P LA N T H IS TO LO G Y A N D

A N ATO M Y
Introduction
Histology is a branch of biology that deals with the study of
tissues.
Tissue is a group of similarly specialized cells having
common origin, structure and function.
Anatomy is a branch of biology that deals with the study of
internal structure of organ and organ systems.
As we know that all organisms are made up of cells and
their organization starts from Acellular, Cellular, Tissue level,
to the higher organ system level.
In plants, based on the capacity of division/ growth, the
plant tissues are classified in to two major groups, namely
the Meristems and the Permanent tissues.

Meristems

Meristematic tissue (meristos- divide)

Meristematic tissue may be defined as a
group of young undifferentiated cells, capable
of active divisions to produce new cells,
present in the regions of growth. Cells are
compactly packed with thin cell wall made
from Cellulose, cytoplasm is dense without
vacuoles, nucleus is prominent, plastids and
ergastic substances are absent.
They bring about growth, differentiation and
involved in the healing process.

Based on the position, they are classified in

to
Apical meristems: They are found at the
tips/apices of roots, shoots and bring about the
increase in vertical length (primary growth).
Intercalary meristems: They are found in inter
nodal regions (as in grasses) or at the leaf bases
(as in Pinus). It is also responsible for primary
growth. E.g. Bamboo, Sugar cane etc
Lateral meristems: They are found along the
lateral sides of the permanent tissues in the stem
and roots of dicotyledons and gymnosperms.
They are absent in monocots. They are
responsible for increase in girth of the organ
(Secondary growth). Vascular and Cork cambium
are the examples.

Based on the origin, they are classified in

to
Promeristems/ Primordial/ Embryonic
meristems: These are earliest, embryonic
meristematic cells occupying a small area and
give rise to new tissues and organs.
Primary meristems: These meristems are
derived from the promeristems and retain the
capacity to divide through out life of the plant.
They persist through out life. They are found at
the tips of roots, stems and primordia of leaves.
Secondary meristems: these are derived from
primary permanent tissues by a process called
dedifferentiation. It takes place during
secondary growth.

Based on the function, they are

classified in to
Protoderm meristem: it is the outer most
layer of the Apical meristem; in future
differentiate to form epidermal tissue
system.
Ground meristem: it lies between the
protoderm and procambium. It forms
ground tissue.
Procambium: it is a part of apical
meristem develops in to primary
vascular tissues, namely xylem, phloem
and cambium.

Permanent tissues
Permanent tissues may be defined as a group of
cells, derived from meristems after differentiation,
growth and maturation, characterized by definite
shape, size and function.

Classification of Permanent tissues

Based on the cell types the Permanent tissues are
classified in to Simple Permanent tissues and
Complex Permanent tissues.
Simple Permanent tissues: SPT as group of similar
cells having common functions. They are further
classified in to three types:
Parenchyma
Collenchyma and
Sclerenchyma.

Parenchyma: it is the ground/ fundamental/

living tissue generally found in soft parts of
the plants of all categories.
Distribution: Soft regions such as cortex and
pith of root and stem, mesophyll of leaves,
fleshy parts of fruits, endosperm of seeds etc.
Structure: They are thin walled, iso-diametric,
shape variable, with definite nucleus and
vacuolated cytoplasm and certain large
amount of food materials.

Classification of Parenchyma: Based on the cell

structure and function, three types of
parenchyma are noticed-

 Simple parenchyma: Ground tissue possesses all

the characters as above. These cells show

essential metabolic functions like respiration,
secretion, excretion and also storage.
Idioblasts: Parenchymal cells containing mineral crystals

like Raphides (Oils, crystals of Calcium oxalate).

Chlorenchyma: Parenchymal cells containing

chloroplasts usually found in the cells of leaves

and some times the cells of stem cortex.
Palisade parenchyma: these are elongated

chlorenchymal cells as in leaves.

Spongy parenchyma: These are irregularly shaped
chlorenchymal cells with large intercellular spaces.

Aerenchyma: The parenchymal cells with large

intercellular air filled spaces (Helps in Buoyancy

and exchange of gases), usually found in aquatic,
submerged and very few land plants.
Prosenchyma: slightly thick walled elongated parenchymal

Functions of Parenchyma:
They take part in the vital functions like photosynthesis,

respiration, secretion etc.

They are involved in the storage of Starch, Proteins, Crystals

etc.
They are involved in the storage of water as in Xerophytes

(Opuntia, Aloe etc.).

Collenchyma: It is a simple living permanent tissue

found in the hypodermal regions of the plant body
Distribution:
present
below
the
epidermis
(hypodermal region). It also occurs in leaves on
either sides of vein and normally absent in leaf
mesophyll underground storage.
Structure
Cells living appear cylindrical in vertical section
and oval, globular or polygonal in cross section.
More compactly packed, nucleus at corners and
vacuolated cytoplasm. Cellwall uniformly thin
except at corners, where in corners, the secondary
wall is thickened with cellulose, hemi cellulose or
pectin.

Types of Collenchyma:
Based on the Pectinization and arrangement of cells the
Collenchyma is classified in to three types
Angular Collenchyma: Cells are irregularly arranged, compactly
packed and only the corners of the cells are pectinised. E.g. Datura,
Ficus, Morus etc.
Lamellar Collenchyma: Cells are compactly packed and pectin is
deposited layer by layer in Collenchyma. E.g. Sunflower.
Lacunar Collenchyma: Cells are with small intercellular spaces and
pectin gets deposited in there spaces rarely. E.g. Malva, Salvia,
Hypodermis of cucurbita

Functions
It functions as primary mechanical tissue and give support to
young aerial parts of dicots particular Petioles, Pedicel and fruit
stalk.
Like parenchyma it also takes place in wound healing process.
Collenchyma contains chloroplasts that take part in photosynthesis.
It protects the vascular bundles of leaves by forming Bundle caps.
It gives support to the developing organs of the plant.

Sclerenchyma: It is simple dead mechanical tissue,

composed of dead cells at maturity and hence it is
mechanical. The cells of Sclerenchyma are thickened
uniformly and often lignified.
Sclerenchyma consists two types of cells
Sclerenchyma fibers: the fibers are spindle shaped
dead cells with the pointed ends. The cell wall is
lignified and thick. Cells are arranged vertically,
compactly packed and purely mechanical in function.
In cross section the fibers appear mostly polygonal in
out line and show highly lignified walls with pits at few
places. During development, plasmodesmata are
blocked due to lignification.
Occurrence: They are found below the epidermis,
around the vascular bundle in monocot stem, pericycle
of dicot stem. Also present in xylem, phloem, cortex,
pith mesocarp of fruits (E.g. Cocos nucifera, Mango) and
seed coat of Cotton.

Based on the distribution the Sclerenchyma fibers are

recognized in to two types
Intra Xylary -Associated with Xylem and originate
from the same meristematic cells along with the
xylary elements, tracheid vessels and parenchyma.
Extra xylary - These fibers include other than the
fibers associated with Xylem. The fibers associated
secondary phloem, forming a part of pericycle
(Bundle cap) in dicot stem.

Functions:
Sclerenchyma fibers give support, strength and
rigidity to the plant (Mechanical in support).
Sclerenchyma fibers are of commercial importance
used as fibers for coir, jute, cotton etc. in textile
industries.

Sclerids (Stone cells):

Short, highly thick walled, irregular shaped, lignified dead cells. They
develop by secondary thickening of parenchymal cells.
The sclereids have pits in their walls; pits may be simple or bordered.
Distribution: found in groups or single present in the hard covering of
seeds and nuts, endocarp of stony fruits (E.g. coconut) and pulp of
many fruits such as custard apple, Guava, Pear, Sapota etc. they also
occur in cortex phloem, pith of certain plants.
Depending upon the cell size, shape and nature of cell wall
Brachy sclereids: Small unbranched (isodiametric cells with branched
pits. E.g. Guava, Pear, etc.
Macro sclereids: Elongated rod shaped / columnar shape. E.g. Seeds of
Leguminous plants.
Osteo sclereids: Bone shaped cells. E.g. Xerophytic leaves.
Astero sclereids: Star shaped sclereids. E.g. intercellular spaces of
leaves and petioles of certain aquatic plants.
Functions:
Lignified structures give the mechanical support.
Firmness and hardness to plant parts.

Forms the shell/Seeds.

 Complex permanent tissues

Complex permanent tissues are composed of

different kinds of cells and perform common function.

Xylem and phloem are the complex permanent

tissues. They are present in vascular plants; also

called heterogenous tissues, where the different cells
have different structure and function.

Complex permanent tissues are found in

Tracheophyta (Pteridophytes, Gymnosperms and

Angiosperms) helps in translocation of water, solutes,
foods substances and give mechanical support.

 Xylem (Nageli: Xylem = Wood)

It is a complex, dead, mechanical, permanent,

vascular tissue present in the vascular bundles. It is

a principal water conducting tissue.

Based on the origin and development two types of

xylem are found

Primary xylem it is originated from pro cambium
and
Secondary xylem it is originated from intra
fascicular cambium.

Xylem is heterogenous tissue composed of different

types of cell elements tracheids, tracheae
(vessels), xylem fibers and xylem parenchyma.

 Tracheids: most primitive tissue of xylem cells,

originate from pro cambium present in

Tracheophyta (Pteridophytes, Gymnosperms and
Angiosperms)
Tracheids are elongated tube like cells with
tapering ends; cell walls are thick, hard and
lignified.
Lumen is empty, considerably larger; it appears
circular or polygonal in cross section.
Cells lack protoplast (dead) at maturity.

Functions:
Adapted for transporting water and mineral solutes
from roots to the stems and leaves.
The thick and rigid walls of tracheids also aid in
providing mechanical support.

 Tracheae (Vessels): Found mostly in angiosperms,

in few gymnosperms and in pteridophytes.

Vessels consists of long cylindrical, pipe like or
broad, tube like series of cells with transverse
end wall running parallel to the long axis of the
plant body, lumen large, sinuses of cells are
broad and wide. The cell wall is hard, thick and
lignified. Various thickenings are seen.

Functions
Adapted for transporting water and mineral
solutes from roots to the stems and leaves.
The thick and rigid walls of tracheids also aid in
providing mechanical support.

 Xylem fibers: Sclerenchymal fibers in association with xylem

form the Xylem fibers. They are slender, spindle shaped,

dead. The Cells are thick, hard and lignified.

Function: Helps in conduction of water, and are Mechanical in

support.

Xylem parenchyma (Wood Parenchyma): Xylem parenchyma

is produced by pro cambium of apical meristem. Cells living,

compactly packed and more or less cylindrical, placed end to
end, may be thick or thin walled, with or without pits. The
Tyloses are seen.

Tyloses: The parenchyma cells extend to the lumen of xylem

and look like balloons/ or buttons. They hinder the ascent of

sap.

Functions: Helps in conduction of water, store food materials.

Phloem (Gk; phlois = bark)

It is a complex, living, mechanical, permanent, vascular

tissue present in the vascular bundles. It is a principal
Food conducting tissue commonly referred as Bast.
Based on the origin and development two types of

phloem are found

Primary Phloem it is developed from pro cambium,
and
Secondary Phloem it is developed from Vascular
cambium.

Phloem tissue composed of different types of cell

elements Sieve elements, Companion cells, Phloem
parenchyma and Phloem fibers.

 Sieve elements: the conducting elements of phloem are sieve

elements. The less specialized cells are Sieve cells. The more
specialized cells are called Sieve tube members.
Sieve cells: these are narrow elongated cells with tapering ends.
These have sieve areas throughout the lateral walls and end walls.
They are found in Pteridophytes and Gymnosperms.
Sieve tubes:
They are tube like, placed end to end, resemble Xylem vessels,
found in Angiosperms and form the main phloem elements.
The sieve tube members are living with perforated oblique or
transverse end walls. The perforated end walls are like a sieve and
are called Sieve plates. The perforated areas of sieve plates are
known as Sieve areas.
Sieve tubes comprise several to many enucleated sieve tube
members inter connected via the sieve plates or sieve tube in a
series. These are found only in Angiosperms.
The cell wall is thin in most and thick (due deposition of Pectin) in
few and made up of cellulose.
During winter, the sieve areas are covered by the deposit of
colorless shining substance in the form of a layer called callus.

 Sieve tube member: they are found in almost every

part of Angiosperms. They are elongated tube like

cells placed one above the other. The term Sieve
refers to group of pores that are located on the end
walls and some times on the sides.
Function of Sieve cells and Sieve tubes:
Sieve cells and Sieve tubes are the chief food
conducting elements of phloem. They transport the
prepared food materials.

Companion cells: They are living nucleated specialized

parenchymatous cell with dense cytoplasm, associated
with a sieve tube in origin, position and function. They
develop from same Meristematic cell and gives rise to
the sieve element. Usually a single companion cell is
found associated with Sieve tube member. The Sieve
tube members and Companion cells are
interdependent, as the companion cell nucleus directs
the activities of Sieve tube cell. If one dies, the other
will die soon.
Functions:
They are involved in the conduction of food materials.
The companion cell nucleus directs the activities of
Sieve tube cell.
They are involved in the maintenance of pressure
gradient in the sieve tubes.

 Phloem parenchyma: The parenchymatous

cells found associated with phloem and are

living cells with dense cytoplasm and nucleus. It
is found as the component of phloem of
Gymnosperms
and
Dicotyledons
of
Angiosperms. In dicotyledons the phloem
parenchyma store starch, oil, and other food
substances and accumulate tannins and resins.
Phloem
fibers/
Bast
fibers:
the
sclerenchymatous fibers found associated with
phloem and are long lignified; provide
mechanical strength to the plant body. The
phloem fibers have commercial value used in
industries.

Plant Anatomy.
The study of internal structure of plants is plant anatomy.
Based on the origin vascular tissues are classified as follows:
Primary Vascular Tissues: formed by the pro cambium of
apical meristems. These are primary vascular tissues, present in
primary shoots and roots. E.g. All monocots, few herbaceous
dicots.
It brings about the primary growth.
Secondary Vascular Tissues: formed by the vascular cambium
of vascular bundle (Lateral meristems). They bring about increase
in the diameter of the plant (Secondary growth -girth). E.g. Dicots
and Gymnosperms.
Exarch xylem: the differentiation of cells occurs centripetally.
That is the first formed xylem (Proto Xylem) present far away
from the center of the axis or it faces towards the peripheral/
marginal side and later formed xylem present at center/ towards
the pith. E.g. Roots.
Endarch Xylem: the differentiation of cells occurs centrifugally.
That is the first formed xylem (Proto Xylem) present in the
peripheral/ marginal side and later formed xylem present at
center/ towards the pith. E.g. Roots.

 Proto xylem: first formed xylem, consisting of

tracheary elements, narrow lumen with Annular,

Scalariform and Spiral thickenings. The thickening
begins before the completion and elongation of the
xylem cell.

Meta xylem: later formed xylem, consisting of

tracheary elements with large lumen, with reticulate

and pitted thickenings. The thickening begins after
the completion and elongation of the xylem wall.

Proto phloem & Meta phloem like xylem the

phloem is also differentiated in to protophloem (first

formed phloem) and Meta phloem (Later formed). In
most of the plants the Proto phloem & Meta phloem
not distinguishable.

 Vascualr bundle
The strand of tissue composed of primary xylem and phloem is called vascular

bundle.
Based on the origin, formation and position the vascular bundles are classified
in to Radial vascular bundles and Conjoint vascular bundles.

1. Radial vascular bundles: In this bundle the xylem and phloem form equal
number of separate bundles, which lie on different radii alternating with each
other, as in roots.
2. Conjoint vascular bundles: In this the xylem and phloem are joined to
form a bundle. Conjoint bundles may be
a. Collateral vascular bundles: In this the xylem and phloem of each bundle
lie on the same radius, with xylem on the inner side and phloem towards the
exterior. When in collateral vascular bundle, if cambium is present (as in
dicotyledonous stems), the bundle is said to be open, if cambium is absent (as
in monocotyledonous stems), the bundle is said to be closed.
b. Bicollateral vascular bundles: In this type, there is an outer phloem,
median xylem and inner phloem. Bicollateral vascular bundles are always open.
E.g. Cucurbitaceae, Solanaceae, Apocyanaceae etc.
c. Concentric vascular bundle: If one kind of vascular tissue is surrounded by
the other. There are two typesAmphivasal: In this type, the xylem is surrounded by phloem. E.g. Selaginella.
Amphicribal: In this type, the phloem is surrounded by xylem. E.g. Draceaena.

Secondary Growth
An increase in the thickness of the plant body by the
addition of secondary permanent tissues is called
Secondary growth.
It occurs after primary growth.
Observed in stem and root of shrubs and trees of
dicot and gymnosperms.
Secondary permanent tissues formed were
Secondary xylem, Secondary phloem and Secondary
cortex.
The secondary xylem formed is called as Wood and
the external cork layer is called Bark.
Lateral meristems namely Vascular cambium and
cork cambium are involved in secondary growth.
Secondary growth in Dicot stem takes place in Stele
(Intrastelar growth) and in the Cortex (Extrastelar
growth).

Intrastelar Secondary growth

In the stele of dicot stem, vascular bundles are arranged in the
form of a broken ring. The space between the vascular bundles
is occupied by medullary rays composed of parenchyma cells.
There is a strip of cambium between the xylem and phloem of
each vascular bundle. It is called intrafascicular cambium.
On the on set of secondary growth, some parenchyma cells of
the medullary rays undergo dedifferentiation and meristematic.
This strip of cells is now called interfascicular cambium.
The meristematic cells of the intrafascicular cambium fuse to
form a continuous cambial ring called vascular cambium of
fascicular cambium. It initiates intrastelar secondary growth.
The cambial ring at this stage has primary xylem on its inner
surface and primary phloem on its outer surface.
The fascicular cambial cells multiply continuously to form
numerous daughter cells, on inner and outer surface. The cells
of on the inner surface differentiate in to secondary xylem,
while those on the outer surface differentiate into secondary
phloem

 The secondary phloem pushes the primary phloem outwards, which

becomes crushed. And the secondary xylem pushes the primary xylem
towards the pith but being hard, it remains uncrushed.
As a result of secondary growth, a continuous ring of secondary xylem and
a ring of secondary phloem are formed every year. The successive rings of
secondary phloem are pushed outwards and become crushed. There fore,
there is no increase in the bulk of secondary phloem. The secondary
xylem, which is hard, cannot extend inwards to a great extent because;
this area is already occupied by the primary xylem and pith. Hence, it
extends outwards year after a year resulting in an increase in the girth of
the stem. With the addition of these tissues inside the stele, a pressure is
exerted on the pericycle and endodermis. There fore, they are crushed.
The mitotic activity of the cambial ring is seasonal. It occurs only twice
during every year, once in the spring and once in the autumn. Thus, every
year two sets of secondary xylem and two sets of secondary phloem are
formed. Each year, the mitotic division of the cambial ring usually begins
in the spring season. The secondary xylem that is formed in the spring
season is, there fore, known as spring wood or early wood, while the
secondary xylem formed in the autumn is known as autumn wood or late
wood. The springwood is generally characterized by the presence of
xylem vessels having wider lumen. This is because, spring is the ideal
season for the growth and the water requirement of the plant is more in
the spring. The autumn wood has xylem vessels with narrow lumen, since
water requirement in the winter is less.

 The inner springwood and the outer autumn wood together represent

the annual ring. One such ring is formed every year due to secondary
growth. Thus, it is possible to ascertain the age of a dicot tree by
counting the number of annual rings. The branch of biology, which
deals with the ascertaining the age of the tree by counting the number
of annual rings, is called Dendrochronology. While every year two sets
of secondary xylem and two sets of secondary phloem are formed, only
one set is visible because the secondary phloem formed later grows
over and masks the secondary phloem formed earlier.
As secondary growth continues, several changes take place in the
secondary xylem. In an older stem, the secondary xylem shows two
regions-Duramen and Alburnum.
Duramen, also known as Heartwood formed earlier, represents the
centrally located, inactive portion of the secondary xylem. It appears
dark in colour due to accumulation of pigments resulting from the
oxidation of Oils and Tannins. It is incapable of conducting water and
provides only mechanical support to the trees. It is this part of the
secondary xylem that is used as the commercial wood.
Alburnum, also known as Sapwood formed later, represents the
peripheral, active portion of the secondary xylem. This portion is
capable of conducting water. Due to its merge mechanical strength and
lack of durability, the sapwood cannot serve as commercial wood.

Extra stelar secondary growth

It takes place due to the activity of secondary meristem called

cork cambium in the cortex.
To keep pace with the expanding stele and to maintain the area
equilibrium between stele and cortex, the cortex also enlarges.
Some of the parenchyma cells in the peripheral layers of the
cortex undergo dedifferentiation and become meristematic.
They constitute the cork cambium or phellogen. The cork
cambium undergoes mitotic divisions on both outer and inner
surfaces, just as the cambial ring in the stele. The mitotic
activity on the inner surface of the cork cambium results in the
formation of cells, which undergo differentiation in to a living
tissue, called Secondary cortex or Phelloderm, just above the
primary cortex. The mitotic activity on the outer surface results
in the formation of cells, which undergo differentiation in to a
dead tissue, called cork or Phellum, just below the epidermis

 The Phellogen, Phelloderm and Phellum together

constitute Periderm.
The periderm along with the primary cortex
represents the bark. In several dicot plants, the bark
peels off regularly.
Due to the formation of periderm, the epidermis is
subjected to pressure and, as a result, it breaks at
several places to form openings called Lenticels. The
Lenticels, also known as aerating pores, enclose a
group of living cells called Complementary cells.
Through these cells, exchange of respiratory gases
and some extent transpiration, takes place.
Thus, secondary growth in the cortex results in the
formation of Periderm due to which there is an
increase in the girth of the cortex.