PRIMARY STRUCTURE OF STEM, ROOT AND LEAF

Anatomy of stem

The main functions of stems are to support and elevation of leaves, fruits, and flowers. Stem arranges
leaves in a way that it gets direct sunlight to perform photosynthesis. Xylem and Phloem conduct
water across the plant. Stems stores food, water, and nutrients. Cells of a stem, meristems, produce
new living tissues. Underground stem, Aerial stem, and subaerial stem are three different types of
Stem. A stem has many important functions it performs other than letting you climb a tree. Let us
take an in-depth look at the stem of plants. A plant stem is one of the two main structural axes of a
vascular plant. It is the part of the plant that lies above the ground. Few stems are also
found underground and are considered to be stem modifications.

Functions of Stem

 It supports and holds leaves, flowers, and fruits.

 The stem allows the leaves to arrange in a way that they are able to receive direct sunlight in
order to efficiently perform photosynthesis. The arrangement and position of leaves also
allow for gas exchange.

 The xylem and phloem present in the vascular bundles of stems conduct water and minerals
across the plant.

 Stems bear flowers and fruits in a position that facilitates the processes of pollination,
fertilization, and dispersion of seeds.

 Some stems undergo modification to store food and water. Example: succulents.

 Few green stems contain chloroplasts and are capable of carrying out photosynthesis as well.

 Some stems are modified to carry out vegetative propagation which is a form of asexual
reproduction seen in plants.

Structure of a Stem

The stem divides into nodes and internodes. The nodes give rise to the leaves and hold the buds
which grow into branches. The internodes separate two nodes.

Internally, it contains three basic types of tissues: Dermal tissue, Ground tissue, and Vascular tissue
all of which are made of simple cells.

 Epidermis: The epidermis is a single layer of cells that make up the external tissue of the
stem called dermal tissue. This tissue covers the stem and protects the underlying tissue.
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 Woody plants have an extra layer of protection on top of the epidermis known as bark. In
some cases, the bears’ multi-cellular hairs and a few stomata.

 Ground tissue divides into two- the central portion is known as the pith and the cortex which
lies between the vascular tissue and the epidermis

The cortex can be further divided into three layers:

 Hypodermis: It is the outermost layer of the cortex. It is formed of 4 to 5 cell thick layer
of collenchymatous cells. These cells are living and contain chloroplasts.

 General cortex: Lies below the hypodermis. It consists of thin-walled parenchymatous

cells with intercellular spaces. Some of the cells have chloroplasts and are known as
chlorenchyma.

 Endodermis: The innermost layer of the cortex. It is made up of a single row of compact
barrel-shaped cells without intercellular spaces. The cells of endodermis store starch
grains and so they are known as the starch sheath. Casparian strips are distinctly visible in
endodermal cells.

 The vascular tissue of the stem consists of the complex tissues xylem and phloem which
carry water and nutrients up and down the length of the stem and are arranged in distinct
strands called vascular bundles. Cambium is a strip of thin-walled cells that lie between the
xylem and phloem in dicot plants. Cambium is made up of merismatic cells and is
responsible for secondary growth. It is absent in monocots.

1. Dicot Stem

A thin transverse section of a young stem reveals the internal structure when observed
under me microscope:

1. Epidermis: It forms the single-celled outermost layer of the stem. The outer wall of
epidermal cells is cutinized. It bears multi-cellular hairs and a few stomata. It is
protective in nature.

2. Cortex: Cortex lies below the epidermis. It is differentiated into three zones-

a. Hypodermis: It is formed of 4 to 5 cell thick layer of collenchymatous cells. These

cells are living and contain chloroplasts.

b. General cortex: It lies below the hypodermis. It consists of a few layers of thin-
walled parenchymatous cells with intercellular spaces. Some of the cells have
chloroplasts and they are known as chlorenchyma.

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 c. Endodermis: It is the innermost layer of cortex. It is made up of single row of
compact barrel-shaped cells without intercellular spaces. Since the cells of
endodermis contain starch grains, it is also known as starch-sheath. Casparian strips
are distinctly visible in endodermal cells.

3. Pericycle: It lies below the endodermis. It is formed of semilunar patches of

sclerenchyma. The sclerenchyma cells are dead and rigid with their walls. Pericycie
provides mechanical support to the plant and protects the vascular bundles.

4. Vascular bundles: They are many in number and arranged in a ring enclosed by the
pericycle. The vascular bundles are conjoint, collateral, open and endarch. Each vascular
bundle is composed of Xylem, Phloem and Cambium.

a. Xylem: It is the innermost layer of vascular bundles and lies towards the
centre of the stem. Xylem consists of vessels, tracheids, wood fibres and wood
-parenchyma. The smaller vessels which He towards the centre comprise the
protoxylem and the bigger ones which lie away from the centre are known as
metaxylem.

b. Phloem: It lies below the pericycle and is composed of sieve tubes,

companion cells and phloem parenchyma. The phloem cells store starch,
protein and fats.

c. Cambium: It is a strip of thin-walled cells lying in between the phloem and

xylem. The cambial cells consist of a single layer of meristematic cells.

5. Pith or medulla: It is the central part of the stem, composed of parenchymatous cells
with conspicuous intercellular spaces. Its main function is storage of food and transverse
conduction of food materials.

Example - Helianthus Annuus (Family-Compositae)

T.S. shows the following details:

It is circular in outline and remains differentiated into epidermis, cortex, vascular bundles and
pith.

Epidermis:

1. It is the outermost layer of the stem.

2. Externally it is surrounded by a well-defined cuticle.

3. It is single-layered with tangentially flattened cells.

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4. From some of the cells develop multicellular epidermal hairs.

5. The continuity of the epidermal layer is broken by certain stomata.

Cortex:

6. It consists of outer few layers of collenchyma, some layers of parenchyma and an

innermost layer of endodermis.

7. Collenchyma region is 3 to 6 layered, and the cells are thickened at the corners due to the
deposition of pectin and cellulose.

8. Parenchyma is present inner to the collenchyma in the form of few layers.

9. The cells are thin walled and the parenchymatous region contains many intercellular
spaces.

10. Endodermis is the innermost layer of cortex and consists of barrel shaped cells.

11. Cells of the endodermis contain casparian strips and many starch grains.

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Pericycle:

12. It is present in the form of semi-lunar patches of sclerenchyma outside the phloem of
each vascular bundle.

13. Sclerenchymatous patches of pericycle are interrupted by parenchyma.

Vascular Bundles:

14. These are conjoint, collateral, open and endarch.

15. These are arranged in a ring, and each consists of phloem, cambium and xylem.

16. Phloem consists, of sieve tubes, companion cells and phloem parenchyma.

17. Sieve tubes are narrow and associated with the companion cells.

18. Cambium is present in between xylem and phloem of the vascular bundles and consists of
thin walled, rectangular cells arranged in radial rows.

19. Xylem: It consists of vessels, tracheids, wood fibres and woody parenchyma:

i. Big vessels represent metaxylem while the smaller vessels represent the protoxylem.

ii. Metaxylem consists of reticulata vessels while the protoxylem consists of spiral or
scalariform vessels.

iii. Trachcids are thick walled cells and surround the metaxylem.

iv. Wood fibres are thick walled and lignificd.

v. Wood parenchyma is present in the form of thin walled cells.

Pith:

20. A well developed pith is present in the centre consisting of thin walled, rounded or
polygonal, parenchymatous cells.

Identification:

 Presence of vessels in xylem. (Angiosperms)

 Vascular bundles are conjoint, collateral, open and endarch. (Stem)

 Presence of cambium.

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  Vascular bundles are arranged in a ring.

 Well differentiated cortex.

 Well developed pith. (Dicotyledones)

Special Point: Pericycle is present in the form of semilunar patches of sclerenchyma outside the
phloem

2. Monocot stem

Example - Zea mays – Stem (Family – Graminae)

T.S. exhibits following tissues from outside within:

It is circular in outline with a well-defined epidermis, hypodermis, ground tissue and many
scattered vascular bundles.

Epidermis:

1. It is the outermost layer of stem.

2. The outer wall of cells is covered by a thick cuticle.

3. The continuity of the layer is broken by few stomata.

4. Epidermal hairs are absent

Hypodermis:

5. It is two to three cells thick, sclerenchymatous and present just below the epidermis.

6. Cells are polygonal in shape.

Ground tissue:

7. It is not differentiated into cortex, endodermis, pericycle and pith.

8. The cells are parenchymatous and extend from below the sclerenchyma to the centre.

9. The cells are small and compactly arranged below the hypodermis but they are large, round
and loosely arranged in the centre.

Vascular Bundles:

10. Vascular bundles are many and scattered in the ground tissue with no definite arrangement.
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11. They are small and more in number towards the periphery than the centre of the section.

12. Each vascular bundle is conjoint, collateral, closed and endarch.

13. A well-developed sclerenchymatous sheath surrounds each vascular bundle which is more
prominent at its upper and lower faces.

14. Xylem and phloem constitute the vascular bundle.

15. Phloem:

(i) Consists of only sieve tubes and companion cells.

(ii) Phloem fibres and phloem parenchyma arc absent.

14. Xylem and phloem constitute the vascular bundle.

15. Phloem:

(i) Consists of only sieve tubes and companion cells.

(ii) Phloem fibres and phloem parenchyma arc absent.

(ii) Vessels are in the from of ‘Y’.

(iii) Metaxylem is present at the divergent ends of ‘Y’

in the form of two big oval vessels.

(iv) Protoxylem is present at the lower arm of ‘Y’,

consisting of two small vessels.

(v) Protoxylem is surrounded by tracheids and xylem

parenchyma.

(vi) Inner protoxylem vessel and parenchyma break

down and form a water-containing cavity called
lysigenous cavity.

Identification:

 Presence of vessels in the xylem. (Angiosperms)

 Vascular bundles are conjoint, collateral and endarch. (Stem)

 No differentiation of ground tissue.

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  Sclerenchymatous hypodermis.

 Vascular bundles are closed.

 Scattered vascular bundles.

 Absence of secondary growth. (Monocot)

Special Points:

1. Scattered vascular bundles.

2. ‘Y’ -shaped vessels.

3. Presence of protophloem and metaphloem.

Anatomy of Root

1. Anatomy of Dicot root

(gram)

The plants whose seeds have only one cotyledon are

called dicots. The structure of dicot root varies greatly
from that of the monocots. Dicot roots of gram shows
following distinct region in its Transverse section with
following features:

1. Epiblema

2. Cortex

3. Endodermis

4. Pericycle

5. Vascular bundles

6. Pith

1. Epiblema or Epidermis - It is the outermost

unilayered with several unicellular root hairs. It
consists of thin walled, compactly arranged living
parenchymatous cells. Usually epiblema is
characterised by absence of stomata and cuticle.

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Sometimes the epiblema may be less cuticularised. It provides protection to the roots due to
presence of unicellular root hairs it also helps in absorption of water and minerals from soil

2. Cortex - It is thin walled, multilayered region made from circular or polygonal

parenchymatous cells. they usually have intercellular spaces. The cortical cells have no
chloroplast but may contain leucoplast for storage of starch grains. The cortex is responsible for
transportation of water and salts from the root hairs to the center of the root.

3. Endodermis - It is the innermost layer of cortex and covers the stele. It consists of compactly
arranged barrel shaped parenchyma without intercellular spaces. Most of the cells are
characterised by the presence of special thickening of suberin and lignin on their radial and
tangential walls called casparian strips. Some endodermal cell near protoxylem has no
casparian strips and called passage cells or transfusion cells. These cells allow radial diffusion of
water and minerals through the endodermis.

4. Pericycle - It is the outermost layer of stele and composed of uniseriate layer of

parenchymatous cells without intercellular spaces. Some dicots and hydrophytes do not bear
pericycle. Several lateral roots and lateral meristem arise from pericycle region (hence lateral
roots are endogenous in origin). At the time of secondary growth, it produces secondary
cambium or phellogens.

5. Vascular bundles - They are 2-8 in number, radial and arranged in ring. Xylem and
phloem bundles are separated from each other by parenchymatous cells called conjuctive
or complementary tissue.

 Xylem is exarch (i.e. protoxylem towards the periphery and metaxylem towards the centre)
and consists of tracheids, vessels, xylem parenchyma and xylem fibres.

 The pholem forms oval masses beneath the pericycle, alternating with xylem bundles.
Pholem consists of sieve tubes, companion cells and pholem parenchyma. Usually pholem
fibres are absent or reduced.

6. Pith - it is feebly developed and centrally located. It consists of thin walled, polygonal
parenchyma cells with intercellular spaces. In dicots roots, it may be reduced or absent. It helps
in storage of food materials.

Distinguishing Features of Dicot Root

The typical dicot roots show following features.

 Epiblema is uniseriate, thin walled, colourless without intercellular spaces and produce
unicellular root hairs, hence also called as piliferous layer or rhidodermis.

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 Cortex is homogenous (without differentiation).

 Endodermis consists of barrel shaped compact parenchymatous cells. It contains

both casparian stripes and passage cells.

 Pericycle uniseriate and become meristematic to give secondary roots and secondary tissues.

 Vascular bundles are radial; Xylem is exarch, number of xylem bundles varies from 2 to 4
rarely more (upto 6-8). Metaxylems are angular arranged in linear.

 Usually conjuctive tissues are well developed.

 Pith is very small or completed obliterated.

2. Anatomy of Monocot roots

That plant whose seed contains only one cotyledon or embryonic leaf is known as
monocotyledon or simply monocot.

Primary Structure of Monocot roots

The typical monocot roots show following features:

 Epiblema is single layered, thin walled, colorless, polygonal without intercellular spaces,
with presence of unicellular root hairs, hence also called as piliferous layer or rhizodermis.

 Cortex may be heterogeneous with outer dead exodermis.

 Endodermis consists of barrel shaped parenchyma without intercellular spaces. Casparian

stripes are little present but passage cells are absent.

 Pericycle gives lateral roots only, secondary growth is absent.

 Vascular bundles are radial; Xylem is exarch, bundles more than six. Metaxylem elements
are oval or circular.

 Conjunctive tissues are limited or even absent.

 Pith is large or rarely reduced.

Features of Different Regions of Monocot Root

1. Epiblema is the outermost single layer made from compactly arranged parenchymatous cells
without intercellular space. Usually Epiblema has no stomata but bears unicellular epidermal
root hairs and less amount of cutin. It contains more cuticle than dicot roots. The root hairs

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 and thin walled epidermal cells take part in the
absorption of water and minerals from the soil.
The epiphytes have several layered hygroscopic
epidermis, called velamen tissues. It is made
from spongy dead cells which helps in
absorption of water from atmosphere. It also
checks excessive loss of water from cortex.
Usually the wall of velamen has spiral or
reticulate secondary thickening of cellulose,
pectin and lignin.

2. Cortex is a multi-layered well developed and

made from oval parenchymatous cells with
intercellular spaces. The intercellular spaces
usually help in gaseous exchanges, storage of
starch, etc. In monocots and several old roots,
few layers of cortex just below epiblema give
rise to a single or multilayered cuticularised
sclerenchymatous region called exodermis.
Cortex helps in mechanical support to the roots
(like hypodermis to stem).

3. Endodermis is innermost layer of cortex made

from barrel shaped parenchyma. It forms a
definite ring around the stele. These cells are
characterized by the presence of casparian
stripes. It is deposition of suberin and lignin,
and their radial and tangential walls. Usually
passage cells are absent in monocot roots. Due
to presence of casparian stripes, endodermis forms water tight jacket around the vascular
tissues, hence it is also called biological barrier. It regulates the inward and outward flow of
water and minerals and prevents diffusion of air into xylem elements.

4. Pericycle is uniseriate (multiseriate in Smilax) and made from thin walled parenchymatous
cells. It is outermost layer of stellar system. Usually it is made from parenchymatous cells
but it may become sclerenchymatous in older roots. Several lateral roots arise from this
layer. Hence, lateral roots are endogenous in origin.

5. Vascular bundle is radial, arranged in a ring (except mangrove, which also contains
lenticels), polyarch (presence of many alternating xylem and phloem bundles). Xylem and
phloem are found at different radii alternating with each other (radial). The number of xylem

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 and phloem vary from, 8 to 46 (100 in pandanus). The xylem is exarch, i.e. the protoxylem
lies towards periphery and metaxylem toward center. The protoxylem has smaller vessels
with spiral or annular thickening, whereas the walls of metaxylem contains pitted thickening.
Phleom consists of seive tubes, companion cells and phloem parenchyma. Usually phleom
sclerenchyma or fibers are absent. The phloem is also exarch (protophloem towards the
periphery and metaphloem towards the center). Secondary growth is absent in monocot roots
due to lack of vascular and cork cambium. Conjunctive tissue is parenchymatous tissues
which separates xylem and phloem bundles. It may become sclerenchymatous in older roots.

6. Pith is large, well developed portion of monocot root. It occupies the central portion and
made from thin walled parenchymatou tissue with intercellular spaces. It contains abundant
amount of starch grains.

Monocot root of maize have bands of vascular bundles. Bundles are not separate and vessels are
not found in linear rows, but arranged in V-shaped structure.

Anatomy of Leaves

The foliage leaves are characterised by green colour, thinness and flatness. They develop as
protrusions from the shoot apex and are organs of limited growth. Leaves are very important
vegetative organs, as they are chiefly concerned with the physiological process, photosynthesis
and transpiration. Like other organs they also exhibit three tissue systems’

Epidermal tissue system consists of the epidermal layers occurring on the adaxial (upper) and
abaxial (lower) sides. Occurrence of stomata and outgrowths are distinctive features. The ground
tissue system, is known as mesophyll tissue. It is often differentiated into columnar palisade
parenchyma on the adaxial side and irregular or isodiametric spongy parenchyma on this
differentiation in mesophyll is referred to as dorsiventral, what is very common in dicotyledons.
One with undifferentiated mesophyll, as commonly found in the monocotyledons, is known as an
isobilateral leaf. Presence of conspicuous air spaces in the mesophyll is another marked feature.
The gaseous exchange between the internal photosynthetic tissues and outside atmosphere thus
becomes easy.

The vascular tissue system is composed of vascular bundles which are usually collateral and
closed. But the bundles entering the leaf occupy such a position that xylem occurs on the upper
side and phloem on the lower. The vascular tissues, in fact, form the skeleton of the leaf, on
which other tissues—the ground tissues, remain inserted. The petiole may continue into the
midrib which bears branches and sub-branches ultimately ramifying in the leaf lamina in both
reticulate and parallel type of venation.

The ultimate branches are very small and terminate in what are known as bundle ends. Often
these ends bend into minute specialised photosynthetic areas known as vein islets or they may

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just extend into the mesophyll. Bundle-ends vary considerably in the leaves, but commonly it
consists of a single tracheid with a single sieve element or specialised parenchyma representing
xylem and phloem respectively, surrounded by a parenchymatous bundle sheath.

1. Anatomy Dorsiventral Leaves (Dicot leaves)

Example - Leaf of Mango:

I. Epidermis: There are two epidermal layers on adaxial and abaxial surfaces of the leaf. Each is
uniseriate, composed of a row of compactly-set tabular cells. The outer walls are cutinised and
possess thin cuticle, the thickness being more pronounced in the cells of the upper epidermis than
those on the lower side. Stomata occur on the lower epidermis.

II. Mesophyll:

The ground tissue forming the mesophyll is differentiated into palisade and spongy cells. The
palisade cells occur towards upper epidermis. They are columnar cells with scanty intercellular
spaces and remain arranged more or less at right angles to the upper epidermis. Chloroplasts are
abundantly present, which particularly occur along the radial walls of the cells.

There are two layers of palisade cells. The spongy cells occur towards the lower epidermis. They
are quite loosely arranged with conspicuous intercellular spaces. The number of chloroplasts is
naturally much smaller here, which explains the pale green colour of the lower surface of the
leaf.

III. Vascular bundles:

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Bundles are collateral and closed. They are located in the mesophyll. The size of the bundle
depends on the position one chooses to take in making a section.

A bigger bundle is composed of xylem and phloem, the former occurring towards upper
epidermis and the latter towards the lower side. The xylem is made of tracheary elements, and
the phloem of sieve tubes and companion cells. The bundle remains surrounded by a row of
colourless parenchyma cells.

This band is referred to as bundle sheath or border parenchyma. Thus the bundle is not in direct
contact with the mesophyll cells. Parenchyma and often collenchyma cells are present on the
outer and inner sides of the bundle which may reach up to the two epidermal layers. These cells
constitute what is known as bundle sheath extension.

2. Isobilateral Leaves (Monocot leaves):

These leaves are common in the monocotyledons. Here stomata occur on both the epidermal
layers, though they are more abundant on the abaxial side. Mesophyll hardly shows
differentiation between palisade and spongy cells. A few common isobilateral leaves have been
selected for the study of internal structures.

Example -1- Leaf of Maize:

I. Epidermis:

Both upper and lower epidermal layers are uniseriate and composed of more or less oval cells
with cuticularised outer walls. Upper epidermis may be easily identified due to presence of large
and empty bulliform cells. Stomata occur on both the epidermal layers.

II. Mesophyll:

The mesophyll does not show differentiation into palisade and spongy cells, but is made of rather
compactly-arranged isodiametric cells.

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III. Vascular bundles:

The bundles are collateral and closed ones which remain arranged in parallel series. Majority of
the bundles are small, but fairly large bundles occur at regular intervals. Small bundles have
xylem on the upper and phloem on the lower sides surrounded by large parenchyma cells
forming the bundle sheath.

The cells of the sheath contain plastids, often with starch grains. It is assumed that this layer
serves as a temporary storage tissue, apart from-conducting the products of photosynthesis to the
phloem. Xylem, as usual, consists of tracheary elements, and phloem of sieve tubes and
companion cells.

A large vascular bundle practically resembles that of a stem. Sclerenchyma cells occur in patches
on both edges of the bundles, obviously for giving mechanical strength.

Example – 2- Leaf of Bamboo:

I. Epidermis:

As usual there are two epidermal layers. The upper epidermis possesses a number of conspicuous
bulliform cells. The lower one bears stomata and remain covered with strong cuticle.
Considerable deposition of silicon is a distinctive character. Stiff sharply pointed hairs are also
present.

II. Mesophyll:

It is composed of compactly-arranged cells, without showing any differentiation into palisade

and spongy cells. The cells are of rather palisade type, though not much elongate. Some distinct
cavities are present here and there.

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III. Vascular bundles:

These are collateral and closed ones which remain arranged in parallel series. Xylem occurs on
the adaxial and phloem on the abaxial sides. Larger bundles have more distinct xylem and
phloem surrounded by a bundle sheath, and has patches of sclerenchyma cells on the two sides.

NORMAL SECONDARY GROWTH IN DICOT STEM AND ROOT

The growth in length of main axis by the activity of apical meristem is called primary
growth. Increase in thickness or girth of the aixs due to the formation of secondary tissue is
called secondary growth.

The further growth in thickness is completed by cambial activity, called secondary

growth in thickness. The tissues, formed during secondary growth are called secondary tissues.
The cambium forms secondary tissues in the stelar region and cork-cambium form secondary
tissues into cortical regions. Secondary growth occurs only in dicot stem and root. It is usually
absent in monocot root and stem.

In dicot stem, secondary growth takes place through the following steps:

1. Formation of Cambium Ring: Formation of the cambium ring is the first step of
secondary growth. The cambium of vascualr bundles becomes meristematic. At the same
time some of the medullary ray cells lying at the level of cambium also become
meristematic and form a strip of interfasciular cambium together with intrafasciular
cambium form a complete circular ring, which is called cambium ring. The cambium ring
forms the secondary tissues in the stelar region.
The vascualr cambium consists of two types of cells, the fusiform initials and the ray
initials. The fusiform initials are vertically oriented and divide to form the elements of
xylem and phloem. The cells of ray initials are smaller and isodiametric which give rise
to vascular rays of parenchymatous cells.
2. Formation of Secondary Vascualr Tissues: The cambium ring cuts off new cells, both
on outer and inner sides. The new cells formed on the outer side gradually modify into
the elements of secondary phloem. The cells formed on the inner side gradually modify
into secondary xylem.

Structure of Cambium:

The vascular cambium (plural cambia) is a plant tissue located between the xylem and
the phloem in the stem and root of a vascular plant, and is the source of both the secondary
xylem growth (inwards, towards the pith [material at the center of plant, often dead and/or
deteriorated, that is composed of parenchyma tissue][1]) and the secondary phloem growth
(outwards [to the bark, rough or smooth, of the plant]). It is a cylinder of

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unspecialized meristem cells that divide to give new cells which then specialize to
form secondary vascular tissues.

Vascular cambia are found in dicots and gymnosperms but not monocots, which usually lack
secondary growth. A few leaf types also have a vascular cambium. Vascular cambium does not
transport water, minerals, or dissolved food through the plant. It does, however, produce the
phloem and xylem, which do perform these functions

Vascular meristems generate cells which differentiate into xylem and phloem. The apical
meristems in the shoot and root contain procambium, the primary vascular meristem. During
secondary growth, a lateral vascular meristem called cambium develops mainly from the
procambium embedded between the differentiated xylem and phloem. The cambium present
between primary xylem and primary phloem is called intrafasicular cambium. At the time of
secondary growth, cells of meduallary rays, in a line with intrafasicular cambium, become
meristematic and form interfascicular cambium. The intrafascicular and interfascicular
cambiums, therefore, represent a continuous ring which bisects the primary xylem and primary
phloem and is known as cambium ring. The vascular cambium then produces secondary xylem
on the inside of the ring, and secondary phloem on the outside, pushing the primary xylem and
phloem apart.

Vascular Cambium

1. The Vascular cambium is the remnant part of the apical meristem. It is present between the
xylem and phloem of Vascular bundle.

2. This cambium continues to divide and adds secondary phloem on its outer side surrounded
secondary xylem on its innerside.

3. The vascular cambium produces primary xylem and medullary rays respectively.

Cork Cambium:

1. The cork cambium is a true secondary meristem which develops in the region outside the
vascular tissues.

2. This cambium gives rise to cork and secondary cortex towards outer and inner sides
respectively.

3. The cork cambium produces phellogen, phellem and phelloderm collectively known as
periderm

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There are two different types of cambium cells:

1. The ray initials, which are more or less isodiametric and give rise to vascular rays; and
2. The fusiform initials, the elongate tapering cells that divide to form all cells of the
vertical system.

The cambial cells are highly vacuolated, usually with one large vacuole and thin peripheral
cytoplasm. The nucleus is large and in the fusiform cells is much elongated. The walls of
cambial cells have primary pit fields with plasmodesmata. The radial walls are thicker than
tangential walls, and their primary pit fields are deeply depressed.

Secondary Growth in Dicot Stem

Primary growth produces growth in length and development of lateral appendages.

Secondary growth is the formation of secondary tissues from lateral meristems. It increases the
diameter of the stem. In woody plants, secondary tissues constitute the bulk of the plant. They
take part in providing protection, support and conduction of water and nutrients.

The vascular bundles of dicotyledonous stems are collateral and open, and arranged in a
ring. They contain a single layer of cambium cells, which separate the xylem from the phloem,
called fascicular cambium, i.e., the cambium of the vascular bundle, (fascicle = bundle). When
the primary xylem and primary phloem are first differentiated there is no cambium across the
pith rays or medullary rays to connect the edges of the cambium within vascular bundles.

Secondary tissues are formed by two types of lateral meristems, vascular cambium and cork
cambium or phellogen. Vascular cambium produces secondary vascular tissues while phellogen
forms periderm.

Secondary growth occurs in perennial gymnosperms and dicots such as trees and shrubs. It is
also found in the woody stems of some herbs. In such cases, the secondary growth is equivalent
to one annual ring, e.g., Sunflower.

A. Formation of Secondary Vascular Tissues:

They are formed by the vascular cambium. Vascular cambium is produced by two types of
meristems, fascicular or intra-fascicular and inter-fascicular cambium. Intra-fascicular cambium
is a primary meristem which occurs as strips in vascular bundles. Inter-fascicular cambium arises
secondarily from the cells of medullary rays which occur at the level of intra-fascicular strips.

These two types of meristematic tissues get connected to form a ring of vascular cambium.
Vascular cambium is truly single layered but appears to be a few layers (2-5) in thickness due to
presence of its immediate derivatives. Cells of vascular cambium divide periclinally both on the
outer and inner sides (bipolar divisions) to form secondary permanent tissues.

18
The cells of vascular cambium are of two types, elongated spindle-shaped fusiform initials and
shorter isodiametric ray initials (Fig. 6.29). Both appear rectangular in T.S. Ray initials give rise
to vascular rays.

Fusiform initials divide to form secondary phloem on the outer side and secondary xylem on the
inner side (Fig. 6.28 B). With the formation of secondary xylem on the inner side, the vascular
cambium moves gradually to the outside by adding new cells.

The phenomenon is called dilation. New ray cells are also added. They form additional rays
every year (Fig. 6.28 D). The vascular cambium undergoes two types of divisions— additive
(periclinal divisions for formation of secondary tissues) and multiplicative (anticlinal divisions
for dilation).

Ray initials produce radial system (= horizontal or transverse system) while fusiform initials
form axial system (= vertical system) of secondary vascular tissues.

If the cell that is differentiated is next to the xylem it forms xylem, while if it is next to phloem it
becomes phloem towards the outer side of the cambium. The cambium cells divide continuously
in this manner producing secondary tissues on both sides of it. In this way, new cells are added to
the xylem and the phloem, and the vascular bundles increase in size.

While there is more or less alternation in the production of xylem and phloem cells from a
cambium cell, more cells are formed on the xylem side than on the phloem side. The cells
formed from the cambium in the region of the pith rays become pith-ray cells. The activity of the
cambium thus increases the length of the pith rays grow equally.

19
20
The formation of new cells from the cambium result in an enlargement of the stem that is known
as the secondary thickening. The formation of new cells in secondary thickening continues
throughout the life of the plant. It is in this way that the trunks of trees continue to grow in
diameter.

Secondary Xylem:

21
The cambium ring cuts off new cells on its inner side are gradually modified into xylary
elements, called the secondary xylem. This tissue serves many important functions, such as
conduction of water and nutrients, mechanical support, etc. The secondary xylem of tree trunks
is of great economic value, since it constitutes the timber and wood of commerce.

The secondary xylem consists of scalariform and pitted vessels, tracheids, wood fibres and wood
parenchyma. These elements of secondary xylem are more or less similar to those occur in
primary xylem. Vessels or tracheae are most abundant and are usually shorter than those of
primary xylem. Mostly the vessels are pitted. Annular and spiral tracheids and vessels are
altogether absent.

Xylem parenchyma cells may be long and fusiform, but sometimes they are short. They are
living cells and usually meant for storage of food material (starch and fat) in them. Tannins and
crystals are frequently found in these cells. Xylem parenchyma may occur either in the
association of the vessels or quite independently. The fibres of secondary xylem possess thick
walls and bordered pits.

Xylem Rays:

Ray initials of the cambium ring form some narrow bands of parenchymatous cells. These cells
extend radially from the pith to the phloem. These are called secondary medullary
rays or vascular rays. The rays present in xylem are xylem rays and the rays present in phloem
are called phloem rays.

The xylem rays or wood rays extend radially in the secondary xylem. They are strap or ribbon
like. They originate from the ray initials. The xylem rays run as a continuous band to the
secondary phloem through the cambium, thus forming a continuous conducting system. All
vascular rays are initiated by the cambium and, once formed, are increased in length indefinitely
by the cambium.

22
The xylem rays traverse in the secondary xylem and establish communication with the living
cells of the vascular tissue. In gymnosperm wood where no wood parenchyma is present, every
tracheid is in direct contact with at least one ray. Vessels also in their longitudinal extent come
into contact with many rays.

The xylem rays help in the exchange of gases. They also aid in the conduction of water and food
from phloem to the cambium and xylem parenchyma.

Secondary Phloem:

They consist of sieve tubes, companion cells, and phloem parenchyma and phloem fibres. The
primary phloem present on the outside gets crushed and is represented by small patches.

23
Secondary growth in extra stelar zone:

This zone extends from hypodermis to endodermis and made up of primary ‘issues during
primary growth.

Secondary growth takes place in following steps:

(a) Origin of cork cambium (Phellogen):

First or second layer of hypodermis or outer cortex becomes meristematic and known as cork
cambium.

(b) Activity of cork cambium:

It divides in tangential plane cutting cells towards inner as well as outer side. The outer cells are
compactly arranged and have thin wall in the beginning but as they mature there is gradual loss
of living matter and the cell wall becomes thick due to deposition of fatty substance suberin. It is
known as cork (phellem). Thin wall cell cuts of towards the inner side of the cork cambium are
known as phelloderm (secondary cortex). Its cells are living with cellulose cell wall.

(c) Formation of periderm:

Cork (phellem), cork cambium (phellogen) and secondary cortex (phelloderm) together form
periderm.

Bark:

All the tissues outside the vascular cambium are collectively known as Bark. It includes
secondary phloem, primary phloem, pericycle, endodermis, primary cortex and periderm.

(d) Lenticel:

It is a small portion of periderm where the activity of phellogen (cork cambium) is more. Cells
are loosely arranged, thin walled with numerous inter cellular spaces known as complimentary
cells. Lenticel (breathingpores) helps in gas exchange in woody part of the plant and helps in
lenticular transpiration.

The Secondary Growth in Dicot Root

The secondary growth in root also takes place by the activity of the cambium and cork
cambium. It is a usual feature of dicotyledonous and gymnospermous roots, where it generally
starts at a very early stage, so much so that it is difficult to get the roots without secondary
growth in most of the cases.

24
1. Activity of Cambium: Certain of the cells of conjunctive tissue just beneath the phloem
become meristematic and form strips of cambium. The number of strips produced depends upon
thus number of phloem bundles present. In a diarch root two, in triarch root three and in tetrarch
root four such strips are formed. These strips exertend both ways in between phloem and xylem
and ultimately unite with the pericyclic cells lying just outside the protoxylem.

The pericyclic cells divide tangentially and produce two layers of which the cells of inner layer
also become meristematic and unite with the strips of cambia and thus, a continuous wavy band
of cambium is produced extending down the phloem and over the xylem (Fig. 13.6). It becomes
active and forms new cells.

It divides by periclinal divisions and then by anticlinal divisions for increase in circumference.
The strip of cambium below the phloem becomes active earlier and the activity is much faster on
the inner side. Because of this, phloem and cambium strip below it are pushed outward and the
wavy band of cambium now becomes circular to form a cambium ring. Now the entire of the
cambium becomes active.

The cells formed on the inner side get differentiated into secondary xylem. It consists of
comparatively large vessels, tracheids, a little wood fibres and well evolved xylem parenchyma.
The activity of cambium is so fast on the inner side that after secondary growth xylem forms the
main bulk of the root and is present in the form of solid core.

The primary xylem bundles can remain intact up to the last or the crushed. The annual rings like
the stem are not visible in roots. The pith is entirely crushed, or if some part is left the cells
become thick walled.

The secondary vascular tissues produced by the activity of cambium do not form a continuous
ring but are interrupted by the bands of radially-elongated, parenchymatous cells, known as
primary medullary rays. These are formed above each primary xylem patch and extend up to the
phloem. Sometimes, other smaller medullar rays can also develop from other parts of cambium
and may be known as secondary medullary rays. The number of rays goes on increasing with the
increase in the size of the vascular cylinder.

The secondary xylem cells vary in quantity if different roots, they have only tracheids in
gymnosperms, only vessels in willow and both tracheids and vessels in most of the plants. In
some storage roots storage parenchyma develops in the secondary xylem. The cells of secondary
xylem are arranged in definite rows when first produced become irregularly disposed due to
differential enlargement of various tracheid elements.

25
Like the stem, in roots of perennial trees, shrubs and woody climbers also, the xylem elements
produced in the bringing of each season are larger and thin walled, while those formed during
late are smaller in size and are thick walled. Thus, annual rings are produced.

2. Activity of Cork Cambium: The secondary tissues produced by the activity of cambium
exert a pressure on the outer tissue. To withstand this pressure, the cells of pericycle become
meristematic and function as the phellogen or cork cambium. The cells of pericycle divide
tangentially. Similar to stem, here also it produces layers of cork or phellem on the outside and
secondary cortex or phelloderm on the inner side.

The bark in the case of roots includes cork, endodermis, cortex and epiblema. In certain cases,
the cork cambium may be formed from the phloem cells. In this case, the pericycle also produces
the part of bark. Subsequent barks have only cork. Lenticels can also be formed here and there.
When the bark is removed, the new cork cambium layer is formed from the parenchyma
produced by the previous cork cambium.

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PERIDERM

In roots and stems having secondary growth, the epidermis is replaced by a protective layer of
secondary origin known as periderm. It generally develops in gymnosperms and dicotyledonous
axis and is rarely produced in leaves or monocotyledons. The periderm is also formed along
surfaces exposed after abscission of plant parts, such as leaves or branches. It also evolves as
protective layer near injured parts (wound periderm).

The term periderm is more distinct than bark. The latter designates all tissues outside the
vascular cambium. In secondary state, it consists of secondary phloem and all tissues outside it.
It can be distinguished into outer non-living and inner living parts. The functional phloem is the
innermost part of the living bark.

Structure of Periderm:

The periderm consists of the phellogen or cork cambium, the meristem that produces the
periderm; the cork or phellem, the protective tissue produced outside by the phellogen, and the
inner cortex or phelloderm, the living parenchyma, formed inside by the phellogen. Because of
the formation of cork, the tissues outside it usually die out.

27
The phellogen is simple in structure and it has only one kind cells. They appear as continuous
tangential layer (lateral meristem) of rectangular, radially flattened cells in cross section. In
longitudinal section they are rectangular or polygonal in outline.

The phellem or cork cells are often prismatic in shape and can be elongated vertically, radially or
tangentially to form irregularly shaped structures. These are compactly arranged and absent
inter-cellular spaces. These are dead cells at maturity. The cell walls are suberized.

The suberin, a fatty substance, generally occurs as a distinct lamella that covers the original
primary cellulose wall. The thickness of cells walls of cork is variable. In thick-walled cells
lignified cellulose layer exists on the inner side of suberin lamella. The walls of cork cells are
brow or yellow in colour, while coloured resinous or tanniferous material can be filled in the
lumina.

The commercial cork generally has thin walls and air-filled lumina. It is highly impervious to
water and resistant to oil. It has thermal insulating quality and is light in weight. The mature cork
of type is also a resilient and compressible tissue. These properties also make it useful to protect
the plant parts.

In many species viz., Rhododendron maximum, the phellem includes non-suberized cells, called
phelloid cells together with cork. These may also be thin or thick walled. When thick walled
these are known as sclereids.

The phelloderm is a typical parenchyma which may be distinguished from other parenchyma by
being present in the same radial files as the phellem cells.

Development of Periderm:

The first periderm commonly appears during the first year of growth of stem and root. In stem
most usually it originates in the sub-epidermal layer. In some species, the first periderm appears
rather deep in the stem, usually in the primary phloem viz., Berberis, and Vitis etc. In roots, in
other word, the first periderm originates in the pericycle. In some cases, where the root cortex
serves for food storage, it can originate near the surface also. The subsequent periderms may
appear the same year or later in the successively deeper layers beneath the first, i.e., from the
parenchyma of the phloem, including ray cells.

28
The first phellogen is generally initiated uniformly around the circumference of the axis.. The
subsequent periderms appear as discontinuous but overlapping layers. These can finally appear
as continuous, or partly so, layers, around the axis.

The phellogen arises from living cells which are potentially meristematic. In these cells, the
initiating divisions can start in presence of chloroplast and ergastic substances, viz., starch,
tannins. But gradually these structures disappear. The phellogen is initiated by periclinal
divisions and it forms the phellem and phelloderm by the same type of divisions.

The activity of the phellogen is more on the outside and thus, the amount of phelloderm formed
is generally very small, sometimes restricted only to few layer of cells. A given phellogen cell
usually produces a few cork cells every year. There may or may not be certain difference in the
size of cells formed during earlier part of the season or later part of the year.

On the basis of manner of function, two kinds of barks are distinguished—scale bark and ring
bark. The former occurs when subsequent periderms exist in restricted overlapping strata, each
cutting out scale of tissue, e.g., Finns and Pyrus etc. Ring bark results from the formation of
successive periderms approximately concentrically around axis, in the produce of sheets e.g.,
Vitis, and Lonicera, etc.

Lenticels:

The limited part of the periderm with more active phellogen producing a tissue with intercellular
spaces can be called a lenticel. The phellogen in this part itself also has intercellular spaces. Due
to this special structure the lenticels are used for entry of air through the periderm. The lenticels
are commonly produced in stems and roots.

29
The protrude out of the surface of axis. Their size is also variable, becoming even up to 1
centimeter. Generally they are irregularly distributed but sometimes they exist in vertical rows
opposite the rays.

The phellogen in the lenticels is a continuation of the rest but is bent inwards in this area. The
loose tissue produced outside is called filling tissue or complementary tissue. On the inner side
the phelloderm is produced in the normal fashion.

Bark:

In large trees, due to continued formation of secondary tissues in the intrastelar region the
periderm may not be adequate to withstand the inwardly generated pressure. In that case addi-
tional layers of periderm are formed successively in the deeper regions of the cortex, pericycle
and even phloem.

The outwardly cut-off phellem or cork cells by the phellogen ultimately get devoid of water and
food supply and eventually become dead. All these dead tissues lying outside the active
phellogen constitute the bark of the trees.

The term bark is loosely used and the term rhytidome is used for the outer bark covering all the
tissues external to the innermost phellogen. Actually, the term bark is given to all tissues external
to the vascular cambium.

Due to the formation of successive layers of periderm in the deeper regions of the stem the bark
is formed in concentric rings surrounding the entire stem, which is known as ring bark. In some
plants the periderm is formed as overlapping scale-like layers, known as scale bark. In
Eucalyptus, Platanus, etc. the bark is intermediate between these two types, where the outer
layers of the bark peel off in the form of sheets

ANOMALOUS SECONDARY GROWTH

Many dicotyledons show secondary growth that deviates considerably from the normal
secondary growth. The deviating methods of secondary thickening are known as abnormal or
anomalous- although the normal and abnormal produces of growth are not sharply separated
from one another.

A. Anomalous Position of Cambium:

Stems of many unusual shapes or types are formed by the unusual position of the cambium.

B. Abnormal Behavior of Normal Cambium:

30
Sometimes when the normal cambium starts cutting cells at many places irregularly, and forms
at certain places much larger portions of xylem than of phloem, and at other places more phloem
than xylem, a ridged and furrowed xylem cylinder is produced. This can be of simple structure
(e.g., some Bignoniaceous genus) or very complex (viz., in Bignonia sp.)

Secondary Growth in Bignonia

In some type of stem as in Bignonia and other members of family Bignoniaceae, the cambium is
normal in disposition and activity to begin with, but it soon cuts off different proportions of
xylem and phloem in different points.

At four points arranged in form of a cross, formation of secondary xylem is reduced and that of
secondary phloem correspondingly increased. As a result the woody cylinder appears to have
four longitudinal grooves which become increasingly deeper with secondary growth.

The cambium breaks up into a number of strips, widest ones occurring opposite the four
projecting ridges of wood and the narrow ones at the bases of the grooves. As a result peculiar
structure with ridged and furrowed xylem cylinder is formed. Deeply ridged vascular cylinder is
formed in some plants due to the fact that the cambium produces only ray parenchyma cells at
some points.

Anatomy of Bignonia – Stem (Family – Bignoniaceae):

Epidermis

1. Single-layered epidermis consists of rectangular cells.

2. A thick cuticle is present.

3. A few multicellidar hairs are also arising from some cells.

Cortex:

4. Ii is well-differentiated into collenchyma and parenchyma.

5. Collenchyma is present below the epidermis in the ridges in young stem but at maturity there
develops sclerenchyma.

6. Parenchyma is present below the sclerenchyma or collenchyma in the ridges and directly
below the epidermis in the grooves.

7. In old stem cortex consists of cork, cork cambium and cortex.

8. Endodermis is undistinguishable from cortical cells. The cells lack casparian strips.

31
Pericycle:

9. It is in sclerenchymatous patches.

Vascular system:

10. It phloem, secondary phloem, xylem and primary xylem.

11. Four longitudinal furrows of secondary phloem arc present which are wedged in between the
secondary xylem cylinder.

12. Vascular bundles are conjoint, collateral, open and endarch.

13. Primary phloem is crushed and present in small patches.

14. Secondary phloem is in the form of a ring which remains intruded into the secondary xylem
at four places.

15. Intruded furrows (four) of secondary phloem are arranged in the form of a cross.

16. In Bignonia unguis-catae, bars of sclerenchyma are present in the furrows of secondary
phloem.

17. Cambium is single layered, present in between xylem and phloem and bent towards inner
side along the furrows of secondary phloem.

18. Secondary xylem consists of vessels, tracheids, fibres and xylem parenchyma.

19. Due to the intrusion of the phloem at four places, secondary xylem is ridged and furrowed at
four places.

20. Primary xylem is present close to the pith facing its protoxylem towards the centre. Its
location is just opposite to the patches of primary phloem.

Pith:

21. It is thin walled and parenchymatous.

Secondary Growth:

Formation of four furrows of secondary phloem in the secondary xylem is due to the abnormal
functioning of cambium which was behaving normally sometimes earlier.

At four or more places cambium produces less amount of secondary xylem towards inter side
and large amount of secondary phloem towards outer side. Thus four wedges of secondary

32
phloem are formed. They intrude into the secondary xylem and so xylem cylinder appears ridged
and furrowed.

33
The most interesting anatomical feature in Bignonia of Bignoniaceae is the occurrence of
anomalous secondary structure. Such is as follows:

a. Presence of Phloem Wedges in the Xylem: The young stems which exhibit such type
of structure when mature are provided with a normal ring of vascular bundles. As soon as
maximum growth is reached four furrows at four equidistant points appear in the xylem,
extending almost to the pith. The cambium is situated on the inside of the furrows.
b. Presence of Fissured Xylem: The fissured xylem may only be visible in fairly old stems.
First of all wedges of phloem are produced and thereafter the xylem strand becomes
fissured by dialation and cell division in wood parenchyma and pith.
C. Accessory cambium formation and its activity and medullary bundles:

In the stems of Bougainvillaea, and other members of Nyctaginaceae (viz. Boerhaavia

diffusa, Mirabilis, etc.), many cambia arise successively in a centrifugal direction. Every
cambium produces xylem and conjunctive tissue to the inside, and phloem and conjunctive tissue
to the outside. The resulting tissue gives the appearance of concentric rings of vascular bundles
embedded in conjunctive tissue.

Secondary Growth in Boerhaavia diffusa

This anomalous type of secondary growth in thickness takes place by means of evolution of the
successive rings of collateral vascular bundles.

In Boerhaavia, the anomalous secondary thickening exists in the form of succession of lings of
vascular bundles. In the members of family Nyctaginaceae the secondary vascular bundles
remain embedded in parenchymatous, prosenchymatous or lignified conjunctive tissue as the
case can be.

Anatomy of Boerhaavia – Stem (Family – Nyctaginaceae):

Epidermis:

1. Single layered epidermis consists of small, radially elongated cells.

2. Multicellular epidermal hairs arise from some cells.

3. A thick cuticle is present on the epidermis.

4. Some stomata are also present.

Cortex:

5. It is well differentiated and consists of few layered collenchymatous hypodermis followed by

chlorenchyma.

34
6. Collenchyma is 3 to 4 cells deep, but generally near stomata it is only one layered.

7. Chlorenchyma is present inner to collenchyma in the form of 3 to 7 layers.

8. Chlorenchymatous cells are thin walled, oval, full of chloroplasts and enclose many
intercellular spaces.

9. Endodermis is clearly developed and made up of many, tubular, thick-walled cells.

Pericycle:

10. Inner to the endodermis is present parenchymatous pericycle but at some places it is
represented by isolated patches of sclerenchyma.

Vascular System:

11. Vascular bundles are present in three rings. In the innermost ring are present two large
bundles; in the middle ring the number ranges from 6 to 14 while the outermost ring consists of
15 to 20 vascular bundles.

12. Vascular bundles of innermost and

middle rings are medullary bundles.

13. Vascular bundles are conjoint, collateral

and endarch.

14. Two vascular bundles of the innermost

ring arc large, oval and lie opposite to each
other with their xylem facing towards
centre and phloem outwards.

15. Middle ring consists of 6-14 small

vascular bundles.

16. Vascular bundles of inner and middle

rings may show a little secondary growth.

17. Phloem consists of sieve tubes,

companion cells and phloem parenchyma
while the xylem consists of vessels,
tracheids and xylem parenchyma.

35
18. Outermost ring of the vascular bundles contain inter-fascicular cambium which is absent in
other two rings.

19. Cambium develops secondarily from the pericycle and becomes active. It cuts secondary
phloem towards outer side and secondary xylem towards inner side. Due to these changes the
primary phloem becomes crushed and present next to pericycle. Primary xylem is situated near
the pith.

20. Interfascicular cambium also soon becomes active and cuts internally the row of cells which
become thick walled and lignified and are known as conjunctive tissue.

Pith:

21. It is well developed, parenchymatous and present in the centre.

Secondary Growth in Dracena

Normally the vascular bundles of the monocotyledonous stems are closed ones. Thus due to
absence of the cambium” they lack secondary growth in thickness and the vascular system is
wholly composed of primary tissues.

Some monocotyledons belonging

to the family Liliaceae, mainly the
arborescent ones like Dracaena,
Yucca, Cordyline, Agave, Aloe and
others exhibit a peculiar type of
secondary increase in thickness.

Anatomy of Dracaena Stem:

The cross-section of Dracaena stem

shows the following structure. The
single layered epidermis remains
covered with thick cuticle. The
lenticels are also visible on the
epidermis. Beneath the epidermis
and hypodermis, the cork cambium
arises which produce the cork
towards outside. Below the cork
cambium, well developed parenchyma is present.

36
Anomalous Structure:

Dracaena shows anomalous secondary growth. In the regions which have ceased to elongate
some cells occurring outside the vascular bundles become meristematic and form the cambium.
The secondary meristem originates in the cortex, in fact, deep layers of cortex or pericycle,
though it is difficult to locate those zones in many stems.

The cambial cells are fusiform or rectangular in shape. Instead of forming phloem and xylem on
the outer and inner sides, as in normal condition, the cambial cells go on dividing and producing
secondary tissues on the inner side first, and later small amount of new tissues are cut off on the
outer side as well.

Those formed on the inner side differentiate into oval-shaped vascular bundles and radially
arranged parenchyma cells. These parenchyma cells in which the vascular bundles remain
embedded are said to constitute the conjunctive tissue.

The radial arrangement of the parenchyma cells of conjunctive tissue is due to their origin by
tangential divisions of the cambial cells. So they may be easily distinguished from the irregularly
arranged parenchyma of the primary ground tissues. They may be thin-walled or thick- walled.

A few derivatives of the cambium divide longitudinally, initially anticlinally, then periclinally
and even haphazardly to form xylem and phloem elements of the secondary vascular bundles.
These differ from the primary ones in presence of small amount of
phloem and in absence of annular and spiral protoxylem elements.

The secondary bundles are mostly amphivasal, some of them may be collateral as well. The
small amount of phloem consists of short sieve tubes, companion cells and parenchyma. The
xylem is made of only tracheids, usually with scalariform thickening and small amount of xylem
parenchyma which have lignified walls.

The tissues cut off by the cambial cells on the outer side are scanty in amount and are
parenchymatous in nature. The primary bundles are comparatively larger. They are also mostly
amphivasal or rather rarely collateral ones.

37