The vascular cambium is a lateral meristem that develops either as a longitudinal

or as a hollow cylinder. The vascular cambium produces the secondary xylem and
secondary phloem.
Origin of Vascular Cambium:
The vascular cambium is develops from the Procambium.
There is a sequence of differentiation of procambium from the apical meristem of
the axis to a single cylinder of vascular bundles. The vascular differentiation is
detectable above the level of leaf or leaf initiation. This is called Promeristem.
Beneath the insertion of leaf primordial, the longitudinal divisions without
considerable increase in width of cells produce somewhat elongated cells. n this
way, with the progressive differentiation of the promeristematic cells, three
meristems!the Protoderm, Ground Meristem and Procambium are recogni"ed.
The procambium may be in the form of a solid or hollow cylinder or strips.
#evelopment of procambium is acropetal. The differentiations of procambial cells
into the cells of vascular tissue ta$e place in different plants at different stages of
procambial development.
Development on Cambium:
n certain plants %including the monocotyledons& all the cells of procambium
undergo differentiation into primary vascular tissues. But in dicotyledons and
gymnosperms, a portion of procambium remains meristematic even after the
completion of the primary growth and developed into the cambium of secondary
body.
The cambium that arises within the bundle of the primary vascular tissue of stem is
called Fascicular cambium.
The strips of fascicular cambium usually become 'oined by additional strips of
cambium which form the Interfascicular cambium.
The nterfascicular cambium develops from nterfascicular parenchyma. Thus a
complete hollow cylinder of cambium is developed. The cells cut off from the
cambium on its inner face become secondary xylem. (nd those which are cut off
from its outer face develop into secondary phloem. The region of vascular
cambium in which cell divisions are actively ta$ing place called cambial zone. t
is composed of many layers of xylem and phloem. The cells that are actively
dividing in cambial "one are called cambial initials.
The cambium is usually consists of two types of cells.
(a). Fusiform initials. (b). Ray initials.
Fusiform initials:
These are elongated cells with tapered ends. They are long and give rise to all cells
of xylem and phloem.
!rrangement of fusiform initials:
Two types of cambium can be distinguished on the basis of arrangement of
fusiform cells.
"torie# cambium: n this type, the fusiform cells are arranged in hori"ontal
rows. )o their ends are approximately at the same level. This type is common
in plants with short fusiform initials e.g. in Tamarix.
$onstorie# cambium: n this type, the fusiform initials partially overlap one
another and are not arranged in hori"ontal rows. t is found in plants with long
fusiform initials. t is found in the Pteridophytes and living *ymnosperms.
Cell #ivisions in Fusiform:
There are four types of cell divisions in fusiform initials.
%roliferative #ivisions: +usiform initials divide with longitudinal, periclinal
walls to produce new derivatives. This division is called proliferative
divisions. The cells which are produced by this division ma'ority of them
become xylem mother cells and few are phloem mother cells.
&ultiplicative #ivisions , (s the xylem matures inside the vascular cambium,
it expands and the cambium is pushed outward and its circumference is
increased. The fusiform initials divide with longitudinal, anticlinal walls to
produce more initials rather than the derivatives. t is called multiplicative
divisions.
%seu#o transverse #ivisions , The multiplicative divisions in non-storied
cambium are called pseudo transverse divisions.
Ray Initials:
These are short cells. They form the transverse or ray system of wood.
!rrangement of Ray initials:
The ray initials are small than the fusiform initials. n conifers, they are
arranged in uniseriate groups and produce uniseriate rays. n some cases, the
multiseriate condition is also found. There are following ways by which the
ray initials are form in the cambium.
( single cell may be cut off the side of fusiform initials.
( single cell may be cut off the end of fusiform initials by a special
transverse division.
( declining fusiform initial may be reduced to a single ray initial.
.n the other hand, ray initials can begin to elongate and convert themselves
to fusiform initials.
Cambial activity:
The vascular cambium shows great variation in the period and intensity of activity.
These variations are the result of many internal and external factors.
"easonal activity:
n tropical regions, many trees do not become dormant and their cambial activity
remains occur during the entire year.
But in sub-tropical and temperate regions, the cambium stops its activity with the
beginning of the un-favorable conditions usually in autumn. t remained inactive
during summer. n coming spring, the cambium becomes active again.
n dormant state, it stops all cell divisions and many of xylem mother cells and
phloem mother cells mature. n spring, as the division resumes, the first cells to
divide/reactivated are those which are below the swelling buds.
The vascular cambium remains dormant during the times of the stress such as
drought, high and low temperature. n arid regions, the availability of water in the
soil is important factor in the control of cambial activity. The cambium become
active when there is plentiful water and become dormant during the period of water
shortage.
Thus the intensity and amount of cambial activity varies in different seasons and
climates.
"econ#ary 'ylem:
)econdary xylems are produces toward the inner side. t pushes the primary xylem
bundles towards centre. 0ltimately, the pith is completely filled with the secondary
xylem and primary xylem.
)econdary xylem consists of axial system and ray system.
!(ial system: The axial or vertical system comprises the tracheary elements, fibers
and wood parenchyma.
Ray system: The ray or hori"ontal system is composed of xylem rays.
Both are interpenetrating systems closely integrated with each other in origin,
structure and function. The living cells of both systems are interconnected with
each others.
"econ#ary (ylem of )ymnosperms:
a). !(ial system:
n large ma'ority of conifers, the axial system is composed of only tracheids.
1owever, some conifers develop fiber tracheids at the end of growing season.
2ibriform fibers are altogether absent in conifer wood and therefore called as
)oftwood.
b). Ray system:
The ray system of gymnosperms is composed of parenchyma cells.
n gymnosperms, rays are commonly one cell wide %uniseriate&. The presence of
resin ducts in a ray ma$es the ray more than one cell wide. )uch rays are called
fusiform rays.
c). Formation of *oo#:
The secondary xylem accumulates. t forms wood. 3ood has two parts.
"ap *oo#: The narrow, peripheral functional part of the secondary xylem is called
sap wood.
+eart *oo#: The ma'or central part forms the heart wood. t is non-functional.
)ometimes, the outgrowth is form which is called Tylosis.
!nnual rings: n spring, the cambium is more active. t produces wide and loosely
elements. t is called spring *oo#. The activity of cambium slows down during the
autumn. t produces narrow and closely pac$ed tracheal elements. t forms autumn
*oo#. The difference b/w the spring wood and autumn wood produce the (nnual
rings. These rings can be seen in the T.) of wood.
"econ#ary (ylem of Dicot *oo#:
a). !(ial system:
0nli$e the conifers, dicot wood typically contains the parenchyma cells in axial
system. These derived from fusiform initials. There are two basic types of axial
parenchyma.
!potrac,eal parenc,yma: The parenchyma is typically independent of vessels.
%aratrac,eal parenc,yma: The parenchyma is associated with the vessels.
These are further subdivided into followings.
-ypes of !potrac,eal parenc,yma:
i). Diffuse# parenc,yma: f the apotracheal parenchyma cells occur as isolated,
uniseriate bands or in the form of single cells scattered irregularly among the wood
fibers they are called diffused parenchyma.
ii). .an#e# parenc,yma: f they form concentric bands or clusters they are called
banded parenchyma.
-ypes of %aratrac,eal parenc,yma:
i). "canty paratrac,eal parenc,yma: n this type, the paratracheal parenchyma
does not form a complete sheath around the vessel.
ii). /nilateral paratrac,eal parenc,yma: n this case, the paratracheal
parenchyma located on only one side of vessels.
iii). Vesicentric parenc,yma: n this case, parenchyma surrounds the vessels
completely
b). Ray system:
The dicot typically contains only cells in rays. .n the basis of orientation of the
long axis of the cell, there are two types of ray parenchyma cells. %rocumbent
%square& and /prig,t %vertically elongated&. f all the cells in rays are procumbent,
the rays are called +omogenous and when the rays consist of upright and
procumbent cells, they are called +eterogeneous. The heterogeneous rays may be
uniseriate or multiseriate.

"econ#ary %,loem:
)econdary phloem also consists of hori"ontal or ray system and vertical or axial
system.
a). !(ial system:
The secondary axial system contains sieve tubes, companion cells and phloem
parenchyma. 4ew phloem elements are added each other. t increases the
circumference only. Therefore, the layer of secondary phloem never becomes very
thic$. The secondary phloem of previous year is peeled off with the bar$ of the
periderm in many plants.
b). Ray system:
The characters of rays are the same as in the secondary xylem.
!bnormal "econ#ary )ro*t,: