition of successive layers) and increase in
length, when the cell wall increases its surface and the cell increases in
size. The growth of the plants is mainly due to cell size growth, which
is produced by the force of the hydrostatic pressure, that is, by the
force exerted by the water from inside the cell outwards on the primary
cell wall. Although it depends on the cell type, cells can grow up to 10
times their size. In some cases up to 100 times.
The cells can grow homotrophically, the entire surface of the cell
wall expands although it can be at different rates, or heterotropic, only
a part of the surface of the cell wall expands (for example in the pollen
tubes, or in the trichomes). A cell grows where there is less resistance,
which depends on the resistance opposed by the cell wall. This
conditions where the cell of the plant will grow, which will determine,
for example, the shape of the stems and leaves, or that they will grow
towards a light source or not. The resistance of the primary cell wall is
determined by the orientation of the fibers of cellulose and by the
consistency of the primary cell wall assembly.
The primary cell wall is anisotropic, a consequence of the irregular
orientation of the fibers of cellulose. These fibers are shorter and more
irregular that in the secondary cell wall (see below). Usually this
irregular orientation occurs in cells that grow or have grown in all
directions. When a cell expands in a preferred direction, the
microfibrills of cellulose are oriented perpendicular, in the manner of
rings, with respect to the growth axis, and the external ones, which
were already there, are arranged longitudinally to that axis. It is normal
to find layers in the primary cell wall where the microfibrills are
oriented helically and with a certain rotation of angle with respect to
the next layers.
An interesting aspect of the primary cell wall synthesis is how the
cell manages to orient the molecules and microfibrills of cellulose,
since this will determine the orientation of the fibrils and fibers of
cellulose. The orientation of the molecules of cellulose is conditioned
by their synthesis and by how they are deposited on the plasma
membrane, which is determined by the spaces through which the
enzymatic complex that synthetizes it can move through the plasma
membrane: the rosette of cellulose synthase. One theory suggests that
this movement depends on the orientation of the cortical microtubules
that are located just under the plasma membrane, in the cytosol. These
microtubules act like barriers that cannot be crossed by the rosettes of
cellulose synthase. The enzymes move through the membrane, driven
by the polymers of cellulose that are synthetized but only where

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microtubules allow them. In this way, by depolymerizing and
polymerizing microtubules again, the cell can control the orientation of
the fibers of cellulose. It has been shown that microtubules
can rearrange in a matter of minutes to accommodate these changes of
orientation. Other extracellular and intracellular factors can also
condition the direction of movement of these enzymatic complexes.

Synthesis and orientation of cellulose fibrils guided by microtubules.

Cellulose synthesizer complexes move as they synthetize cellulose
following the pathways marked by microtubules (Modified from
McFarlane et al., 2014)
The consistency of the cell wall also determines how the cell will
grow. A softening or relaxation of the cell wall must be produced by
secretion of substances to certain areas of the wall. It has been
observed that it becomes softer in certain places through the chemical
modification of the pectins and by acidification; it is in these places
where there is also less resistance and, therefore, where the cell grows.
Pectins play an important role in the relaxation of the cell wall for
growth. They can be hydrated a lot by adding plasticity to the wall. In
particular, during growth, enzymes are released that change the
molecular form of the pectins initially released or different types of
pectins are released directly. All this leads to a relaxation of the cell
wall and promotes cell growth. Calcium is important for pectins since
it favors the union between them, and is released after the elongation
of the cell. For example, the walls of meristematic cells are poor in
calcium.

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Auxins, a class of plant hormones (or plant growth regulators),
cause acidification of the cell wall and its relaxation by activating the
expansins, the methyl-esterase of pectin and the endoglucanases.
The expansins do not have enzymatic activity and their effect seems to
hinder hydrogen bonds, while the endoglucanases decrease the number
of bonds between cellulose-cellulose and cellulose-hemicellulose.
Although microtubules seem involved in determining how a cell
grows, it has been found that non-uniform (anisotropic) growth begins
to be detected even before cellulose fiber orientation occurs. Prior to
the organization of the microtubules and the orientation of the cellulose
fibers, a local or irregular alteration of the pectins takes place.
Therefore, the initiation of heterotropic growth would not begin with
the reorientation of the microtubules, but with the modification of the
pectins. It is even suggested that the orientation of the microtubules
would be a consequence of the alteration of the pectins and, therefore,
a secondary response.

SECONDARY CELL WALL

Those cells that have the mission of support and those conductors
that are part of the xylem develop an additional wall layer called
secondary cell wall. It is deposited between the plasma membrane and
the primary cell wall and the process involves the synthesis of
numerous layers of cellulose fibers that are added one after the other
by a mechanism called apposition. The synthesis of the secondary cell
wall begins during the final phase of the growth and extension of the
primary cell wall. Once the cell wall is synthetized, the cells die
by apoptosis. They are probably one of the few cell types whose
function, mechanical resistance and sap transport, is performed when
they die. Plants that grow in thickness and height need a great support
and develop what we call wood. The secondary wall is the main
component of the wood.

Composition
The secondary cell wall is composed mainly of cellulose (40-60%
of the dry weight), hemicellulose (10-40% of the dry weight, especially
the xylan) and lignin (10 to the 35% of the dry weight). It has very few
pectins and lacks glycoproteins as structural proteins and enzymes, or
at least they are not abundant. The proportion of cellulose in the
secondary wall is greater that in the primary wall and it also has
hemicellulose in lower proportion.
A typical substance of the secondary cell wall is lignin, which is
the most abundant biopolymer in plants after cellulose. The lignin is
deposited between the microfibrills of cellulose to give consistency.

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This molecule allowed the plants gain a consistency and resistance
hitherto unknown. When lignification of the secondary wall occurs,
part of these molecules can also be deposited in the primary cell wall
and in the middle lamella. Even in conifers, the largest amount of
lignin is found in the middle lamella of the conductive cells. The cross-
linked framework that these molecules form seems to favor the
elimination of water from the wall and, therefore, the access of
hydrolytic enzymes.

Orientation of the fibers of cellulose in the

different layers of the cell wall.

The secondary wall have a thickness of 2 to 10 µm, is poorly

hydrated and is rigid. Its thickness is greater than that of the primary
wall, sometimes so thick that it obliterates the interior of the cell. In
some cells, three layers can be distinguished in the secondary cell
wall: S1, S2, and S3, each with a different orientation of its fibers of
cellulose. Usually layer S2 varies in thickness between cell types.
Cellulose fibers are usually arranged helically. When the secondary
wall develops, the rest of the cytoplasm adhere the layer S3 forming a
layer called the warty layer, due to the irregularity of its surface. The
secondary cell wall deposition is not very regular so there
are interruptions in it. Sometimes, it is possible to find cells with a part
of their wall that is secondary and another part that is primary.

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Diagram showing various forms of thickening in the cell walls of the

conductive cells of the xylem.

The irregular deposition of secondary cell wall on the cell surface

creates irregular structures that are characteristic of some cell types.
For example, the cells of the protoxylem and of the metaxylem present
a thickening of the secondary wall that is arranged helically along the
cell. This arrangement resembles the tracheae of the insects and thus
the conductive cells of the xylem are called tracheal elements. While in
the secondary xylem, cells there have other types of irregularities.

Pits
Although the entire cell may be surrounded by a secondary cell
wall, there are always interruptions or channels in it that are called
pores or pits, which originate when the secondary cell wall is being
deposited. These pits are created simultaneously in the two
neighboring cells, leaving a channel that allows to communicate the
interior of both cells. A thin membrane, which is called membrane of
the pit, separates the two aligned pits of neighboring cells; this
membrane is formed by the middle lamella and the primary walls of
the two cells. The pits, one or more, are formed where there were
primary pits in the primary cell wall.

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Types of pits
Though pits are usually simple and complementary, a few more pit
variations can be formed:
 Simple pits: A pit pair in which the diameter of the pit chamber and
the diameter of the pit aperture are equal.
 Bordered pits: A pit pair in which the pit chamber is over-arched by
the cell wall, creating a larger pit chamber and smaller pit aperture.
 Half bordered pits: A pit pair in which a bordered pit has a
complementary simple pit. Such a pit pair is called half bordered pit
pair.
 Blind pits: A pit pair in which a simple pit has no complementary pit.
 Compound pits: A pit pair in which one cell wall has a large pit and
the adjacent cell wall has numerous, small pits.

Main morphologies of pits.

The pits have different morphology. In the simple pits, the pit
chamber or hollow area that crosses the cell wall has a similar diameter
over its entire length or a little wider in the pit apertures. These pits are
found in parenchyma cells, extraxilar fibers and sclereids. The
bordered pits are those in which a ridge is formed in the apertures, half
bordered pits when only an aperture shows the ridge (typically
established between conductive and accompanying cells)
and bordered with “torus” are those in which in the middle lamella
there is a thickening of the primary wall called a bull, which acts as a
valve. The bordered pits with “torus” are absent in flowering plants
(angiosperms). On the other hand, there are pits called blind when the
pit opens to the intercellular space; that is, it has no complementary pit.

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CELL WALL FORMATION DURING CELL DIVISION
A new plant cell does not generate its entire cell wall completely
from scratch, that is, it is not born naked. When a cell is going to
divide, its two daughters cells inherit all the cell wall that produced its
progenitor except in the area where the cytoplasms will separate. The
formation of this cell wall from scratch begins at the late anaphase of
the mitosis, beginning with this the cytokinesis or division of the
cytoplasms. The first thing observed during the cytokinesis of plant
cells is the transport of vesicles from the Golgi apparatus with content
to build the new wall, mainly polysaccharides and glycoproteins. This
transport is given from the two zones close to the nuclei to the
intermediate zone where the new wall will be formed. The vesicles are
transported by motor proteins along bundles of microtubules remaining
from the mitotic spindle. There is a bundle for each nucleus and both
bundles overlap in the intermediate zone. In this division zone, there
are also actin filaments oriented perpendicularly to the microtubules.
The set of microtubules, actin filaments, and vesicles is
called phragmoplast. The phragmoplast is the structure that is
responsible for forming the new cell wall. When the vesicles reach the
intermediate zone of division, where the new cell wall will form, they
fuse together to form a plate-like structure that will separate the two
cells and that is oriented perpendicular to the mitotic spindle. This
plate is called a cell plate. The cell plate grows in a centrifugal manner,
that is, the center of the plate is formed first and then more material is
added to the edges so that it grows in extension, but not in thickness.
The phragmoplast then adopts, by depolymerization and
polymerization of new microtubules, an annular form and the vesicles
are added in the periphery of the cell plate.

Different phases of mitosis from prophase (left) to telophase (right). It

can be observed how a new cell wall is gradually created that separates
both groups of chromosomes, which will form the nuclei of the daughter
cells. This structure under construction is called phragmoplast (arrow). It

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is also seen in this figure.

Formation of the new cell wall thanks to the activity of the phragmoplast,
which is formed by microtubules belonging to the two daughter cells;
they transport vesicles from the Golgi apparatus to the central plate. The
phragmoplast becomes increasingly peripheral until it touches the original
wall of the stem cell and the central plate merges with that wall.
The cell plate edges will be extended, making the cell plate grow,
until they come into contact and merge with the cell walls parallel to
the mitotic spindle that the mother cell already had. In this way, each

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daughter cell is completely surrounded by a cell wall. The cell plate,
with the arrival of more substances from the Golgi apparatus,
especially pectic substances, will be transformed into the middle
lamella. It seems that once the contact of the edge of the cell plate with
the middle lamella of the mother cell has occurred, it is when there is a
transformation of the cell plate in the middle lamella. The growth of
the middle lamella is mainly centripetal, that is, it will form from the
edges to the interior, where the cell plate began to form. The middle
lamella is the layer of the cell wall that is shared between the two
daughter cells and both contribute to its formation. It is a very fine
layer to which later the others will be added to form a mature cell wall.
Regardless of whether the cell wall is synthetized again or new layers
are added during maturation, the process is always from the outside in,
that is, the more recent parts are always closer to the plasma
membrane.
An interesting aspect of the formation of a new cell wall during
cytokinesis is where and how the division plane will be oriented. For
example, if it will be periclinal or anticlinal, or any other orientation.
The position and orientation of the dividing plane, and therefore of the
cell plate, is established even before the formation of the mitotic
spindle. In the majority of the cells, before the formation of the mitotic
spindle, a network of microtubules, actin filaments and endoplasmic
reticulum cisterns appear in the cortical region of the cytoplasm near
the plasma membrane, forming a belt or ring called the preprophase
band. This ring disappears when the mitotic spindle begins to form.
However, the cell plate will be formed where this preprophase band
was located. In such a way that this initial band conditions the
formation and orientation of the mitotic spindle.
During cytokinesis, intercellular spaces are also formed; they are
important for the diffusion of gases and store secretion substances.
Most of these spaces are formed when the growth edge of the new
middle lamella reaches to the proximities of the primary cell wall of
the mother cell. This growing middle lamella branches out and then
two growth fronts are created that will cross the primary cell wall of
the mother cell. When these fronts reach the middle lamella of the
mother cell, a space surrounded by middle lamella is created. This
space will become an intercellular space. During the maturation of
tissues, the intercellular spaces increase, being greater in adult tissues.
The normal form is by schizogenesis, that is, by a physical separation
between cells produced first by degradation of the middle lamella that
allows physical separation, and by a subsequent cell differential
growth. These spaces are very evident in the spongy parenchyma of the
leaves.

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During the formation of the cell wall, cisterns of endoplasmic
reticulum are trapped; these cisterns inhibit cell wall formation and
remain as discontinuities, which will later become plasmodesmata.
Torus and margo

A simplified diagram of a bordered pit-pair with a torus and margo. The

top shows an unobstructed pit and the bottom shows an aspirated pit, with
the margo flexing under stress.
The torus and margo are characteristic features of bordered pit-
pairs in gymnosperms. In other vascular plants, the torus is rare. The pit
membrane is separated into two parts: a thick impermeable torus at the
center of the pit membrane, and the permeable margo surrounding it. The
torus regulates the functions of the bordered pit, and the margo is a cell
wall-derived porous membrane that supports the torus. The margo is
composed of bundles of microfibrils that radiate from the torus.
The margo is flexible and can move towards either side of the pit
while under stress. This allows the thick, impermeable torus to block the
pit aperture. When the torus is displaced so that it blocks the pit aperture,
the pit is said to be aspirated.
Plasmodesmata
Plasmodesmata are thin sections of the endoplasmic reticulum that
traverse pits and connect adjacent cells. These sections provide an avenue
of transport through the pits and facilitate communication.
Plasmodesmata are not restricted to pits however, as plasmodesmata often
cross a cell wall of constant width and occasionally the cell wall is even
wider in areas where plasmodesmata traverse it.
REFERENCES
Evert RF. 2006. Esau's plant anatomy: meristems, cells, and tissues of the plant body: their
structure, function, and development, 3rd edition. John Wiley and Sons, Inc. ISBN:
9780471738435.
Hartholt J, Suttangkakul A, Scheller HV 2010. Biosynthesis of pectin. Plant physiology.
153: 384-395.
McFarlane HE, Döring A, Persson S. 2014. The cell biology of the cellulose synthesis.
Annual review of plant biology. 65:69-94.
Charles B. Beck (2010). An Introduction to Plant Structure and Development: Plant Anatomy
for the Twenty-First Century (second, revised ed.). Cambridge University Press.
p. 31.ISBN 9781139486361.

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Advance Plant anatomy / Master student Plant tissues/ Meristematic tissues

PLANT TISSUES
Introduction
Growth takes place in two stages in plants: first there is the division of
cells of an undifferentiated type (simple, thin-walled parenchyma) adding to
the number of cells; then there is the enlargement of some of the cells
produced by these divisions.
Dividing cells of the undifferentiated type are not present throughout
the plant, but are concentrated in particular places. In addition to these, certain
cells in most organs remain relatively undifferentiated and may begin to
divide if the appropriate conditions arise and after they have undergone a
process known as dedifferentiation. Such cells give rise to adventitious roots
and buds, or to the callus tissue which forms during wound healing. They are
of great importance to the horticulturalist. The ability of such cells to divide is
a basic requirement for the success of many forms of vegetative propagation
and grafting.
Cells that divide actively to produce the primary plant body are associated
together in meristems.
From the above it is clear that the plant tissues :
A tissue is a group of cells having a common origin and usually
performing a common function. A plant is made up of different kinds of
tissues. Tissues are classified into two main groups, namely, meristematic and
permanent tissues based on whether the cells being formed are capable of
dividing or not.

MERISTEMATIC TISSUES (MERISTEMS)

Growth in plants is largely restricted to specialized regions of active cell
division called meristems (Gk. meristos: divided). Plants have different kinds
of meristems. The meristems which occur at the tips of roots and shoots and
produce primary tissues are called apical meristems.

Characterization
 Meristematic tissue is a group of cells that has power of continuous
division.
 Cells are immature and young
 Meristematic tissue is commonly called as meristems.
 Shape of cell: each cell is oval, rounded, polygonal or rectangular
 Size: small
 Intercellular space: Absent
 Cell wall: thin walled made up of cellulose
 Nucleus: single large and prominent
 Reserved food: cell do not store food
 Cell division: high capacity and continuous
 Metabolic activity: very high

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TYPES OF MERISTEMATIC TISSUE

1. Based on origin
2. Based on position
3. Based on function

Types of meristematic tissue on the basis of origin

i- promeristem (primodial meristem)

 Origin: embryonic origin

 It is earliest and youngest meristematic tissue
 It is present in growing root and shoot tip.
 It give rises to primary meristem,
ii- Primary meristem:
 Origin: from Promeristem
 Cell are always active and dividing
 Present below promeristem in the shoot and root tip, and also in
intercalary position
 It give rises to secondary meristem, and primary permanent tissue.
iii- Secondary meristem:
 Origin: from primary meristem
 It is developed later on life
 It give rises to secondary permanent tissue

Types of meristematic tissue on the basis of Position

i. Apical meristem
ii. Intercalary meristem
iii. Lateral meristem

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i- Apical Meristems
Apical meristems are located at the shoot apex, where primary stem,
leaves and flowers differentiate, and at the root apex (Fig. below), where
primary root tissue is produced. Subsequent elongation of the shoot axis may
occur by random cell divisions and growth throughout the youngest
internodes. This region of diffuse cell division is termed an uninterrupted
meristem, and is continuous with the apical meristem. However, in some plant
stems, particularly in grasses, most cell divisions contributing to stem
elongation occur in a limited region, usually at the base of the internode,
which is then termed an intercalary meristem. Both intercalary and
uninterrupted meristems represent growth in regions of already differentiated
tissues.

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ii- Lateral Meristems

Lateral meristems are located parallel to the long axis of a shoot or root,
most commonly in the pericyclic region, at the junction between vascular
tissue and cortex. Examples of lateral meristems include primary and
secondary thickening meristems (PTM and STM) and vascular cambium.
Primary and secondary thickening meristems produce both ground tissue and
vascular bundles . Vascular cambium produces secondary xylem and phloem.
Adventitious roots are typically formed in the root pericycle; in these cases the
pericycle could be termed a lateral meristem. The phellogen (cork cambium)
is a lateral meristem that occurs in the stem or root cortex, where it forms a
protective corky layer. A phellogen may also develop in the region of a
wound, or at the point of leaf abscission.

ii- Intercalary meristem

A meristem developing between regions of mature or
permanent tissue (as at the base of the grass leaf) compare apical meristem,
lateral meristem.
Intercalary meristems are capable of cell division, and they allow for
rapid growth and regrowth of many monocots. Intercalary meristems at the
nodes of bamboo allow for rapid stem elongation, while those at the base of
most grass leaf blades allow damaged leaves to rapidly regrow.
Meristems are classified by their location in the plant as apical
(located at root and shoot tips), lateral (in the vascular and cork cambia),
and intercalary (at internodes, or stem regions between the places at which

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leaves attach, and leaf bases, especially of certain monocotyledons—e.g.,

grasses).
Monocots, like grasses, have intercalary meristems which allow the
leaves to grow back after mowing. In addition to the apical meristems located
in the shoot and root tips, plants in the DICOT class have lateral meristems.
Intercalary Meristems – The intercalary meristems are located at the
internodes or the base of the leaves. The intercalary meristems help in
increasing the length of the internode. This is usually seen in
monocotyledonous plants.
Intercalary Growth. a lengthwise growth in plants as a result of cell
division in the formative tissue (meristem), located below the top of the
organ—for example, in the internodes of the stalks of grasses and at the base
of the leaves.

Types of meristematic tissue on the basis of function:

i. Protoderm
 Function: protection from mechanical injury
 It gives rise to epidermis layer.
 It is the outermost meristematic tissue
ii. Procambium
 Function: transport of water and nutrition
 It gives rise to vascular tissue (xylem and phloem)
 It is the innermost meristematic tissue
iii. Ground meristem
 Function: various functions
 It gives rise to cortex, endodermis, pericycle and pith in dicot and
hypodermis, ground tissue in monocot.

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MERISTEMOIDS AND ASYMMETRIC CELL DIVISION

Meristemoids are individual cells that are responsible for the
differentiation of distinct structures. In many cases meristemoids represent the
smaller, densely cytoplasmic, daughter cell that results from an unequal
(asymmetric) cell division; the larger daughter cell is less active. Asymmetric
divisions are caused by cell polarization resulting from organized arrays of
actin filaments in the dense cytoplasm during determination of cell plate
alignment42. Examples of unequal cell divisions include cleavage of the
microspore into a larger vegetative cell and smaller generative cell, formation
of a root hair initial (trichoblast), a protophloem division to form a larger sieve
tube element and smaller companion cell, and division of an epidermal cell
into two cells of unequal sizes, the smaller of which is the meristemoid that
will divide to form the guard cells of a stoma.

Several theories have been to explain the mode of growth found in shoot
apical meristem
1- APICAL CELL THEORY
This theory proposed by Nagli 1878. According this theory, a single
apical cell is the structural and functional unit apical meristem which governs
the entire process of apical growth. Such organization has been found only in
lower plants like algae, ferns.

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2- HISTOGEN THEORY
Hanstein postulated this theory revealing that three distinct
meristematic zone (layer) are found in apical region. Each zone consists of a
variable number of layers called histogen or tissue builder.
i- Dermatogen
The outermost uniseriate layer. Dermatogen cells divide anticlinally and
develop into unilayerd epidermis.
ii- Periblem
The middle region composed of isodiametric cell, forms the cortex.
iii- Plerome
The central or inner mass, forms the stele (vascular tissues, pericycle,
pith, rays). Disadvantage of this theory:
It is not possible to distinguish these three histogen layers in gymnosperms
and angiosperms. Hence this theory was rejected for shoot apical meristem.

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3- TUNICA-CORPUS THEORY
Schmidt 1924 proposed tunica – corpus theory to explain shoot apex
organization. According to this theory, there are two zone in apical meristem.
i- Tunica: the outer zone consisting of one or more peripheral layers of cells,
forming
the outer region by anticlinal divisions.
ii- Corpus: the central undifferentiated multilayered mass of cells surrounded
by tunica which forms the central part of shoot by irregular divisions.
The tunica cells are smaller and corpus cells are larger. The number of tunica
layers may vary even in same species due to influence of seasonal growth
changes.

4- CYTOHISTOLOGICAL ZONATION
It was found that in gymnosperms, tunica like layer is not found which led
foster 1939 to divide shoot apex organization of ginkgo biloba on basis of
cytohistological zonation, which react differently to staining. He recognized four
inter related zones.
i- Apical initial zone
The outermost layer of the apical meristem undergoes periclinal and
anticlinal divisions and contributes cells to the peripheral and interior tissues of the
shoot.
ii- Peripheral layer
Encircles the other zones. The peripheral zone typically is the most
meristematic of all three zones and has the densest protoplasts and the smallest cell
dimensions. It may be described as a eumeristem. Leaf primordia and the
procambium arise here, as well as the cortical ground Tissue.
iii- Central mother cell
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iv- Rib meristem

Appears directly below the central zone and is centrally located in the apex.
It usually becomes the pith after additional meristematic activity has occurred.

REFERENCES
1- CUTLER, D.F. ; BOTHA, C.E.J. and STEVENSON, D.W. (2007). Plant Anatomy An
Applied Approach, First published by Blackwell Publishing Ltd BLACKWELL
PUBLISHING , UK.
2- PAULA, J. and RUDAL, L. (2007). Anatomy of Flowering Plants An Introduction to
Structure and Development, The Edinburgh Building, Cambridge CB2 8RU, UK.

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2. Permanent tissues
Tissues that stopped active division & differentiate to functions. It may be
living cells contain cytoplasm & nucleus, keep in division ability depending on
conditions, some of the permanent tissues lose it's nucleus but keep cytoplasm and
still active like sieve tube, some of the permanent tissues lose activity because of
thickening with Lignin, suberin & die as like as fibers, tracheid.

Permanent tissues on the base of Topographic continuity

Tissues forming plant body classified into systems which located certain
parts in plant body, therefore three types of systems forming plants body

1- Dermal tissue system

All tissues which covered plant body such as Epidermis in primary growth
parts and Periderm in roots and stems with secondary growth
2- Vascular tissue system
Consist of xylem and phloem in primary and secondary growth of plant
body.

3- Ground tissue system (Fundamental)

Consist of all tissues between two systems above such as Cortex, pith,
Medullary rays in roots and stems and Ground tissue in stems of monocot. and
Mesophyll in leaves , Paranchyma, Collanchyma and Sclerenchyma are the most
important forming of this system.

1- Dermal tissue system

This term refers to all peripheral tissues envelop all plant body organs in
primary and secondary growth, dermal tissue is a protective layer. Separate plant
from the outer environment & mechanical protection, gaseous exchange through
stomata & keeping water & metabolic products, it is represent in:
1- Epidermis: primary growth.
2- Periderm: secondary growth.
1- Epidermis
The term epidermis designates
the outermost layer of all parts of on
the primary plant body. It is derived
from the (Greek epi, for upon, and
derma, for skin). Including roots,
stems, leaves, flowers, fruits, and
seeds. An epidermis is considered to
be absent, however, on the root cap
and not differentiated as such on
the apical meristems.

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EPIDERMIS CELLS CHARACTERISTICS
1. Leaving cells, clear nucleus contain cytoplasm, vacuoles, chloroplasts are
absent inspect of guard cells, hydrophytes, shadow plants.
2. Primary cell wall and some time secondary cell walls like in Pinus. containing
primary pit-fields and plasmodesmata generally occur in the
anticlinal and inner periclinal walls of the epidermis, although the
frequency of plasmodesmata between the epidermis and mesophyll
of leaves is relatively low. For a time plasmodesmata were thought
to occur in the outer epidermal walls and were called ectodesmata.
3. Intercellular spaces absent so that block water & gases passage inspect of
stomata.
4. In roots the epidermis may have a common origin with cells of the
root cap or differentiate from the outermost cell layer of the cortex.
5. The difference in origin of the epidermis in shoots and roots has
convinced some investigators that the surface layer of the root should
have its own name, rhizodermis, or epiblem.
6. Aerial parts epidermis walls saturated from the outer side with cutien, which is
added in two ways:
A- Cutien adding throw cells wall this way called ( Cutinization )
B- Cutien adding as continues outer layer this way called (Cuticularization)
and the layer called cuticle.
7- Cuticles are formed in cortical cells and give rise to a protective
tissue called cuticular epithelium.
8- The cuticle occur in the aerial part and absent in the ground part, and cuticle
thickness is very important from the ecological part,
Xerophytes plant = thick cuticle
Hydrophytes plant = thin cuticle or absent
9- Epidermal cell walls vary in thickness in different plants and in different
parts of the same plant. In the thinner walled epidermis, the outer
periclinal walls are frequently thicker than the inner periclinal and
anticlinal walls.

INITIATION OF EPIDERMIS
Plant epidermal initials varies with plant groups as following :
1- In low vascular plants there is no independent origin of the epidermis, but only
one or a few cells forming of all the tissues of the plant.
2- In high vascular plants the method of epidermal initials depends on how
regular structural cells in the developing apex.
a- most of gymnosperm and few of angiosperm there is no independent origin of
the epidermis thus the term Protoderm is called on surface layer of the apex
that will later become the epidermis.
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b- In some of dicot. and most of monocot. plants which shoot apex on it
Characterized into clearly coated layers, epidermis forming from outer coated
layer as a result of vertical divisions (anticlinal) epidermis has an independent
initials called Dermatogen as in Hestorgen theory.
3- In roots, epidermis rarely has an independent initials layer, in some of root
apical meristem on monocot. plants there are four initials zones one of them
specialized to forming epidermis

TYPES OF Epidermis
1- Simple epidermis (uniseriate)
The epidermis is usually one layer of cells in thickness

2- Double epidermis (double seriate)

3- Multiple epidermis (multiseriate)
In some leaves the protodermal cells and their derivatives divide
periclinally (parallel with the surface), resulting in a tissue consisting of
several layers of ontogenetically related cells. (Sometimes only
individual cells of the epidermis undergo periclinal divisions.) such a
tissue is referred to as a multiple, or multiseriate epidermis.

FUNCTIONS OF THE EPIDERMIS

The common functions of the epidermis of the aerial plant parts
are :
1- considered to be reduction of water loss by transpiration.
2- mechanical protection.

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3- gaseous exchange through stomata. Because of the compact
arrangement of the cells and the presence of the relatively tough
cuticle.
4- The epidermis also offers mechanical support and adds stiffness to
the stems
5- In stems and coleoptiles the epidermis, which is under tension, has
been regarded as the tissue that controls elongation of the entire
organ.
6- The epidermis is also a dynamic storage compartment of various
metabolic products
7- The site of light perception involved in circadian leaf movements
and photoperiodic induction.
8- In the sea grasses and other submerged aquatic angiosperms, the
epidermis is the principal site of photosynthesis.
9- In some leaves the upper epidermal cells act as lenses, focusing light
upon the chloroplasts of the underlying palisade parenchyma cells
10- Epidermal cells of both the shoot and root are involved with the
absorption of water and solutes.
11- Although the mature epidermis is generally passive with regard to
meristematic activity, it often retains the potentiality for growth for a
long time.
12- Epidermal cells have living protoplasts and may store various
products of metabolism.
13- They contain plastids that usually develop only few grana and are,
therefore, deficient in chlorophyll. Photosynthetically active
chloroplasts, however, occur in the epidermis of plants living in
deep shade, as well as in the epidermis of submerged water plants.
Starch and protein crystals may be present in epidermal plastids,
anthocyanins in vacuoles.

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TYPES OF EPIDERMIS CELLS

1- Ordinary epidermal cells : (also called ground cells, epidermal cells
proper, unspecialized epidermal cells, pavement cells), and of more
specialized cells dispersed throughout the mass.
2- guard cells : more specialized cells are the guard cells of the stomata
and a variety of appendages.
3- epidermal hairs or trichomes : the trichomes, including the root hairs,
which develop from epidermal cells of the roots.

1- Ordinary epidermal cells

Mature ordinary epidermal cells (often simply referred to
hereafter as epidermal cells) are variable in shape, but typically they are
tubular, having little depth. Some, such as the palisade -like epidermal
cells of many seeds, are much deeper than they are wide. In elongated
plant parts, such as stems, petioles, vein ribs of leaves, and leaves of
most monocots, the epidermal cells are elongated parallel with the long
axis of the plant part. In many leaves, ovaries, and ovules, the
epidermal cells have wavy vertical (anticlinal) walls. The pattern of

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waviness is controlled by local wall differentiation, which determines
the pattern of wall expansion.

2- Guard cells
Cells occur like couples with a stoma between each pear leaving cells
contain nucleus & chloroplasts its occur either kidney shape or dumble shape may
be contact with assistant cells or may not, its function is open and close the
stomata.
Stomata
Stomata are specialized pores in the epidermis through which gaseous
exchange (water release and carbon dioxide uptake) takes place. They occur on
most plant surfaces above ground, especially on green photosynthetic stems and
leaves, but also on floral parts. Each stoma consists of two guard cells
surrounding a central pore. Cuticular ridges extend over or under the pore from
the outer or inner edges of the adjacent guard cell walls.

Stomata kinds:

1- Monocot – Dicot type of stomata

The normal and most common type is called the type of dicotyledon and the
dicotyledon. It is found in all dicotyledons and all monocots except for Gramineae
and cyperaceae family and in this type the sense cells are kidney-shaped in the
surface view. In the vertical section, the guard cells appear to contain a horn-like
on the outside only or the outer and inner sides of the stoma, these parts are
bounded on the outside by a cavity called the front cavity, and on the inner side
another cavity called the back cavity, and the latter is located between the stoma
opening and the internal air chamber that lies directly inside the stoma.

2- Gramineae - cyperaceae type

In this type, the s guard cells are dum-bell-shaped in surface view and the
cell appears narrow in the middle and widened and bulging at the two ends. The
narrow middle part is thick in wall and the swollen ends are thin wall. Opening
and closing the gap in this type depends on the shape of the guard cell and the
uneven thickening in its wall. When the cell is full, the thin edges below the thick-
walled midsection swell and the stoma opens. In the case of decreased osmotic
pressure, the peripheral parts become less swollen, the thicker, intermediate parts
of the sentinel cells converge, and the stoma closes.

3- Coniferales type
The third type is found in Coniferales plants, including Pinus, and it is
called the gymnosperm type. Stomatal complexes in this type are sunken and
equipped with subsidiary cells. The guard cells appear in the vertical section in an

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inclined position, as their walls are thickened in their parts and thin in other parts,
and the opening and closing of the stoma depends on the type of thickening of the
guard cells walls and the placement of the subsidiary cells in relation to the guard
cells

subsidiary cells
The epidermal cells immediately adjacent to the guard cells are termed
subsidiary cells. it they differ morphologically from surrounding epidermal cells.

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Type of subsidiary cells
Classifications of stomatal
types are based either on the
arrangement of mature subsidiary
cells, or on their patterns of
development. Types of mature
stomata include :
1- Anomocytic : stomata lack
subsidiary cells entirely Ex.
Apocynaceae, Boraginaceae,
Chenopodiaceae, Cucurbita
etc.
2- Anisocytic : stomata possess
three unequal subsidiary cells
Ex. Cruciferae, Solanum,
Petunia, Sedum and
Nicotiana etc.
3- Diacytic : stomata possess one
or more pairs of subsidiary
cells with their common walls
at right angles to the guard
cells Ex. , Acanthaceae,
Hygrophila, Dianthus etc.
4- Paracytic : stomata possess
one or more subsidiary cells
at either side of the guard
cells Ex. Rubiaceae,
Convolvulaceae, Phaseolus,
Arachis and Psoralea etc.
5- Actinocytic: stoma remains
surrounded by a circle of
radiating subsidiary cells Ex.
Ancistrocladus and Euclea
pseudebenus (Ebenaceae)
6- Gramineous: Gramineous
stoma possesses two guard
cells that are shaped like
dumb-bells. Each guard cell
has a narrow middle portion
and two bulbous ends. The
narrow middle portion is
strongly thickened.

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The subsidiary cells occur parallel to the long axis of pore. Ex.
Gramineae and Cyperaceae.
7- Hemiparacytic: The stoma is accompanied by a single subsidiary cell,
which is placed parallel to the long axis of the pore and this cell may
be long or short in length in contrast to the guard cells. Example:
Glinus latioides and Trianthema lancastrum etc.
8- Hexacytic: The stoma is surrounded by six subsidiary cells among
which two are situated on the two polar sides and rest two pairs occur
on the two lateral sides being parallel to the long axis of the guard
cells.

Stomata occurrence
The stomata are common on green aerial parts of plants
particularly the leaves, they occur either on both surfaces (amphistomatic
leaf) or on one only either the upper (epistomatic leaf) or more commonly
the lower (hypostomatic leaf).

Formation of Stomata
Stomata formation either by the oblique division for the
protodermal cell (in dicotyledon) or by unisometric division for
epidermal cell (in monocotyledon) temperature have a direct effect on
division.

EPIDERMAL HAIRS OR TRICHOMES

Trichomes (from the Greek, meaning a growth of hair) are
highly variable epidermal appendages. They may occur on all
parts of the plant and may persist throughout the life of the plant
part or may fall off early. Some of the persisting trichomes
remain alive; others die and become dry. Although trichomes
vary widely in structure within families and smaller groups of
plants, they are sometimes remarkably uniform in a given taxon
and have long been used for taxonomic purposes trichomes are
usually distinguished from emer-gences, such as warts and
prickles, which are formed from both epidermal and
subepidermal tissue and typically are more massive than
trichomes. The distinction between trichomes and emergences is
not sharp, however, because some trichomes are elevated upon a
base consisting of subepidermal cells. Thus a developmental
study may be necessary to determine whether some outgrowths

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are solely epidermal in origin or both epidermal and
subepidermal in origin.
They vary widely in both form and function, and include
unicellular or multicellular, branched or unbranched forms, and also
scales, glandular (secretory) hairs, hooked hairs and stinging hairs.
Glandular trichomes usually possess a unicellular or multi-cellular
stalk and a secretory head with one to several cells. Secreted substances
such as volatile oils collect between the secretory cells and a raised
cuticle, which later breaks to release them. Trichomes can be classified
into:

1- Unicellular:
a. unbranched in cotton seed
b. branched in Mathiola
2- Multicellular:
a. uniseriate
b. multiseriate in begonia

3- Dendroid (tree-like) in platanus

4- Stellate in Malva
5- Peltate in Olea
6- Bladders in Atriplex
7- subsurface emergence in some
roses.
8- Stinging hairs in Urtica

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The Periderm
Periderm is a protective tissue of secondary origin. It replaces the epidermis
when the axis increased in growth and the epidermis is destroyed. Periderm
contains:
a. Cork cambium (phellogen)
b. Cork (phellem) , mostly dead cells with cell
walls containing much suberin.
c. phelloderm
Cork cambium (phellogen): Secondary
meristem, lateral position, rectangular cells,
meristematic cells which have so more vacuoles,
simple tissue (one type of cells, thermoisolation,
to form cork (outside) and secondary cortex
(inside).
The origin:
- In the root from the pericycle.
- In the stem from the hypodermis region.

Cork (phellem): permanent tissue, it cells dead in

mature because of thicking its wall with (suberin)
(max substance) in perviousness to water & gases exchanged & prevent
temperature effect on the inner tissues.

Phelloderm: parenchymatic cells differ from ordinary cortex cells in its regular
strips & its composed of few layers some time one layer.
The average of phellem formation is higher from the average of phelloderm
formation and it may reaches to 20 strips in one season of growth.
Origin:
The first periderm from sub-epidermal layer.

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Rhytidom: a term mean all the dead tissues which occur outside the phellogen
when the phellogen get in deeper layers. (outer Bark) (a layer of cork alternating
with a layer of tissue cut off by the cork).
Bark: a term applied most commonly to all tissues outside the vascular cambium
of the axis, in either primary or secondary state growth. (Like primary and
secondary phloem, cortex, periderm and any other tissue outside the periderm,
Bark contain a sclerenchymatic cells, phloem fibes&sclerides. We can see two
kinds of Bark:
- Cyclic Bark: when the cork cambium occur as a complete cylinder. The Bark
is formed and fall in cyclic form as like as in vitis.
- Scaly Bark: its formed when the cork cambium occur in linear shape and the
Bark fall like as scales as like as in Pinus& Oak.

Lenticels:
Cells without subern in on its walls called complementary tissue which
contain wide intercellular spaces their function is gas use exchange instead of the
guard cells.
Lenticels are structurally
differentiated portions of the periderm
characterized by a relatively loose
arrangement of the cells.

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3- Ground Tissue System (Fundamental)
This system includes all the tissues excepting the epidermis and the
vascular bundles. Thus it is the largest or the most exhaustive system, which
begins from the layer next to epidermis and continues right up to the center of
the organs in cylindrical bodies such as Cortex, pith, Medullary rays in roots
and stems and Mesophyll in leaves , Paranchyma, Collanchyma and
Sclerenchyma are the most important forming of this system.

Characterization
1- Ground tissue system is permanent tissue which is heterogeneous in nature,
consisting of diverse types of cell elements adapted to carry on different
functions.
2- In the axis of higher plants the vascular bundles occupy a restricted position
inside the stele or central cylinder.
3- The ground tissues occurring outside the stele, and, in fact, surrounding it,
form the cortex, what may be called external or extrastelar ground tissue.
4- Similarly there are internal or intrastelar ground tissues inside the stele, e.g.,
pith.
5- Both external and internal ground tissues are further differentiated to
specialized zones.
6- These tissues are derived from the ground meristem of the embryonic region.

Types of ground tissues

A- Parenchyma
B- Collenchyma
C- Sclerenchyma

A- Parenchyma
The term parenchyma refers to a tissue composed of living cells variable
in their morphology and physiology, the word parenchyma is derived from the
Greek para, beside, and en-chein, to pour, a combination of words that
expresses the ancient concept of parenchyma as a semiliquid substance “poured
beside” other tissues that are formed earlier and are more solid.

Characterization of parenchyma cells

1- Parenchyma may be considered the phylogenetic precursor of all other
tissues.
2- Living cells at maturity with central nucleus, cytoplasm and vacuoles
surrounded by thin primary cell wall (0.2 to 2 μm in diameter), sometimes
is thick in the endosperm of date palm and persimmon.
3-These cells keep their ability to divide or to resume (meristematic
activity)developmentally, parenchyma cells are relatively undifferentiat

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(unspecialized morphologically and physiologically) compared with such
cells as sieve elements, tracheids, and fibers because, in contrast to these
three examples of cell categories, on other word (parenchyma cells are
able to resume meristematic activity: to dedifferentiate, divide, and
redifferentiate). parenchyma cells may change functions or combine
several different ones. However, parenchyma cells may also be distinctly
specialized, for example, with reference to photosynthesis, storage of
specific substances.
4- they are surrounded by intercellular spaces or larger cavities for effective gas
exchange.
5-Shapes of cells differ with the differentiation of tissue.
i. Isodiametric: Helianthus annus ii- Spherical: Geranium

iii- Columinal: like in mesophyll in leaf iv- Stellate: like in Alkana

v- Lobed: like in Pinus

6- Middle lamellae may or not be distinguishable, plasmodesmata are

common.
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7- Parenchyma cells are the pith and cortex of stems and roots, the
photosynthetic tissue (mesophyll) of leaves .
8- Some parenchyma cells develop secondary walls (lignified) make it
difficult to distinguish from sclerenchyma.

Origin
a- The parenchyma of the cortex and pith, of the mesophyll of leaves, and
of flower parts, differentiates from the ground meristem.
b- The parenchyma cells associated with the primary and secondary
vascular tissues are formed by the procambium and the vascular
cambium respectively.

Classification of parenchyma tissues according to function

1- Ordinary parenchyma : not differentiated to any function.

2- Chlorenchyma:

It is specialized parenchyma
tissue specialized for photosynthesis
contains numerous chloroplasts. The
greatest expression of chlorenchyma
is represented by the mesophyll of
leaves, but chloroplasts may be
abundant also in the cortex of a stem.
Chloroplasts may occur in deeper
stem tissues, including the secondary
xylem and even the pith. Typically
photosynthesizing cells are
conspicuously vacuolated, and the
tissue is permeated by an extensive
system of intercellular spaces.

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3- Storage parenchyma
It is characterized by accumulating
specific kinds of substances like starch-
storing cells such as those of the potato
tuber, the endosperm of cereals, and the
cotyledons of many embryos, the
abundant starch-containing amyloplasts
may virtually obscure all other
cytoplasmic components. In many seeds
the storage parenchyma cells are
characterized by an abundance of protein
and/or oil bodies. In various parts of the
plant, parenchyma cells may become
conspicuous by accumulating
anthocyanins or tannins in their vacuoles.

4- Aernchyma
It is type of parenchyma
which found in the leaves, stems
and roots of some plants like
aquatic, the cells are
characterized by small size, thin
cell wall and air filled large
cavities which provide low
resistance internal pathway for
the exchange of gases between
plant organs above the water and
submerged tissues.

5- Water storage parenchyma

Parenchyma cells in the tissue system, parenchyma may be rather
specialized as a waterstorage tissue. Many succulent plants, such as the
Cactaceae, Aloe, Agave, contain in their photosynthetic organs
chlorophyll free parenchyma cells full of water. This water tissue consists
of living cells of particularly large size and usually with thin walls. The
cells are often in rows Transfer cell, a specialized form of parenchyma
cell, is readily identified by elaborate ingrowths of the primary cell wall.
The increase in the area of the plasma membrane beneath these walls

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facilitates the rapid transport of solutes to and from cells of the vascular
system. The ingrowths develop relatively late in cell maturation and are
deposited on the original primary wall; hence they may be considered a
specialized form of secondary wall.

6- Secretory parenchyma
In contrast, secretory types of parenchyma cells have dense
protoplasts, especially rich in ribosomes, and have either numerous Golgi
bodies or a massively developed endoplasmic reticulum, depending on
the type of secretory product formed.

B- Collenchyma tissues
The terms parenchyma and collenchyma are also related, but in the
latter the first part of the word, derived from the Greek colla, glue, refers
to the thick glistening wall characteristic of collenchyma.
Collenchyma cells and parenchyma cells are similar to one another
both physiologically and structurally. Both have complete protoplasts
capable of resuming meristematic activity, and their cell walls are
typically primary and nonlignified. The difference between the two lies
chiefly in the thicker walls of collenchyma cells; in addition the more
highly specialized collenchyma cells are longer than most kinds of
parenchyma cells.

Characterization of collenchyma
1- It is a living simple permanent tissue composed of more or less
elongated cells, with thickened primary walls. for it consists of a
single cell type.
2- Its cells contain nuclei, cytoplasm and chloroplasts.
3- It occurs in the peripheral regions of stem and leaf, directly beneath
epidermis or a few layers removed from it.
4- Frequently forms a continuous layer around circumference of axis
(stem and leaf petiole), may occur in strands
5- In leaf blade occurs in ribs especially the larger one, occurs on both
sides or one side of ribs.
6- Roots rarely have collenchyma.
7- Collenchyma may or may not contain intercellular spaces. If spaces
are present in the angular type of collenchyma, the thickened walls
occur next to the intercellular spaces.
8- Cell Walls of Collenchyma are thick and glistening in fresh sections,
and often the thickening is unevenly distributed. They contain, in
addition to cellulose, large amounts of pectin and hemicelluloses and
no lignin, so it is nonlignified primary walls.

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9- Its function is supporting in living organs

Origin
a- Procambium
b- Ground meristems

TYPES OF COLLENCHYMA ACCORDING TO WALL

THICKENED NATURE

The distribution of wall thickening in collenchyma shows several

patterns, these are :

1- Lamellar Collenchyma
or plate parenchyma
Collenchyma with the
thickenings on two opposite sites
of the tangential walls. And radial
walls still thin, These thicknesses
are in the form of layers or
overlapping sheets lying on top of
each other, cells without
intercellular spaces completely.
Lamellar collenchyma is especially
well developed in the stem cortex
of Sambucus nigra and Helinthus
annuus stem.

2- Angular collenchyma

It is with wall thickenings

localized in the corners
commonly, angular collenchyma
are found in the stems of
Cucurbita. and Vitis. It is the
most common type, cells without
intercellular spaces completely.

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3- Lacunar, or lacunate,
collenchyma
It is characterized by the
presence of intercellular spaces
between cells, and the thickness
is concentrated on the parts of the
walls facing these voids, Cells of
this type may contain
chloroplasts and have the
capacity of dedifferentiation, It
is the least common type. Occur
in Lactuce plant.

C- Sclerenchyma

The word is derived from the Greek skleros, meaning “hard” and
enchyma, an infusion; it emphasizes the hardness of sclerenchyma walls. The
individual cells of sclerenchyma are termed sclerenchyma cells. In addition to
comprising sclerenchyma tissue, sclerenchyma cells like parenchyma cells may
occur singly or in groups in other tissues. both parenchyma cells and
collenchyma cells may become sclerified.

Characterization of Sclerenchyma

1- Simple permanent tissues

2- Thick walled cells, often lignified dead when get mature.
3- Principal function is mechanical (Supporting), so the cells are supposed to
enable plant organs to withstand various stress, such as bending, weight &
pressure, without damage to the thin-walled softer cells.
4- The elastic, highly hydrated primary walls of collenchyma, distinguish this
tissue from sclerenchyma with its hard elastic secondary walls.
5- Sclerenchyma cells are usually divided into two categories, fibers and
sclereids.

Origin
1- Procambium
2- Vascular cambium
3- Differentiation of parenchyma cells.

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TYPES OF SCLERENCHYMA CELLS

1- FIBERS 2- SCLEREIDS

1- FIBERS
Fibers typically are long, spindle-shaped cells, with more or less thick
secondary walls, and they usually occur in strands, the fiber walls are not highly
hydrated. They are therefore harder than collenchyma walls and are elastic
rather than plastic. Fibers serve as supporting elements in plant parts that are no
longer elongating. The degree of lignification varies, and typically the simple or
slightly bordered pits are relatively scarce and slit-like. Many fibers retain their
protoplasts at maturity.
Origin: Procambium or vascular cambium

Distribution of fibers in the plant body

1- Bundle sheath in Gramineae stems such as Zea mays stem.
2- Bundle cap which is part of pericycle in Helianthus annuus stem.

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3- Fibers as continues ring in pericycle of Ruscus sp. and Cucurita .
4- Fibers are separated bundles in Yucca leaf.
5- Fibers as Tapes Opposite of vascular bundles in Zea mays leaf.

1 2 3

4 5

There are two kinds of fibers:

1- Hard fiber: occur in monocotyledone& leaves up and down vascular bundle

and usually its not pure fibers mixed with other tissues, heavily lignified walls.
2- Soft fiber: (Phloem, Pericycle, Cortex fibers) found in dicotyledonae plants,
this kind of fibers utilized economically since ancient times in plant like (Flax,
hemp, ramie, jute). The epidermal hairs of cotton seed and the kapok seed pod
are also termed fibers.

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Fibers may be divided into two large groups depending on its position:
1- Xylary (Xylem fibers or Wood fibers).
2- Extraxylary : fibers located outside the xylem
1- Xylary (Xylem fibers or Wood fibers) : Wood fibers are commonly
divided into two main groups :

a- The libriform fibers

b- The fiber-tracheids
both of which typically have lignified cell walls. The libriform fibers
resemble phloem fibers. Libriform is derived from the Latin liber, meaning
“inner bark,” that is, phloem. Although the distinction between the two groups
of wood fibers has long been based primarily on the presence of simple pits in
libriform fibers and of bordered pits in fiber-tracheids, truly simple pits in fiber
walls are extremely uncommon. The extremes of the two types of wood fiber
are easy to distinguish, but imperceptible gradations occur between them.
Fiber-tracheids also intergrade with tracheids, which have distinctly bordered
pits. Commonly the thickness of the wall increases in the sequence of tracheid,
fiber-tracheid, and libriform fiber. In addition, in a given sample of wood, the
tracheids are usually shorter and the fibers longer, with the libriform fibers
attaining the greatest length.
Although commonly regarded as dead cells at maturity, living protoplasts
are retained in the libriform fibers and fiber-tracheids in many woody plants
(Fibers with living protoplasts have been found in bamboo culms over nine
years of age) These fibers often contain numerous starch grains; hence, in
addition to support, they function in the storage of carbohydrates. The
secondary walls of wood fibers differ from those of phloem fibers in that they
consist of three layers designated S1, S2, and S3 for outer, middle, and inner,
respectively. In addition the walls of wood fibers typically are lignified.
2- Extraxylary : fibers located outside the xylem
i. The cell walls of the extraxylary fibers are often very thick. In the phloem
fibers of flax, the secondary wall may amount to 90% of the cross-sectional area
of the cell.
ii. The secondary walls of extraxylary fibers have a distinct polylamellate
structure, the individual lamellae varying in thickness from 0.1 to 0.2 μm.
iii. Not all extraxylary fibers have such wall structure. In mature bamboo culms,
some fiber walls show a high degree of polylamellation, whereas others have no
clearly visible lamellae.
iv. Some extraxylary fibers have lignified walls; the walls of others contain little
or no lignin (flax, hemp, ramie). Some extraxylary fibers, notably those of the
monocots, are strongly lignified.

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Extraxylary consist of :
a- the phloem fibers :
Phloem fibers occur in many stems. The flax (Linum usitatissimum) stem
has only one band of fibers, several layers in depth, located on the outer
periphery of the vascular cylinder. These fibers originate in the earliest part of
the primary phloem (the protophloem) but mature as fibers after this part of the
phloem ceases to function in conduction. Flax fibers are, therefore, primary
phloem fibers, or protophloem fibers. The stems of, Tilia (basswood), Vitis
(grapevine), and many others, have both primary phloem fibers and secondary
phloem fibers, which are located within the secondary phloem , In addition the
secondary walls of the secondary phloem fibers of most woody angiosperms
and conifers consist of only two layers, a thin outer (S1) layer and a thick inner
(S2) layer .
b- Cortical fibers
Cortical fibers, as the name implies, originate in the cortex, do not
originate as part of the phloem tissue but outside it. Perivascular fibers are
commonly referred to as pericyclic fibers. However, the designation pericyclic
is often used with reference to the primary phloem fibers as well
C- Perivascular fibers
Perivascular fibers are located on the periphery of the vascular cylinder
inside the innermost cortical layer. do not originate as part of the phloem tissue
but outside it.
d- Extraxylary fibers also include the fibers of the monocots, whether or not
associated with the vascular bundles.

2- SCLEREIDS
The distribution of sclereids among other cells is of special interest with
regard to problems of cell differentiation in plants. They may occur in more or
less extensive layers or clusters, but frequently they appear isolated among
other types of cells from which they may differ sharply by their thick walls and
often bizarre shapes.
Sclereids typically are short cells with thick secondary walls, strongly
lignified, and provided with numerous simple pits. Some sclereids have
relatively thin secondary walls, however, and may be difficult to distinguish
from sclerified parenchyma cells. The thick-walled forms, on the other hand,
may strongly contrast with parenchyma cells: their walls may be so massive as
almost to occlude the lumina, and their prominent pits often are ramiform. The
secondary wall typically appears multilayered, reflecting its helicoidal
construction. Crystals are embedded in the secondary walls of certain species.
Many sclereids retain living protoplasts at maturity. Based on Shape and Size.
Sclereids occur in the epidermis, the ground tissue, and the vascular
tissues.

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Origin: Parenchyma cells differentiate into sclereids.
Sclereids May be Classified into a number of types :
1- Brachysclereids (or stone cells) : The most commonly recognized
categories of sclereids, roughly isodiametric or somewhat elongated cells,
widely distributed in cortex, phloem, and pith of stems, and in the flesh of
fruit.
2- Macrosclereids : elongated and columnar (rod-like) cells, exemplified by
sclereids forming the palisade-like epidermal layer of leguminous seed
coats .
3- Osteosclereids ( bone cells) : also columnar but with enlarged ends as in the
subepidermal layer of some seed coats .
4- Astrosclereids, star-cells, with lobes or arms diverging from a central body,
often found in the leaves of eudicots.
5- Trichosclereids : less commonly recognized types, thin-walled sclereids
resembling hairs, with branches projecting into intercellular spaces.
6- Fliform sclereids : long slender cells resembling fibers .

Fig. : Sclereids. A, B, stone cells from fruit flesh of pear (Pyrus). C, D, sclereids from stem cortex of
wax plant (Hoya), in sectional (C) and surface (D) views. E, F, sclereids from petiole of Camellia. G,
columnar sclereid with ramified ends from palisade mesophyll of Hakea. H, I, filiform sclereids from
leaf mesophyll of olive (Olea). J, K, sclereids from endocarp of fruit of apple (Malus). L,
astrosclereid from stem cortex of Trochodendron. (From Esau, 1977.)

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VASCULAR TISSUES SYSTEM

Introduction
Vascular system occur in higher plants therefor it is called vascular
plants and absent in lower plants, therefor it is called non-vascular plants .
The vascular tissues system in plants carries out two essential functions,
namely the delivery of resources (water, essential mineral nutrients, sugars
and amino acids) to the various plant organs, and provision of mechanical
support. There are two types of vascular tissues system :

1- xylem

2- phloem

1- Xylem
The term xylem was introduced by Nägeli (1858) and is derived from
the Greek xylon, wood. The xylem is a complex permanent tissues, it is the
principal water-conducting tissue in a vascular plant. It is also involved in the
transport of solutes, in support, and in food storage. Together with the
phloem, the principal food-conducting tissue, the xylem forms a continuous
vascular system extending throughout the plant body.

The origin of xylem in primary growth stage in procambium, the

resulting xylem elements at first called protoxylem, & the xylem elements
mature after that called metaxylem which large in diameter than the
protoxylem. Plant which get in secondary growth such as dicotyledonous &
gymnosperms have secondary xylem initiated from vascular cambium.

Xylem elements:
In most of Angiosperms xylem consist of vessels, tracheid, Fibers and
xylem parenchyma.

1. The vessels:
A series of vessel members joined end to end, length about 2-15 ft,
thickened with lignin differ in length, thickening way, diameter depending
on spaces, age, time of vessel formation in plant. Because of Functional

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similarities between vessels and tracheid, the two structures together are
called tracheary elements. The secondary thickening of tracheary elements
are pitted , scalariform , reticulate , spiral , annular.

Diagram showing various forms of thickening in the cell walls of the

conductive cells of the xylem.
There are two types of vessels end :
a. Non-perforated: in this kind the ends of the vessels solute.
b. Perforated:
1- Simple perforation plates (uniperforate) : continent one perforate only
2- Compound perforation plates (multiperforation) : continent more than
one perforate in different shapes that are :
- Foraminate or Ephedronal
- Reticulate
- Scalariform

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FIGURE : Perforation plates. Scanning electron micrographs of the perforated end walls of
vessel elements from secondary xylem. A, a simple perforation plate, with its single large
opening, in Pelargonium vessel element. B, the ladder-like bars of a scalariform perforation
plate between vessel elements in Rhododendron. C, foraminate perforation plate, with its
circular perforations, in Ephedra. D, contiguous scalariform and reticulate perforation plates
in Knema fur-furacea. (A–C, courtesy of P. Dayanandan; D, from Ohtani et al., 1992.)

2- The Tracheid
Dead cells at maturation more narrow than the vessels with a thickened
wall with the lignin, imporforated, ends attenuated, contain so many border
pits, tracheids similar to vessels but it have thin cell wall & wide inner cavity.
Vessels occurrence is characteristic of Angiospermae in contrast tracheid is the
transportation element in Gymnospermae & lower plants xylem.

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3- Xylem fibers
Any of the fibers made up of dead sclerenchyma cells in between
the xylem vessels and tracheids of the xylem tissue, and chiefly provide
mechanical support.
The xylem fibers are non-living sclerenchyma cells as they lose
their protoplast at maturity. These cells are found in between the tracheids and
xylem vessels of the xylem tissue. Sclerenchyma cells are narrow and elongated
cells with tapering ends. They are former parenchyma cells that developed
secondary cell walls. Their cell walls become lignified. Their elasticity and
great tensile strength make them an essential component in the xylem as they
protect and provide mechanical support to the major water conducting tissues
of the xylem.
There are two major types of xylem fibers
(1) fiber tracheids
(2) libriform fibers. Fiber tracheids are rather involved in providing mechanical
support than in conduction.

4- Xylem parenchyma:
Parenchyma cells associated with xylem are called “xylem parenchyma”.

 Only living cells of xylem.

 The cell wall is cellulosic and thin.
 They have prominent nucleus and protoplast.
 Cells are colorless and have large vacuoles.
 Living parenchyma cells are found in both primary and secondary xylem.
 Storage of food material in the form of starch, fats, tannins and crystals.

There are two major types of parenchyma cells in secondary xylem :

1. Axial parenchyma (Vertical): derived from fisiform cambial initials, living
tall cells divided into shorter cells.
2. Radial parenchyma (Horizontal): derived from ray initials of the cambium
tall with thickened wall contain simple & border & half border pits.

Tylosis
A tylosis (plural: tyloses) in many plants, the axial and the ray
parenchyma cells develop protrusions that enter tracheary cells when these
become inactive or the xylem tissue is injured, such outgrowth from

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parenchyma cells, Tyloses development occur through the pit – pairs
connecting the parenchyma cells with the tracheary elements. Tyloses have
bladder shape &completely fill the lumen of the tracheid or vessel element, the
nucleus of the originating parenchyma cell & part of the cytoplasm appear in
the cytoplasm.

Primary & secondary xylem

Primary xylem originate from procambium during the primary growth
period, and differentiate to protoxylem and metaxylem, while the secondary
xylem originate from vascular cambium and composed of vessels, tracheids,
fibers, parenchyma xylem, & contain vertical system & horizontal system.

Annual Rings (Growth Rings)

Cambium activity in plants is seasonally because of the weather
change therefore growth cycle appear and consist of :
i- Spring wood / Early wood: wide elements, thin cell wall, most of them are
vessels.
ii- Summer wood/ Late wood: narrow elements thickened cell wall most of
them are fibers, few vessels.
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Heart wood and Sapwood

Heart wood
no active cells, cell walls
thickened Low water ratio, color
dark, cells saturated with oils, gums
resin, pigments. important,
economical.
Sap wood
leaving cells, high water ratio,
thin cell walls, light colour, function
is transportation and mechanical.

Secondary Xylem in Gymnospermae

1) Simpler than Angiosperm xylem composed of tracheids, fibers, parenchyma.
Vessels only occur in one order (Gnelales).
2) In some of gemnospermae contain (Grassulae) up and down border pit.
3) Xylem ray contain homocellular ray composed of parenchyma and
Heterocellular ray composed of parenchyma + tracheids.

Resin Ducts
Occur in gemnospermae xylem, derived in schizogenous intercellular
space among parenchyma cells which produce resin after that convert to
epithelium cells. Sometimes resin duct may become closed by enlarging
epithelial cells. These tylosis – like intrusions are called tylosoids they differ
from tyloses in that they do not grow through pits.

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2- The Phloem
Phloem is the living tissue in vascular plants that transports the soluble
organic compounds made during photosynthesis to parts of the plant where
needed. This transport process is called translocation.

Phloem can be classified into:

1. Primary phloem : It is origin from Procambium
a- Protophloem
b- Metaphloem
Protophloem Metaphloem
1. initiate at early stage of growth initiate at latest age of growth
external position of the vascular occur in determined position of
2.
bundle & occur pressed the vascular bundle of larger
sieve elements are thin sieve elements are more and
3.
enlargement
sieve tube may be conjugated with sieve tube conjugated with
4.
companion cell companion cell
5. Fibers present Fibers absent & some sclerides

2. Secondary phloem : It is origin from vascular cambium, It is assorted as :

a- Axial or vertical system
b- Radial or horizontal system
Vascular rays = Phloem rays + Xylem rays
It is an important characteristic of the secondary xylem & phloem.

In Angiosperms Phloem consist of :

1- sieve tubes, 2- companion cells, 3- fibers, 4- parenchyma cells.
In Gymnosperms Phloem consist of: 1- sieve cells, 2- fibers, 3- phloem
parenchyma, 4- Albuminous cell and other cells or tissues like secretory cells.

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Phloem elements:
1. Sieve Elements: They consist of sieve – tube & sieve cells. The term sieve
tube refer to a longitudinal series of sieve tube members. In both classification,
the characteristic of the wall structures – pits and perforation plates, sieve areas
and sieve plates.

The sieve areas (the term refer to a sieve) are a wall areas with clusters of
pores through which the adjacent sieve elements are inter-connected by strand
like prolongation of their protoplast. Thus the sieve areas are comparable to the
primary pit – fields with plasmodesmata that occur in primary walls of living
parenchyma cells. The sieve area strand are commonly seen associated with the
carbohydrate callose , a polymer of glucose residues united into spirally wound
chain.
Young sieve tube contain nucleus & cytoplasm & other organs but when
it mature nucleus &vacuoles membrane (Tonoplast) analyze and mix gradually
as well as Slim bodies form in cytoplasm which is a small gel, protein
structures, granulosis or filiform, crystal or tubular in shapes.

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Sieve plates: a plates that separate sieve tubes units and consist of porous, its
occur at the ends of sieve tube units. Sieve plates either be simple sieve plates,
or be compound sieve plates.
The sieve units considered to be a life in spite of absent their nucleus
because of:
1. have the ability to form callose & analyses the callose.
2. have the ability to form connecting strand.

2. Companion cells: Specialized parenchyma cells, arise from the same

meristematic cell as the associated sieve tube member.
Companion cells & other living cell form a systematic function to
transport soluble substances through plasmodesmata to the sieve tube.
Companion cells may regulate biological activity for the sieve tube, if the
companion cell die, then the sieve tube function will be absent.
The companion cell is a characteristic of Angiosperms phloem while in
Gymnosperms have a similar cells called Albuminous cells with the sieve
cells, Albuminous cells differ than companion cells in:
1. Origin: albuminous cells does not arise from the same origin of the sieve
cells while the companion cells arise from the same origin of the sieve tube.
2. Position: albuminous cells occur in the ray system while the companion cells
occur within the axial system.
3. Contents: albuminous cells contain high ratio of the albumins.

The sieve cells: occur in gymnospermae phloem, connect with each other with
sieve aerial, cylindrical shape & tall, keep its ability for many years because it
does not lose it nucleus.

3. Phloem fibers: occur in both primary & secondary phloem, in primary

development stage fibers still elongating so the primary fibers may be very

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long. It have secondary cell wall thickened with the lignin contain simple pits.
Fibers may be separate as well as Vitis. Fibers in phloem occur as groups or
layers alternate with sieve elements. Fibers in phloem is very important
economical.

4. Phloem parenchyma: occur in primary & secondary phloem. It has primary

cell wall & contain primary pit fields.
Function of phloem parenchyma is storage (store starch resin substances,
crystals phloem parenchyma may die if the sieve element die, sometime
became septet with cross wall as well as in xylem parenchyma.

Vascular bundles

Xylem & phloem units in monocots vascular bundle are less than in
dicots, arranged as the letter(V/Y) and also contain tracheid, protoxylem have a
large intercellular space called (protoxylem cavity) vascular bundle in
monocots covered with a layer or more of fibers forms (bundle sheath) in some
monocots plants vascular bundle as concentric bundles

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Monocotyledonae Dicotyledonae
1. ground tissue differentiate to cortex,
1. Does not differentiate
pith, medullary rays.
2. Does not found 2. Pericycle found
3.Vascular bundle arranged in one –
3. diffused in ground tissue
two cycles
4. Closed collateral vascular
4. Open collateral vascular bundle
bundle does not contain
contain vascular cambium
vascular cambium
5. xylem vessels arranged in
5. Xylem vessels arranged in strips
form Y or V.
6. vascular bundle covered 6. Vascular bundle does not covered
with bundle sheath (fibers) with sheath
7. no secondary thickening 7. secondary thickening occur

Types of vascular bundles

The following points highlight the four main types of vascular bundle.
The types are:

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1. Collateral Bundle
A vascular bundle in which a
strand c f phloem is present external
to the strand of xylem on the same
radius side by side is known as
collateral bundle. Cambium may be
present or absent in between xylem
and phloem, and so there are the
following two types of collateral
bundle:

(a) Closed collateral bundle

In this type cambium is
absent in between xylem and
phloem. Therefore stems having this
type of bundle do not have normal
secondary growth. Ex.
Monocotyledonous stem. Usually
these bundles are enclosed within
bundle sheath made up of
sclerenchyma and those that lack the
sheath are considered as anomalous
(e.g. Asparagus stem).

(b) Open collateral bundle

An open collateral vascular
bundle has cambium called
fascicular cambium between xylem
and phloem. The bundles can
increase in diameter by normal
secondary growth with the help of
fascicular cambium. Ex.
Dicotyledonous stem.

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2. Bicollateral Bundle
A vascular bundle with phloem situated on the peripheral and inner side
of xylem is known as bicollateral bundle. A strip of cambium termed outer
cambium is present between the peripheral phloem and xylem; another strip of
cambium, termed inner cambium, is also present between inner phloem and
xylem.
The peripheral or external phloem is termed as outer phloem whereas the
inner or internal phloem is called inner phloem. The sequence of vascular
tissues in the bicollateral bundles from periphery toward center is outer phloem,
outer cambium, xylem, inner cambium and inner phloem.
These bundles are open type as strips of cambia are present but the
secondary thickening occurs only by the outer cambium, i.e. cambium present
between the outer phloem and xylem. Ex. Cucurbita stem.

3. Concentric Bundle
A vascular bundle in which one type of vascular tissue surrounds the
other is known as concentric bundle.
In this bundle xylem either encircles or is encircled by phloem and
accordingly the following two types are recognized:

(a) Amphivasal bundle

A vascular bundle in which xylem encircles the central strand of phloem
is known as amphivasal bundle, also called leptocentric bundle. Ex. Dracaena,
Yucca.

(b) Amphicribral bundle

A vascular bundle in which phloem encircles the central strand of xylem
is called as amphicribral bundle, also known as hadrocentric bundle. Ex.
Selaginella.
The concentric bundles, either amphivasal or amphicribral, are closed as
there is no cambium in between xylem and phloem.

4. Radial Vascular Bundle

A vascular bundle, in which the primary xylem and primary phloem
strands are separated from each other by nonvascular tissues and they are
situated on alternate radii of an axis, is known as radial vascular bundle or
radial bundle.

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Cross section of the stem

Introduction
The part of plant which grow above the ground & bear the flowers,
leaves and of the organs & transport nutrients & water to other parts of plant. In
some cases, stem grew underground, the most common features among stems
in plants are:
1. bearing buds leaves & flowers.
2. formed from nodes & internodes.
3. Branches are externally originates & occur in early stage of growth.
4. Apex does not covered with a cap.
5. Epidermis covered with cuticle.
6. Contain collenchyma & sclerenchyma in the cuticle.
7. Vascular bundle either collateral (xylem & phloem occur on one radial) it is
my be Bicollateral (External phloem then xylem then internal phloem)such
as in Cucurbitaceae, or concentric: this type contain: (1) amphiversal
(phloem in the center), (2) amphicribral (xylem in the center)

The primary Structure of the Stem

A. Dicotyledonae Stem
We can flow different tissues that form a mature stem in the primary
growth stage from outside to inside.

1- Epidermis:
One layer of cells, covered with cuticle, may contain stomata or
trichomes.

2- Cortex:
Under the epidermis directly, less thickness than the cortex in the roots,
consist of parenchyma cells & collenchyma either in group forms or as
continuous form its function is supporting some parenchyma contain
chloroplasts. Commonly the cortex does not contain endodermis their for we
can't distinguish the last layer. In few species of plant may contain casparian
strip.

3- Vascular cylinder
Consist of : (1) pericycle, (2) vascular bundles, (3) medullary rays, (4) pith.

1- Pericycle
If the endodermis is clear in the stem then the pericycle will be clear but
almost pericycle mixed with cortex in most of Angiosperm & gymnosperm, in

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case of differentiate it, its composed of parenchyma & sclerenchyma as a
cycles.

2- Vascular bundles
In most of Dicotyledonae, it is open collateral vascular bundles type/ or
it is bicollateral vascular bundles type.

Phloem: sieve tubes, companion cells, parenchyma & fibers (phloem) and
sclerides.

Xylem: occur in radial strips of xylem vessels which consist of metaxylem &
(exarch) &protoxylem (endarch)

Vascular cambium: as a one strip of meristematic cells. Vascular bundle either

be as a separated units or continuous like a cylinder.

Characterization
1- The vascular cambium is a layer of meristematic cells (or initials) that arises
between primary xylem and phloem.
2- Although it is a single layer of cells, in actual practice it is difficult to
distinguish that layer from its immediate derivatives on either side. Hence,
the term cambial zone is used with few exceptions.
3- The vascular cambium is responsible for increasing the diameter of stems
and roots and for forming woody tissue.
4- In dicot stems, the vascular cambium initially differentiates from procambial
cells within the vascular bundles (Fig. below A).
5- Cell division by the cambium produces cells that become secondary xylem
and phloem. As secondary phloem and xylem tissue accumulates, it both
increases the girth of the stem and forms wood and bark. Because cambial
activity is seasonal in temperate zone plants, the wood and bark are laid
down in distinct annual rings (Fig. below C).

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Dr. HADEEL AHMED AL-AMERI
6- Monocots do not have a vascular cambium, even though some of them, such
as palms and the Joshua tree, exhibit secondary growth. Instead, they have
a thickening meristem that produces secondary ground tissue. This
increases the girth of the stem and additional vascular bundles differentiate
within the secondary ground tissue.

Figure : Secondary growth: the origin and structure of vascular cambium in the stem
The vascular cambium is formed in mature dicot stems after stem elongation stops. (A)
Primary xylem and phloem differentiate from procambial tissue in the vascular bundles, and
a fascicular cambium is formed from procambial tissue separating these tissues. (B) Later, an
interfascicular cambium appears between the vascular bundles that is continuous with the
fascicular cambium. (C) The further development of the cambium results in the formation of
a cylinder of vascular tissue. (D) The vascular cambium is a layer of pluripotent dividing
cells whose derivatives differentiate as either xylem elements (vessel members, tracheids,
fibers, or xylem parenchyma) or phloem elements (sieve tube members, companion cells,
fibers, or parenchyma). (E) The dividing cells of the vascular cambium consist of long,
narrow fusiform initials, from which the tracheary elements are derived, and ray initials,
from which ray parenchyma is formed.

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 Advance Plant anatomy / Master student Cross section of stem
Dr. HADEEL AHMED AL-AMERI

7- This fascicular cambium may contribute additional cells to both

the xylem and the phloem of the bundle. At some point the cambium expands
into the ground tissue between the vascular bundles, forming an interfascicular
cambium, completing the ring of vascular cambium.

The vascular cambium is composed of two kinds of cells :

i- ray initials
ii- fusiform initials
In cross section these look very similar. Both are small, flattened cells
with thin walls. When viewed in tangential section, however, ray initials can be
seen to be relatively short, small cells, whereas fusiform initials are very long
and narrow. In gymnosperms the fusiform initials often are several millimeters
in length. Dicot fusiform initials are much shorter, but some still are up to 0.5
mm in length. Cell division in the fusiform initials usually is tangential and the
cell is partitioned down its long axis, forming two equally long, narrow cells.
Some of the cells produced by the cambial initials continue to divide, whereas
others differentiate. Tracheary elements or sieve elements differentiate from
derivatives of the fusiform initials, and derivatives of the ray initials
differentiate as ray parenchyma. The ray parenchyma permits transport of water
from the xylem into the cambium and the tissues of phloem, as well as transport
of photosynthate from the phloem into the cambium and the living cells of the
xylem.

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 Advance Plant anatomy / Master student Cross section of stem
Dr. HADEEL AHMED AL-AMERI

3- Medullary rays
Parenchymatic cells connect the cortex with the pith. Vascular bundle
among these cells, it may be wide or thin.

4- Pith:
Parenchymatic cells large in size in the center of the stem. Pith occupies
large space in the stem in compare with the root, in some plants pith cells
analyzed during growth.

B- Monocotyledon Stem
We can see the following tissues from the outside to inside in mature
stem:

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Dr. HADEEL AHMED AL-AMERI
1- Epidermis:
One layer of cells covered with cuticle, sometime contain chloroplasts and
trichomes.

2- Ground tissue:
Cortex is not found but there is ground tissue composed of
parenchymatic cells, vascular bundle diffuse within the ground meristem and
the outer layers may be sclerids.

3- Vascular bundle
Occur in large number near the epidermis as well as small number in the
center, vascular bundle in monocots are closed collateral vascular bundle &
does not contain vascular cambium.

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 Advance Plant anatomy / Master student Cross section of stem
Dr. HADEEL AHMED AL-AMERI

Dicotyledonae Monocotyledonae
1. ground tissue differentiate
1. Does not differentiate
to cortex, pith, medullary rays.
2. Pericycle found 2. Does not found
3.Vascular bundle arranged in
3. diffused in ground tissue
one – two cycles
4. Open collateral vascular
4. Closed collateral vascular bundle
bundle contain vascular
does not contain vascular cambium
cambium
5. Xylem vessels arranged in 5. xylem vessels arranged in form Y or
strips V.
6. Vascular bundle does not 6. vascular bundle covered with bundle
covered with sheath sheath (fibers)
7. secondary thickening occur 7. no secondary thickening

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Dr. HADEEL AHMED AL-AMERI
REFERENCES
1- CUTLER, D.F. ; BOTHA, C.E.J. and STEVENSON, D.W. (2007). Plant Anatomy
An Applied Approach, First published by Blackwell Publishing Ltd BLACKWELL
PUBLISHING , UK.
2- 2- PAULA, J. and RUDAL, L. (2007). Anatomy of Flowering Plants An Introduction
to Structure and Development, The Edinburgh Building, Cambridge CB2 8RU, UK.