The below mentioned article provides

study notes on Secondary Growth in Dicot

Stem of plants.

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.

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.
A. Formation of Secondary Vascular
TissuesS:

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 forma 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 dividee
periclinally both on the outer and inner sides
(bipolar divisions) to form secondary
permanent tissues.
rneun
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.
 EPIDERMIS CAMBIUM CELL
CORTEX
PERICYCLE CAMBIUM CELL
PRIMARY PHLOEM XYLEM MOTHER CELL
PITH PRIMAAY XYLEM
INTER PHLOEM MOTHER CELL
FASCICULAR
CAMBIUM CAMBIUM CELL
PRIMARY XYLEM MOTHER CELL
MEDULLARY RAYS
INTER-FASCICULAR B
CAMBIUM

PHELLOGEN OR
CORK CAMBIUM
LENTICLE
EPIDERMIS
PHELLODERM OR .
SECONDARYCORTEX
CORTEX

CRUSHED
PHLOEM
PITH
PITH
PERICYCLE
PRIMARY
PHLOEMA
SECONDARY PHLOEM
VASULAR CAMBIUM
SECONDARY XYLEM
C PRIMARY XYLEM. ANNUAL
FIRST RINGS OF
VASCULAR RAYS D SECOND YEAR
YEAR SECONDARY
XYLEM

. 6.28. A, complete ring of vascular cambium formed by strips of intrafascicular cambium and inter-fascicular
cambium. B, formation of secondary vascular tissue mother cells; C, the beginning o
secondary growth (mostly made up of secondary vascular tissues) of dicot stem
(diagrammatic); D, two-year stage of secondary growth of a dicot stem.
1. Vascular Rays:

The vascular rays or secondary medullary

rays are rows of radially arranged cells which
are formed in the secondary vascular tissues.
They are a few cells in height.

Depending upon their breadth, the vascular

rays are uniseriate (one cell in breadth) or
multiseriate (two or more cells in breadth).
Vascular rays may be homo-cellular (having
one type of cells) or hetero-cellular (with more
than one type of cells). The cells of the
vascular rays enclose intercellular spaces.

The part of the vascular ray present in the

secondary xylem is called wood or xylem ray
while the part present in the secondary
phloem is known as phloem ray. The vascular
rays conduct water and organic food and
permit diffusion of gases in the radial
direction. Besides, their cells store food.
 RAY
INITIALS

FUSIFORM
INITIALS

Fig. 6.29. LS. Vascular cambium

showing fusiform and ray initials.
2. Secondary Phloem (Bast):

It forms a narrow circle on the outer side of

vascular cambium. Secondary phloem does
not grow in thickness because the primary
and the older secondary phloem present on
the outer side gets crushed with the
development of new functional phloerm (Fig.
6.28 D). Therefore, rings (annual rings) are not
produced in secondary phloem. The crushed
or non-functioning phloem may, however,
have fibres and sclereids.

Secondary phloem is made up of the samme

type of cells as are found in the primary
phloem (metaphloem)- sieve tubes,
companion cells, phloem fibres and phloem
parenchyma.
Phloem pairenchyma is oftwotypes-axial
phloem parenchyma made up of
longitudinally arranged cells and phloem ray
parenchyma formed of radially arranged
parenchyma cells that constitute the part of
the vascular ray present in the phloem.
Elements of secondary phloem show a more
regular arrangement. Sieve tubes are
comparatively more numerous but are shorter
and broader. Sclerenchyma fibres occur either
in patches or bands. Sclereids are found in
many cases. In such cases secondary phloem
is differentiated into soft bast (secondary
phloem without fibres) and hard bast (part of
phloem with abundant fibres).

3. Secondary Xylem:

It forms the bulk of the stem and is commonly

called wood. The secondary xylem consists of
vessels, tracheids (both tracheary elements),
wood fibres and wood parenchyma.
Wood parenchyma may contain tannins and
crystals besides storing food. It is of two types
axial parenchyma cells arranged
longitudinally and radial ray parenchyma
cells arranged in radial or horizontal fashion.
The latter is part of vascular ray present in
secondary xylem.
Secondary xylem does not show distinction
into protoxylem and meta-xylem elements.
Therefore, vessels and tracheids with annular
and spiral thickenings are absent. The
tracheary elements of secondary xylem are
similar to those of meta-xylem of the primary
xylem with minor differences. They are
comparatively shorter and more thick-walled.
Pitted thickenings are more common. Fibres
are abundant.

Width of secondary xylem grows with the age

of the plant. The primary xylem persists as
conical projection on its inner side. Pith may
become narrow and ultimately get crushed.
The yearly growth of secondary xylem is
distinct in the areas which experience two
seasons, one favourable spring or rainy
season) and the other un-favourable (autumn,
winter or dry summer).
Conical projection on its inner side. Pith may
become narrow and ultimately get crushed.
The yearly growth of secondary xylem is
distinct in the areas which experience two
seasons, one favourable spring or rainy
season) and the other un-favourable (autumn,
winter or dry summer).

In favourable season the temperature is

optimum. There is a good sunshine and
humidity. At this time the newly formed
leaves produce hormones which stimulate
cambial activity. The activity decreases and
stops towards the approach of un-favourable
season. Hence the annual or yearly growth
appears in the form of distinct rings which are
called annual rings (Fig. 6.30).
Annual rings are formed due to sequence of
rapid growth (favourable season, e.g, spring),
slow growth (before the onset of un-
favourable period, e.g., autumn) andno
growth (un-favourable season, e.g., winter).
Annual rings are not distinct in tropical areas
which do not have long dry periods.

ANNUÁL AINGS

Fg. 6.30. Part of T.S.

old stem showing
annual ings.

Annual Rings (Growth Rings). It is the wood

formed in a single year. It consists of two
types of wood, spring wood and autumn wood
(Fig. 6.31). The spring or early wood is much
wider than the autumn or late wood. It is
lighter in colour and of lower density. Spring
wood consists of larger and wider xylem
elements.
The autumn or late wood is dark coloured and
of higher density. It contains compactly
arranged smaller and narTower elements
which have comparatively thicker walls. In
autumn wood, tracheids and fibres are more
abundant than those found in the spring
wood.

The transition from spring to autumn wood in

an annual ring is gradual but the transition
from autumn wood to the spring wood of thne
next year is sudden. Therefore, each year's
growth is quite distinct. The number of annual
rings corresponds to the age of that part of the
stem. (They can be counted by increment
borer).

Besides giving the age of the plant, the annual

rings also give some clue about the climatic
conditions of the past through which the plant
has passed. Dendrochronology is the science
of counting and analysing annual growth
rings of trees.
 LENTCEL
-EPIDEAMIS
CORK OA PHELLEM

CORK CAMBIUM
PER:0ERM OR PHELLOGEN

sECONDARY
CORTEX
CORTEX oR PHELLODEAS

SCLERENCHYMA CRLSHE
PERICYCLE) PRIAMARY PHLOEMA

PHLOEMRAY SECONDARY
PHLOE
GAMBIUt

AUTUMN WoOD

SPRING ooD
SECONDARY
MEIULLARY RAY

XYLEV RAY

PA:MARYv
MEDULLARY RAY

-PRIMANY xYLEM

-PTH

Fig. 6.31. Part of detailed structure of transverse section of

two year old dicot stem showing secondary growth.
Softwood and Hardwood:

Softwood is the technical name of

gymnosperm w0od because it is devoid of
vessels. Several of the softwoods are very easy
to work with (e.g, Cedrus, Pinus species).
However, all of them are not 'softe. The
softness depends upon the content of fibres
and vascular rays. 90-95% of wood is made of
tracheids and fibres. Vascular rays constitute
5-10% of the wood.

Hardwood is the name of dicot wood which

possesses abundant vessels. Due to the
presence of vessels, the hardwoods are also
called porous woods. In Cassia fistula and
Dalbergia sisso the vessels are comparatively
very broad in the spring wood while they are
quite narrow in the autumn wood. Such a
secondary xylem or wood is called ring
porous.
In others (e.g., Syzygium cumini) larger sized
vessels are distributed throughout spring
wood and autumn wood. This type of
secondary xylem or wood is known as diffuse
porous. Ring porous wood is more advanced
than diffuse porous wood as it provides for
better translocation when the requirement of
the plant is high.
The wood of the older stems
(dalbergia,
Acacia) gets differentiated into two zones, the
outer light coloured and functional
sapwood
or alburnum and the inner
darker and
nonfunctional heartwood or duramen (Fig.
6.33). The tracheids and vessels of the heart
wood get plugged by the in growth of the
adjacent parenchyma cells into their cavities
through the pits. These ingrowths are called
tyloses (Fig. 6.32).
VESSEL PARENCHYMA VESSEL

PIT

TYLOSES
A

6.32. Formation of tyloses in heartwood.

Fig.
A, LS. vessel showing yloses.
B, T.S. vessel showing tyloses.
 BARK
SAP
woOD
HEART
WOOD

Fig. 6.33. Sapwood and heartwood

in TS. of trunk.
Ultimately, the parenchyma cells become
lignified and dead. Various types of plant
products like oils, resins, gums, aromatic
substances, essential oils and tannins are
deposited in the cells of the heartwood. These
substances are collectively called extractives.
They provide colour to the heartwood. They
are also antiseptic. The heartwood is,
therefore, stronger and more durable than the
sapwood.

It is resistant to attack of insects and microbes.

Heart wood is commercial source of Cutch
(Acacia catechu), Haematoxylin
(Haematoxylon campechianum), Brasilin
(Caesalpinia sappan) and Santalin
(Pterocarpus santalinus). Heartwood is,
however, liable to be attacked by wood rotting
fungi. Hollow tree trunks are due to their
activity.
Significance of Secondary Growth:
1. Secondary growth adds to the girth of the
plant. It provides support to increasing weight
of the aerial growth.

2. Secondary growth produces a corky bark

around the tree trunk that protects the interior
from abrasion, heat, cold and infection.

[Link] adds new conducting tissues for replacing

old non-functioning ones as well as for
meeting increased demand for long distance
transport of sap and organic nutrients.
Anomalous Secondary Growth:

It is abnormal type of secondary growth that

occurs in some arborescent monocots (e.g,
Dracaena, Yucca, Agave) and storage roots
(e.g., Beet, Sweet Potato). In arborescent
monocot stems, a secondary cambium grows
in hypodermal region. The latter forms con-
junctive tissue and patches of meristematic
cells. The meristematic patches grow into sec-
ondary vascular bundles.

Anomalous vascular bundles also occur in

cortex (cortical bundles, e.g, Nyctanthes) and
pith (e.g., Boerhaavia). In storage roots (e.g.,
Beet), accessory cambial rings appear on the
outside of endodermis. They produce less
secondary xylem but more secondary phloem.
The secondary phloem contains abundant
storage parenchyma.
Importance of Secondary Growth:

1. It is a means of replacement of old non-

functional tissues with new active tissues.

2. The plants showing secondary growth can

grow and live longer as compared to other
plants.
3. It provides a fire proof, insect proof and
insulating cover around the older plant parts.

4. Commercial cork is a product of secondary

growth. It is obtained from Quercussuber
(Cork Oak).

5. Wood is a very important product of

secondary growth. It represents secondary
xylem.