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BIO 1201 - Lecture 13

Secondary Growth in Dicot Stems

The growth in length of main axis by the activity of apical meristem is called primary growth.
Increase in thickness or girth of the axis due to the formation of secondary tissue is called
secondary growth. The secondary tissue is formed by the activity of cambium and cork-
cambium. 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:
a) Formation Of Cambium Ring
Formation of the cambium ring is the first step of secondary growth. The cambium of vascular
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 vascular 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 isidiametric which give rise to vascular rays of
parenchymatous cells.
b) Formation Of Secondary Vascular 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.

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

Secondary Xylem: Secondary xylem consists of vessels, tracheids, wood fibres and wood
parenchyma. The vessels or trachea are most abundant in secondary xylem and are usually
shorter than that of the primary xylem. The cambium ring forms more tissue on the inner side
than on the outer side. As a result, secondary xylem forms the main bulk of the plant body and is
generally called the wood. Its width increases with age. The primary xylem persists as conical
projections towards the pith.

c) Vascular 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.
d) Annual rings: The activity of vascular cambium is greatly affected by the variations in
the climate. It is more pronounced in temperate regions. The cambium stops dividing in
winter. In the spring season or early summer, the cambium becomes more active and
produces a large number of vessels with wider lumen. These are called spring wood or
early wood.
During the autumn or winter season, the cambium becomes less active and produces vessels with
narrow lumens. Tracheids and wood fibres are formed in large numbers. These woods are called
autumn wood or late wood. Hence, the annual rings are formed year after year. In the oldest part
of the tree, annual rings can be used in determining the age of a tree.

In tropical regions, the climate is more or less uniform. Therefore, the annual rings are not well
developed and do not correlate with the age of tree.

e) Sapwood and Heartwood:

In older stems, the woody trunk is differentiated into two regions. The outer light coloured
region is called sap wood or alburnum and central dark colored region is called heart wood or
duramen. The cells of sapwood are living and functional. They take part in conduction of water
and storage of food. The heartwood consists of dead cells. During the growth process, the rings
of sap wood gradually process, the rings or sap wood gradually convert into heartwood. The
living cells of sap wood lose their protoplast and water content. The lumen of the xylem vessels
get blocked by the ingrowth of the parenchyma cells. The adjacent parenchyma tissue enters
through the pits of vessels and gradually enlarges to form a balloon like structure, which is called
tyloses. The heartwood is stronger and more durable than sapwood. The heartwood becomes
resistant to the attacks of bacteria and fungi due to the presence of antiseptic oils.

Secondary Growth in Dicot Roots

At completion of primary growth (i.e. growth in length), the secondary growth (i.e. growth in
thickness) is initiated in roots due to the activity of the vascular cambium. Secondary growth
includes the production of secondary phloem and secondary xylem by this lateral meristem.
Roots with secondary growth often become woody due to the accumulation of secondary xylem.
If we examine older thick roots of trees, we observe annual growth rings in the wood (secondary
xylem) resembling those of stems.

The vascular cambium appears first on the inner edges of the phloem strands. While these
cambial cells form some secondary elements, the pericyclic cells outside the protoxylem poles
also divide. The inner derivatives of these divisions complete the cylinder of cambium by joining
the strips located on the inner faces of the phloem strands. The vascular cambium loses its wavy
circular outline in transverse sections because on the inner boundary of the phloem, the
secondary xylem is deposited earlier than outside the protoxylem. The secondary vascular tissues
assume the form of a continuous cylinder and complete enclose the primary xylem. The sieve
elements of the primary phloem are crushed, and some of the remaining cells differentiate into
fibers. The cambium which arises in the pericycle outside the xylem poles forms wide vascular
rays. In a few species, the phloem remains internal to the xylem.

The initiation of phellogen and periderm follows the initiation of vascular cambium and
secondary vascular tissues. Phellogen (cork cambium) originates from the pericycle cells located
opposite primary phloem. The primary tissues located outside the pericycle (rhizodermis,
exodermis, cortex and endodermis) become isolated from the secondary body of the root by
periderm formation and die.

The secondary phloem in roots consists of axial elements such as sieve tubes with companion
cells, phloem parenchyma cells and phloem fibers, and radial elements, i.e. ray cells. The axial
elements of secondary xylem which are deposited by fusiform initials include wood parenchyma,
fibers and vessels of various widths with bordered pits arranged in reticulate or scalariform
patterns in their lateral walls. The rays are usually wide in herbaceous dicots and as narrow as in
stems in woody life-forms. The root secondary xylem typically contains more parenchyma
elements than the stem wood.

Periclinal divisions in the pericycle that are not involved in the formation of the vascular
cambium occur not only outside the xylem poles, but spread around the circumference of the
root. Such divisions are preparatory for the formation of the periderm. A phellogen arises among
the outer cells of the proliferated pericycle which forms cork tissue to the outside and phelloderm
towards the inside. The phelloderm is difficult to distinguish from the parenchyma derived from
the pericycle.

Some dicotyledonous roots may retain their cortex for a period of time during secondary
development. Such roots may develop an exodermis or a superficial periderm, and the cortex
later becomes lost.

In some cases, the endodermis of the parent root takes part with the pericycle in forming branch
roots. Sometimes it forms several cell layers by undergoing both anticlinal and periclinal
divisions. In a few cases (e.g. Cucurbitaceae and certain water plants), the innermost cortical cell
layers are also involved in the formation of the lateral root by contributing cells to the lateral

Adventitious roots may be formed in young organs or in older tissues that have not quite lost
their meristematic properties. Most adventitious roots arise endogenously, sometimes from
primordia laid down previously and remaining dormant until stimulated to growth. Usually the
adventitious roots arise from the pericycle, but may also arise from the cambial zone.

Roots in both primary and secondary states of growth may undergo modifications in their
structure due to the acquisition of special functions. In this sense, storage, aerial, mycorrhizal and
contractile roots as well as roots modified by the interaction with parasitic plants are specially
considered in this unit of study.

Many roots may form a symbiotic association with fungi termed mycorrhizae. In
ectomycorrhizae, the fungal hyphae grow as a mantle around root endings and between the cells
of the rhizodermis and cortex, while in endomycorrhizae the hyphae invade the root cells and the
mantle is poorly developed.

Characteristics of Meristematic Tissues

The cells of the meristematic tissues are quite distinct in their cytological and physiological
characteristics from other cells. The following are the characteristics of the meristematic cells.

1. Cells have the power of active division.
2. They are compactly arranged in tissue and there is absolutely no intercellular space.
3. The cell wall is thin and primary in nature, containing only cellulose. It is uniformly thick.
There is no secondary thickening.
4. Cells possess dense protoplasm with a prominent large nucleus compared to other cells of
equal volume.
5. Vacuoles are small or absent totally.
6. Cells do not possess ergastic sub-stances. Ergastic substances are non-protoplasm materials
found in cells.
7. Mitochondria and endoplasmic reticulum are very little differen-tiated.
8. Plastids, when present, are in protoplastid stage.
9. Cells contain relatively more number of ribosomes.
10. Cells are metabolically very active.

When cells of the meristem divide, the daughter cells get differentiated into mature types while
the others re-main meristematic. For this reason, the meristems perpetuate and become
con-tinuous source of cell formation.

Function of the Vascular Cambium

Vascular Cambium produces secondary growth, giving rise to secondary xylem and secondary

The vascular cambium is a lateral meristem in the vascular tissue of plants. It is a cylinder of
unspecialized meristematic cells that divide to give rise to cells that further divide, differentiate,
and specialize to form the secondary vascular tissues.

The vascular cambium usually consists of two types of cells: Fusiform initials (tall cells, axially
oriented); Ray initials (almost isodiametric cells, smaller and round to angular in shape).

Secondary xylem is the xylem that is formed during secondary growth from vascular cambium.

Phloem is the living tissue that carries organic nutrients (known as photosynthate), in particular,
sucrose, a sugar, to all parts of the plant where needed.

The phloem is concerned mainly with the transport of soluble organic material made during


A vascular tissue in land plants primarily responsible for the distribution of sugars and nutrients
manufactured in the shoot.


A vascular tissue in land plants primarily responsible for the distribution of water and minerals
taken up by the roots; also the primary component of wood.

Anomalous Secondary Growth

Anomalous secondary growth does not follow the pattern of a single vascular cambium
producing xylem to the inside and phloem to the outside. Some dicots have anomalous secondary
growth, such as Bougainvillea, where a series of cambia arise outside the oldest phloem. Most
monocots either have no secondary growth or else anomalous secondary growth of some type.
For example, palm trees increase their trunk diameter due to division and enlargement of
parenchyma cells, which is termed "diffuse secondary growth." In some other monocot stems
with anomalous secondary growth, a cambium forms, but it produces vascular bundles and
parenchyma internally and only parenchyma externally. Some monocot stems increase in
diameter due to the activity of a primary thickening meristem, which is derived from the apical
Formation of Cork Cambium
Cork originates from a layer of cambium (=phellogen) that itself is formed as a secondary
meristem from a layer of collenchyma or parenchyma immediately beneath the epidermis. In
contrast to sclerenchyma cells, collenchyma cells are alive and they have retained the potency to
de-differentiate. Cork cambium cells only divide periclinally so that the typical rows of daughter
cells arise: cork cells (=phellem) are mainly generated toward the outside. In a lesser extent also
cork parenchyma (=phelloderm) is made toward the inside. Cork cambium, cork cells and cork
parenchyma together are also named periderm. Mature cork cells are dead; their cell walls
contain suberine, a fatty substance that repels water. The layer of cork provides protection
against desiccation, but it also isolates tissues in the inner parts of the stem or trunk so
thoroughly that exchange of gas with the outer world is impeded. 'Breathing' is yet achieved by
so-called lenticels. The cork cambium starts to generate numerous parenchyma cells toward the
surface, in most cases at the level of a stoma. These thin-walled parenchyma cells, which
eventually degenerate, force an interruption of the sealing created by the cork layer and focus an
opening for gas-exchange.
Lenticel formation
Lenticels originate beneath stomata, simultaneous with the initiation of the first layer or periderm
or just before the initiation of periderm, during first growing season. As the lenticels formation
starts, the parenchyma cells found near substomatal cavity lose their chlorophyll content and
irregularly divide in different plants giving rise to a mass of colourless, rounded, thin walled,
loosely arranged cells called Complementary cells. Sometimes complementary cells produced by
phellogen towards outside instead of producing cork cells.
As the complementary cells increase in number, pressure is exerted against the epidermis and it
ruptures. Outer most cells gradually become dead and may be replaced. Beneath the outer layer
some mass of closely packed cells alternate to loosely arranged cells are formed called as
Closing layer. From inner side continuous production of complementary cells cause rupture of
closing layer at intervals.
i) Lenticels contain profuse inter cellular spaces for which it perform the function of exchange of
gases between the atmosphere and internal tissue of the stem.
ii) Lenticels like Stomata help in transpiration. Called Lenticelar transpiration.
iii) Lenticels are active during night when stomatal transpiration stops.
The periderm is the secondary protective (dermal) tissue that replaces the epidermis during
growth in thickness of stems and roots of gymnosperms and dicotyledons (i.e., secondary
growth). Unlike the epidermis, the periderm is a multilayered tissue system, the bulk of which
usually constitutes the cork, or phellem. There are, however, some exceptions to this inasmuch as
some other structures (e.g. potato skin and apple peel) are also periderm. The lateral meristem,
(cork cambium or phellogen), is one cell layer thick and encircles the stem. It produces periderm
centrifugally. The layer of cork cells formed is impermeable for water and gases, but is
interrupted at certain points by lenticels which function to some extent similar to stomata in the
epidermis, and permit gas diffusion. In some cases parenchyma cells are produced centripetally
(i.e. to the inside of the stem or root) by the phellogen as a part of the periderm. These persistent
living cells are called phelloderm and structurally appear similar to cells of the cortex. The
number of layers of cork and phelloderm varies greatly among different species; some plants
produce no phelloderm. The most important function of the periderm is to reduce the loss of
water and solutes from interior tissues and to protect a plant from unfavorable environmental