Secondary Growth In Dicots

By
Sumayya Abdul Rahim
Department of Botany
Marthoma College, Thiruvalla
 Introduction

• In higher plants, there are two separate growth stages and each of them play a significant role in the plant's life.
• The first series of growth involving an increase in the length of roots and stems from the apical meristems is called
primary growth.
• The growth taking place later on, succeeding the primary growth, which is responsible for the increase of girth or
width is termed as secondary growth.
• This growth is initiated by the two lateral meristems also known as vascular cambium and cork cambium.
• Normally secondary growth takes place in roots and stems of dicotyledons and Gymnosperms.
• Secondary growth begins right after the apical meristems have completed the primary growth process with all
parts developed properly.
• Secondary growth provides the plants or trees with protection against mechanical stress and microbial activity.
• Secondary growth increases the girth of the stem.
• Due to lack of cambium in monocotyledons, secondary growth is absent. But exceptionally secondary growth has
been reported in some monocotyledons. Such as - Palm, Yucca, Dracaena, Smilax, Agave, Coconut etc.
Stages of secondary growth in a typical dicot stem

• Secondary growth in dicot stem can be categorised as-

I. Intrastelar secondary growth- It involves the formation of vascular cambium

in the stelar region and its activity resulting in the formation of secondary xylem,
secondary phloem and secondary medullary rays

II. Extra stelar secondary growth- It involves the formation and activity of cork
cambium in the extra stelar region leading to the formation of cork and secondary
cortex.
(I) Intra stelar Secondary growth:
•The growth in this region starts earlier than the extra stelar region.
A. Formation of ring of vascular cambium:

Stage 1
Vascular bundles of dicot stems are conjoint (xylem and phloem in one bundle), collateral (laterally placed on the same
radius) and open (fascicular cambium present between xylem and pholem), arranged in a ring.
Single layer of fascicular cambium is present between primary xylem and primary phloem. There is no cambium across the
pith rays/ medullary rays.

Stage 2
After differentiation of xylem and phloem, the paranchymatous cells of pith rays between the edges of fascicular cambium
become meristematic and connect to the fascicular (vascular) cambium.
Thus, a complete ring of cambium (intra-fascicular cambium and inter-fascicular cambium) is formed.
B. The activity of the cambial ring
• Two types of cells are found in the ring of this vascular cambium.
(i) Fusiform initials (form secondary xylem and phloem)
(ii) Ray initials (form secondary medullary rays)

• Fusiform initials are long with pointed ends, while ray initials are isodiametric cells. Activity of fusiform initials is
more in vascular cambium.

I. Activity of fusiform initials:

• Periclinal divisions occur in fusiform initials, as a result few cells are formed towards the radius (periphery)
differentiated into secondary phloem or bast and some of the cells are formed towards the central axis and these cells
are differentiated into secondary xylem or wood.
• Secondary xylem is formed 8-10 times more as compared to secondary phloem.
• By the pressure of secondary phloem, primary phloem is pushed towards the outside and gets crushed.
• Primary xylem however remains more or less intact, in or around the center.
II. Activity of ray Initials:
• The ray initials give rise to a narrow band of parenchymatous rays in radial direction known as medullary rays.
• The secondary medullary rays or vascular rays are often rows of radially arranged cells which pass through the
secondary xylem and secondary phloem.
• The primary and secondary rays conduct food,water and minerals from the center to the periphery of the organ in
radial
• The secondary xylem of the tree trunk is of great economical value, since it constitutes the timber and wood of
commerce.
II. Extra Stelar secondary growth in dicot stem (Periderm Formation)
Periderm Formation

• Extra stelar secondary growth is provided by a specific lateral meristem called the cork cambium.
• Due to secondary growth in vascular bundles, the girth of the stem keeps on increasing which leads to damage of the
epidermis and cortex.
• In order to replenish those layers, the cork cambium is necessary.
• Because the cork cambium lies right outside the stele it is known as extra-stelar cambium.
• Cork cambium or Phellogen is multi-layered and thick, it is composed of narrow rectangular cells which are thin-
walled.
• Cork cambium usually originates from the outer layer of cortex because the latter becomes meristematic.
• Cork cambium is formed in the form of a single layered ring.
• It forms secondary tissues in the cortical region.
• Phellogen or cork cambium layer divides and produces cells on either side. The cells that formed on the outer side get
differentiated into cork or phellem. Whereas, the cells formed on the inner side differentiate into the secondary cortex
or phelloderm.
• The cork cells are dead due to deposition of suberin in their middle lamella and are impervious to water.
• Phelloderm is made up of thin-walled parenchymatous cells which are living and exhibit cellulosic cell walls.
• Phellogen, Phellem, Phelloderm are altogether known as Periderm.
• All these layers contribute to the protective layers of the epidermis and cortex after they get damaged due to secondary
growth of the vascular bundles.
• As the cork cambium adds up cells into the phellem and phelloderm, the cells peripheral to phellogen die and fall off.
Bark
• All the tissues that exist peripheral to vascular cambium are considered as a part of bark.
• Bark is not a scientific term, it can represent various tissues for example secondary phloem, periderm, primary cortex,
pericycle etc.
• The dead tissue present outside the cork cambium is generally called outer bark & the tissues present between vascular
and cork cambium constitutes the inner bark.
• All the dead tissues formed outside the innermost cork cambium are called rhytidome.
• The early formed bark, i.e., at the beginning of a season is known as soft bark.
• The later formed bark, i.e., at the end of a season is known as hard bark.

Lenticels
• At places, the phellogen cuts off closely packed parenchyma cells on the
outer side instead of the cork.
• These cells rupture the epidermis and form a lens-like opening on the
outer surface of the stem.
• These openings are known as lenticels.
• These structures are prominently found on woody trees.
• Function: Lenticels mainly perform the function of exchange of gases
between plants and the atmosphere. They are also responsible for
lenticular transpiration
 Secondary growth in dicot root

• The dicot root also exhibits secondary growth.

• The increase in width of a dicot root takes place due to two
different lateral meristems, i.e., vascular cambium and cork
cambium.
• The vascular cambium and cork cambium are formed as a result
of secondary growth.
• The process of secondary growth in dicot roots is accomplished
in two steps,
a) Formation and activity of the vascular cambium
b)Formation and activity of the cork cambium
A. The Formation and activity of the vascular cambium
• The vascular bundles are arranged in a radial manner.
• The xylem and phloem are distinct from each other in separate patches.
• The xylem is of exarch type.
• The vascular cambium forms as a result of secondary growth hence it is not present right from the beginning.
• Conjuctive tissue becomes meristematic below phloem bundles and form separate curved strips of vascular cambium during the secondary
growth in a dicotyledon root.
• Now the cells of the pericycle lying opposite to protoxylem also become meristematic to form additional strips of cambium.
• In this way a complete ring of vascular cambium is formed due to joining of these two.
• The shape of the ring of vascular cambium is wavy in the beginning, but later on it becomes circular due to the pressure of the secondary
xylem.
• The activity of vascular cambium of root is similar to activity of vascular cambium of stem.
• Vascular cambium forms secondary xylem towards the inner side and secondary phloem towards the outer side.
• The portion of vascular cambium which is formed by the pericycle is responsible for the formation of pith rays. These are made up of
parenchyma. These pith rays are known as primary medullary rays (Multiseriate).
• A few medullary or pith rays are also formed from remaining vascular cambium. These are called secondary medullary rays (uniseriate).
Thus two types of medullary rays are found in the secondary structure of roots

B. The formation and activity of the cork cambium

• The pericycle cells divide to generate the cork cambium.
• Thereafter the cork cambium gives rise to the periderm, which is responsible for replacing ruptured epidermis and cortical layers.
• The cork cambium in root is a lot similar to the cork cambium in the stem. This cambium forms a dead cork towards the periphery and the
living, parenchymatous secondary cortex towards the centre.
• Pressure builds upon the peripheral layers of the phellogen, that is the epidermis and the cortical layer as a result they die and finally
slough off.
Other terms to remember

• Polyderm refers to a special type of protective tissue consisting of a uniseriate suberized layer alternating with
multiseriate non-suberized cells in periderm. It is found in the roots and underground stems. Example: Rosaceae

• Rhytidome is a technical term used for the outer dead bark which consists of periderm and isolated cortical or phloem
tissues formed during successive secondary growth.
Example: Quercus
Anomalous Secondary Thickening in Bignonia
• Bignonia is a member of the Bignoniaceae family.
• The young stem is wavy in outline with prominent ridges and furrows.
• The most interesting anatomical feature in Bignonia is the occurrence of anomalous
secondary structure.
T. S. shows many ridges and furrows and reveals the following tissues
Epidermis:
1. Single-layered epidermis consists of rectangular cells.
2. A thick cuticle is present.
3. A few multicellular hairs also arise from some cells.
Cortex:
4. It is well-differentiated into collenchyma and parenchyma.
5. Collenchyma is present below the epidermis in the ridges in the young stem but at maturity
develops sclerenchyma.
6. Parenchyma is present below the sclerenchyma or collenchyma in the ridges and directly
below the epidermis in the grooves.
7. In the old stem cortex consists of a cork, cork cambium, and cortex.
Endodermis is undistinguishable from cortical cells. The cells lack Casparian strips.
Pericycle is heterogenous and the sclerenchyma is present below the ridges.
Pith is prominent and vascular bundles are conjoint, collateral, and arranged in a ring.

Secondary thickening (important)

• Anomalous secondary thickening is due to the abnormal functioning of the
cambium. Cambium is normal in disposition and abnormal in function (adaptive
type or adaptive anomaly).
• During secondary thickening inter and intra-fascicular cambium becomes active to
form a normal cambial ring.
• In the beginning, its activity is normal i.e. it forms more of secondary
• xylem on the inner side and less amount of secondary phloem on the outer side.
• After some time, at four diagonal places the cambium forms internally lesser amount
of xylem and on the outer side greater amount of phloem.
• It results in the formation of four deep wedges of phloem projecting into the xylem.
• Thus, four deep wedges of secondary phloem are supported by four transverse bands of
sclerenchyma cells and four ridges of secondary xylem are formed.
• The sclerenchyma bars give mechanical support to thin-walled phloem tissues
• The cambium is situated on the inside of the furrows and on the outside of the ridges,
while the radial surface is occupied by parenchymatous medullary ray tissues.
• Periderm formation is normal as the activity of the cork cambium is normal
• Bignonia is a woody climber; this anomaly is thus an adaptation to the climbing habit
of the plant. Hence this anomalous secondary growth in Bignonia stem is described as
Adaptive anomalous secondary growth