Unit 6 Plant Anatomy and Embryology

UNIT 6
SECONDARY GROWTH

Structure
6.1 Introduction Economic Value of Secondary
Phloem
Objectives
6.7 Secondary Growth in
6.2 Secondary Growth in Typical Monocot Stem
Dicotyledonous Stem
6.8 Periderm
6.3 Vascular Cambium
Structure
Structure of Vascular Cambium
Phellem
Types of Cambium
Phelloderm
Structure of Protoplast and Cell
Growth Origin and Development of
Periderm
Ray Initation
Bark
6.4 Cambium Activity
Commercial Cork
Formation of Annual Rings
6.9 Distribution of Lenticels
6.5 Secondary Xylem
Development and Structure of
Basic Structure of Secondary
Lenticels
Xylem
6.10 Cambial Variants
Wood Parenchyma
In Stems
Heart Wood and Sapwood
In Roots
Porous and Nonporous Wood
6.11 Summary
Economic importance of Wood
and its Characteristics 6.12 Terminal Questions
6.6 Secondary Phloem 6.13 Answers

6.1 INTRODUCTION
You have studied that among seed bearing plants, herbaceous annuals attain
a limited height and do not need to increase in girth. Primary growth is often
sufficient to meet their structure needs. However, in woody perennials that
reach enormous height and produce large canopies, increase in girth is
necessary to support the weight of the shoot. Secondary growth, derived from
secondary or lateral meristems results in increase in diameter of stems and 107
Block 2 Secondary Growth and Adaptive Features
roots. In the present unit we will describe the structure of vascular cambium,
explain its functioning and role it plays in forming the woody tissues.

The ability of woody plants to undergo secondary growth and produce wood
has many consequences. Woody plants when grow in girth which also contain
conducting tissues gives plant greater capacity to move water and minerals
upward and carbohydrates downward thus number of leaves and roots that
the plant can support increases as does the photosynthetic capacity.

Vascular cambium is a bifacial meristem, which adds to the girth of stem and
root. Vascular cambium varies greatly in its activities during different seasons,
in different plants and on different parts. The active meristematic tissues i.e.,
fascicular cambium regions lies between primary xylem and phloem.
Interfascicular cambium arises from parenchyma cells between vascular
cambia. The fascicular and interfascicular cambium joins to form ring which in
turn give rise to cambial ring. The cells of the cambium differentiate to form the
secondary tissues.

Objectives
After studying this unit you would be able to :

explain the phenomenon of secondary growth in plants;

describe the structure and give function of each of the following

vascular cambia,cork cambium, lenticels;

identify the secondary growth of monocotyledonous and

dicotyledonous stems;

distinguish between types of wood, its annual rings, sapwood,

heartwood and bark;

appreciate the woody anatomy and climate studies (Dendroclimatogy);

explain the commercial uses of cork and different wood; and

list various types of unusual secondary growth in stems and roots.

6.2 SECONDARY GROWTH IN TYPICAL

DICOTYLEDONOUS STEM
You have already read in block 1 that the primary plant body in itself is
structurally and functionally complete, for example the majority of
monocotyledons and pteridophytes. In gymnosperms and most dicotyledons
primary growth is followed by secondary growth. In stem, the secondary
growth in thickness in diameter is confined both intrastelar, i.e, within the stele
and extrastelar regions. The cells that form secondary tissues are produced by
lateral meristems. The lateral meristems grow and join to make a circular ring
known as the vascular cambium which lays down cells that become the
secondary vascular tissues. In the stem, cells which are situated between the
108 primary xylem and primary phloem in the vascular bundles become
Unit 6 Plant Anatomy and Embryology
meristematic and form part of the vascular cambium. Additional cells between
the vascular bundles also become meristmatic. Hence the vascular cambium
can be seen in a cross section of the stem as a continuous ring of tissue, with
the xylem and pith on the inside and phloem, cortex, and epidermis on the
outer side of interfascicular cambium/cambium ring (Fig. 6.1).

Fig. 6.1: Diagrammatic representation of secondary growth in a dicot stem upto

two year (stages (1-4) in T.S.).

The vascular cambium usually, if not always, has a dual origin within the
primary tissues; from provascular strands, and from the "ground" meristem
tissues between those strands. These two modes of origin are termed
intrafascicular (within fascicles) and interfascicular (between fascicles). The
term "provascular tissue" will be used in the text. We should know what this
means. Provascular tissue is the precursor of all vascular tissues and
"Procambium" is that part of the provascular tissue that is the precursor of the
vascular cambium (which may also produce some metaxylem).The transitional
stages between procambium and cambium are denoted as metacambium.

Both procambium and then metacambium differentiate acropetally within the

provascular bundles. Most of the divisions in this layer are periclinal, producing
metaxylem and metaphloem. The cells between the metaxylem and
metaphloem eventually begin to function as cambial initials. Cambia initials
consists of two morphological types of cells-axially short, blocky, ray cells and
long, slender fusiform cells. Procambium at first consists of short cells from
which longer cells may arise in two ways : 109
Block 2 Secondary Growth and Adaptive Features
1) Different cell lengths result from locally different rates of transverse
and/or pseudotransverse cell divisions during growth. Thus the shorter
cells become ray initials and the longer become fusiform initials
2) All procambial cells first become quite elongated. Then some of them by
nonrandom transverse and/or pseudotransverse divisions are
secondarily transformed into sets of axially short ray initials. As soon as
a circle of vascular cambium is completed, its cells divide to produce
new cells.
Those cell formed inside the ring of cambium differentiate into secondary
xylem or wood, about which you would study in the later part of this unit. Most
of the cells of secondary xylem have very thick walls. As the cambium
produces new wood, the stem increases in diameter, the phloem peripheral to
the vascular cambium becomes stretched. In the mean time cells produced
just outside the vascular cambium becomes differentiated into secondary
phloem, and participates in the transport of organic substances. As more
secondary xylem is formed, the first formed secondary phloem is destroyed
and replaced by the newly formed secondary phloem (Fig. 6.2).

Fig. 6.2: Formation of complete vascular cambium. a) After completion of

primary growth some meristematic cells remain between primary xylem
and primary phloem; b) The residual procambium becomes reactivated
to form fascicular cambium and some parenchyma cells of pith become
meristematic to form the interfascicular cambium; c) Formation of
complete cylinder of vascular cambium; d) Cylinders of secondary
phloem and secondary xylem have been formed by vascular cambium.

With the addition of secondary tissues, the stem grows thicker with secondary
110 xylem and phloem, the peripheral primary tissues, cortex and epidermis
Unit 6 Plant Anatomy and Embryology
become compressed and destroyed. The epidermis is replaced by a new
protective layer of secondary tissue. Simultaneously another lateral meristem
called phellogen (formerly termed cork cambium), is differentiated. It divides
and produces new cell towards the outside. These cells become suberised
i.e., impregnated with a waterproof waxy material and die, giving rise to a
protective layer of cork. Secondary growth in roots is similar to that in the
stem. The main roots of a tree are large and woody and provide anchorage to
the plant. The tasks of absorbing water and minerals are performed by
younger roots at the far ends of the root system. A detailed account of the
structure and description of secondary tissues and secondary growth will be
dealt later in the unit.

SAQ 1
a) Define secondary growth.

b) Name the region in a dicot stem where secondary growth occurs.

6.3 VASCULAR CAMBIUM

In certain plants, including monocotyledons, all the cells of the procambium
In gymnosperm initials
undergo differentiation into primary vascular tissues. In almost all the 600-8700µm long was
dicotyledons and gymnosperms, a portion of the procambium remains reported. Fusiform
meristematic even after the completion of primary growth and develops into initials sometimes
the cambium of the secondary body. become very long in
old trunks of Sequoia
The cambium that arises within the bundles of primary vascular tissue of the sempervirens
stem is called fascicular cambium, because it originates within the bundles or e.g.they reach a
larger segments of the primary vascular system. Commonly the bands of maximum length of
fascicular cambium become interconnected by additional bands of meristem, 8700 µm. (Bailey,
the interfascicular cambium, which originates from interfascicular 1923).
parenchyma. A completely formed cambium of the stem has the shape of
hollow cylinder, extending through the nodes and internodes. If the axis is
branched, the main axis of the cambium is continuous with that of the
branches, and it may extend some distance into the leaves.

Fig .6.3: Division in Vascular cambium. 111

Block 2 Secondary Growth and Adaptive Features
The procambium and cambium may be looked upon as two developmental
stages of the same meristem; they intergrade with regard to their
morphological and physiological characteristics. The typical features of
cambium of a woody dicotyledons and gymnosperms is segregation of its
initials into fusiform and ray initials, the occurrence of apical growth, and the
precise method of division in a tangential plane during the formation of xylem
and phloem (Fig. 6.3).

The origin of the interfascicular cambium in the more or less vacuolated

interfascicular parenchyma results from a resumption of meristematic activity
by a potentially meristematic tissue. Usually, no cytological changes are
noticeable in connection with the return to meristematic activity. In most of the
dicotyledons and gymnosperms the cambial cylinder develops between the
axillary xylem and phloem, a position that is retained throughout the life of the
plant. It is from this point that the cambium produces the secondary xylem
centripetally and secondary phloem centrifugally (Fig.6.4). Now we will study
about the strucuture of vascular cambium.

Fig. 6.4: Cambial initials and its derivatives; and Production of new xylem and
new phloem each year. Vascular cambium is shifted away from the
centre of the plant.

6.3.1 Structure of the Vascular Cambium

Active cambial cells are richly cytoplasmic and not wholly undifferentiated and
different in shape from embryonic ground meristematic and apical meristem.
These cells have pitted thin walls while radial walls are thicker than the
tangential walls. The cambial cells possess numerous cytoplasmic
connections. The cytoplasm of the cells is rich in ribosomes, dictyosomes and
endoplasmic reticulum.

The differentiation of the cambial derivatives into xylem and phloem removes
cells from the cambial zone but zone itself does not get removed (Fig. 6.5). If
the rate of radial growth and periclinal division are balanced by the rate of cell
loss through differentiation, the cambial zone thickness remains constant.

The new cell becomes larger than the cell from which it is derived. The
formation of the cell plate is peculiar during the longitudinal division in these
cells.

This meristem, in the anatomical sense of the term usually includes two
112 histologically distinct kinds of cells: i) Fusiform Initials, ii) Ray Initials.
Unit 6 Plant Anatomy and Embryology
i) Fusiform Initials
These are typically axially elongated cells with tapered ends. The mean length
of fusiform cells varies, among taxa and within an individual plant. It tends to
increase with the age of the plant. These cells are shaped somewhat like flat
shoe laces (Fig. 6.5). It would be easy to assume that they have about 8 faces
but a study of Pinus sylveslris has shown that 8 sides are minimum; they can
often have up to 32 faces with an average of 18; they have 14 contact faces
with other such cells. The fusiform initials undergo periclinal division. The
dividing fusiform cells show no phragmoplast function which appears at the
end of the mitosis. It directs the cell plate expansion laterally. Fusiform cell
length is greater in angiosperms. These cells are shorter when present in
storied cambia. Fusiform initials differentiate to form secondary xylem and
secondary phloem.

The periclinal divisions of the fusiform initials result in radially oriented file of
cells. The cells differentiate to form secondary xylem towards the centre of the
axis and form secondary phloem towards the periphery. Few derivatives of the
fusiform initials remain undifferentiated and retain the feature of meristematic
cells. These cells form the cambial zone.

The proportion of ray

initials in the cambium
is variable. In conifers
they are arranged as
vertical, uniseriate
groups that produce
vertical, uniseriate
rays.

Fig.6.5: Schematic drawing: cambial initials cells (a-e) produces secondary

xylem and secondary phloem. f) Fusiform initials can be seen in three
dimensional views. 113
Block 2 Secondary Growth and Adaptive Features
ii) Ray Initials
They are smaller than fusiform initials and are nearly isodiameteric (equal
dimensions, small sides) or only about two or three times as tall as wide. Let
us study how the ray initials are formed in cambium (Fig. 6.5):

i) A single cell may be cut off along the side of a fusiform initial (lateral
division),
ii) A single cell may be cut off the end of fusiform initials,
iii) A declining fusiform initials may be reduced to a single ray initials, and
iv) The last part or whole fusiform initial

Sometimes the fusiform initials can be converted to ray initials in

gymnosperms and dicots. In contrast in dicots ray initials can elongate and
convert themselves into fusiform initials this conversion prevents rays from
becoming too massive and creating large islands of weak parenchyma in
wood. The ray initials can also be broken by intrusive growth of a fusiform
initial from the periphery of the group into the mass of ray initials.

The ratio of fusiform initials to ray initials is high in dicots. In most of the
species 90% of the cambium is reported to consist of fusiform initials. The
ability of a fusiform initial to form ray initial is important because multiplicative
divisions produce a common wall that is not in contact with the ray. A second
division produces an entire cell with no ray contacts. New ray cells need to be
produced to maintain proper horizontal conduction through wood, bark and
cambial zone. Rays in woody dicots are mainly multiseriate while
gymnosperms show the presence of uniseriate rays.

Ray initials are smaller than the fusiform initials they are shorter and
isodiamteric. The ray cells appear quadrilateral and smaller in length in the
tangential section of the stem. They usually appear lens shaped group of ray
initials. Dicots generally have both uniseriate and multiseriate group of ray
initials.

It is interesting to note that the vascular cambium remains dormant during

stress. As it enters dormancy, it stops cell division. Some of the xylem and
phloem mother cells become quiescent and get partially differentiated. After
the stress gets over they start differentiating and quickly form the new tissues.
The cambial initials resume cell division in spring and the mitosis starts in the
cambial cells. This is triggered by the basipetal movement of the auxin. In
tropics the cambial activity of the trees continues throughout the year.

SAQ 2
Fill in the blanks with appropriate word(s).

a) Vascular cambium is composed of …………….. and …………. .

b) Vascular cambium remains dormant during ……………….

c) Ray cell contains ………….... duct.

d) Ray initials are………… than the fusiform initials.

e) ………………. behaves as a permanent meristem.

114
Unit 6 Plant Anatomy and Embryology
6.3.2 Types of Cambium
On the basis of the arrangement of the fusiform cells as seen in tangential
section, cambium is divided into:

i) Storied or Stratified Cambium

The groups of ray initials may become taller either by the loss of fusiform
initials located between two groups of ray initials, allowing them to fuse; or a
fusiform initials can by transverse division, convert itself into a row of ray
initials. All the structural elements which extend radially are produced by the
ray initials (Fig. 6.6 a). In this type the fusiform cells are arranged in tiers, or
stories, That is, the ends of large tangential groups of cells are aligned at the
same levels of axis. If you view them tangentially the ends of cells in axially
adjacent stories generally overlap only slightly making a zigzag pattern.

ii) Non-storied or Nonstratified Cambium

In this type of cambium the ends of cambial fusiform cells typically overlap
much more extensively and in a seemingly random manner. In nonstoried
cambia there is no lateral alignment. In vesselless dicotyledons the fusiform
initials may reach a maximum length of 6200 µm. Thus nonstoried initials are
longer. They are also of more common occurrence (Fig. 6.6 b).

(a) (b)
Fig. 6.6: L.S. views of fusiform initials and ray initials a) Storied cambium; b) Non-storied cambium.

6.3.3 Structure of Protoplast and Cell Growth

Active cambial cells are richly cytoplasmic and are not wholly undifferentiated.
They are obviously different in shape from embryonic ground meristem and
apical meristem cells. They also differ cytologically from the other meristematic
cells. They are more highly vacuolated, have large mitochondria and often
have more highly differentiated plastids. In a cambial fusiform cell, the nucleus
is quite elongated whereas in a ray cell it is usually more nearly spherically.
Active fusiform cells commonly have one or two large vacuoles transverse by
many slender cytoplamic strands, and small vacuoles in the peripheral
cytoplasm.

The cambium initials form pholem and xylem by tangential division. These
vascular tissues are laid down in two opposite directions, the xylem cells
towards the interior of axis and the phloem cells towards its periphery (also
see Fig. 6.3). The consistent tangential orientation of the planes of division
during the formation of vascular tissues determines the arrangement of
cambial derivatives in radial rows. Such radial seriation may persist in the
developing xylem and phloem or it may be disturbed through various kinds of
growth readjustments during the differentiation of these tissues. 115
Block 2 Secondary Growth and Adaptive Features
The thickness of xylem cylinder increases by secondary growth, and the
cambial cylinder also enlarges in circumference. Although active cambial cells
undergo repeated periclinal divisions and radial growth, the width of cambial
zone does not increase indefinitely. Conversely, although differentiation of
cambium derivatives into xylem and phloem continually removes cells from the
cambial zone, the zone itself does not disappear. If the rates of radial growth
and periclinal division are just balanced by the rate of cell loss through
differentiation, the cambial zone thickness remains constant. However, the
balance is often imprecise, and cambial zone thickness tends to vary during
the active season. The rate of production of cambial derivatives depends on
the number of cells in the cambial zone and on the duration of the cell cycle.

Addition of new fusiform initials is brought about by longitudinal anticlinal

divisions of the existing initials in the storied cambium while in nonstoried
cambium, the fusiform initials undergo oblique, pseudotransverse anticlinal
divisions, followed by intrusive growth, and each of the new cells becomes as
long as or even longer than the cell from which it was derived (Fig.6.7).

Fig. 6.7: Cambial development. a) Fusiform cambial initial; b) Ray initial: c) Ray
116 initial as in T.S.
Unit 6 Plant Anatomy and Embryology
Because of the excessive length of the fusiform initials, the formation of the
cell plate during the process of longitudinal division is peculiar to these cells.
The cell plate begins to form between the two nuclei and it spreads slowly.
The plate takes a long time to reach the end walls.

6.3.4 Ray Initiation

With the enlargement of the cambial cylinder new ray intials develop and
single fusiform initials are continuously lost from the cambium and are
replaced by new ones (Fig 6.8).You have studied how ray initials are formed in
section 6.3.1 of this unit.

Fig. 6.8: Origin of a secondary ray. a) Division of fusiform cambial initial to

uniseriate; b) Multseriate ray.

SAQ 3
a) Name the two cambia which join and form a cylinder of cambium.

b) Describe the types of cells found in the vascular cambium.

c) Differentiate between storied and non storied cambium.

6.4 CAMBIAL ACTIVITY

The radial growth is directly correlated with the rate of cambial activity. The
variation in the number of cell across the cambial zone seems to express the
balance between the rate of cell division and the rate of differentiation of the 117
Block 2 Secondary Growth and Adaptive Features
derivatives. However, at the time of cambial activity cell divisions are faster
than cell differentiation. This results in a wide cambial zone. But as soon as
demarcation begins, a balance is established and the width of the zone
remains more or less constant. When the rate of division becomes less and
the rate of differentiation is faster the cambial zone becomes narrower. The
vascular cambium shows variation in the period and intensity of activity. These
variations result from internal and external factors.

6.4.1 Formation of Annual Rings

The secondary xylem in perennial axis commonly consists of concentric
layers, each one of which represents a seasonal growth. If you see a cross
section of the axis, these layers appear as rings, and the terms annual ring
and growth ring or growth layer are applied to each layer.

An annual ring or growth ring of xylem is a layer of secondary xylem formed in

one growing season over the entire plant and is, therefore, an extensive
tubular structure having the general form of the axis of the plant. It is open at
ends where meristems occur. There are some plants in which cambium are
active during the entire life. These plants commonly occur in the tropical
regions where seasons are mot markedly different. However, not all tropical
trees exhibit a continuous cambial activity. In warm temperate climates, the
percentage of ringless trees is still lower.

In geographical regions where there are clear cut seasons, cambium shows
decreased activity with the onset of autumn, and it enters a dormant state
during winter. This may last till the beginning of the following spring. In spring
the cambial activity is resumed and becomes highest during summer. Thus
periodical activity of cambium results in the formation of ring wood. The wood
thus formed in spring is called spring summer wood or earlywood and that
formed in autumn is called autumn wood or latewood.

Fig. 6.9: T.S. of dicotyledonous stem showing annual rings.

It is noted that each annual ring corresponds to one year's growth hence the
age of plant can be determined roughly by counting the total number of annual
rings in a log (trunk) as seen in a transverse section (Fig. 6.9). Tree ring
analysis is also known as Dendrochronology. By analysing the tree rings, a
great deal can be learned about past climatic conditions. This study is known
118 as Dendroclimatology (See box 6.1).
Unit 6 Plant Anatomy and Embryology

BOX 6.1: TREE RING ANALYSIS - A WAY TO TELL THE LIFE HISTORY
OF A TREE.
Every year a new growth ring is added to trunk of the tree. In spring growth is fast and wood is light in colour
and in summer the wood is darker. By counting the dark rings you can tell the age of tree. In addition other
useful information can be obtained by analysing tree rings as weII. For example, the size of each ring varies
depending on environmental conditions, including precipitation and temperature.
Sometimes the variation in tree rings can be due to a single environmental factor. Then similar patterns
appear in the rings of many tree species in a large geographical area. For example, if a certain year is
drought year then in that particular year much smaller wood layer's will be produced. Sometimes locusts may
have eaten the leaves just after they have appeared. This will result in a marked decrease in photosynthesis,
causing low wood production. This will also result in two annual rings being very close each other because
very little growth will take place.
To study the sequence of rings, in trees that have lived for several thousand years, first a master chronology
of complete records of sample of rings dating back as far as possible is developed. Then by matching the
rings one can know the exact age of the living tree or trees (Fig.6.10).
A great deal can also be studied from tree rings about the past climatic conditions. For several years A.E.
Douglass, Harlod C. Fritts and others associated with the Laboratory of Tree Ring Research of the University
of Arizona Phoenix, USA studied ring widths in trees from various sites. They found that a very significant
statistical relationship exists between the growth of trees and climatic condition. With the help of computers
and statistical analysis they developed techniques that take into account of climate and other environmental
variables. They were able to reconstruct relatively precise histories of climatic changes and fluctuations
dating back thousands of years. Presently by ring analysis data are gathered to determine climates of
prehistoric times.

Fig. 6.10: Tree ring dating: A master chronology is developed using progressively older pieces of
wood from the same geographical area. The age of the sample can be accurately
determined by matching the rings of a wood sample of unknown age to the master
chronology.
119
Block 2 Secondary Growth and Adaptive Features
To determine the age of living old tree you do not have to cut it down, to examine the
rings. A simple instrument known as increment borer is used. The increment borer is
primarily made up of a rigid metal cylinder. It is driven in the stem of a tree and a core
of wood is removed. The hole is then treated with a disinfectant and covered up
without harm to the tree. The rings are examined counted from the core and then
analysed (Fig.6.10).

SAQ 4
Which of the following statements are true or false. Write T for true and F for
false.
a) All tropical trees exhibit a continuous cambial activity. [ ]
b) Regions with warm climate have a low per cent of ringless trees. [ ]
c) It is possible to calculate the approximate age of a tree by counting
the total number of rings in one log of wood. [ ]

6.5 SECONDARY XYLEM

The products of the cambium formed towards the centre of the stem and root
constitute secondary xylem. Secondary xylem is composed of tracheids,
vessel members, and different types of fibers, parenchyma cells, xylem ray
cells and sometimes secretory cells (Fig.6.11).
The occurrence and the arrangement of these elements vary in different group
of plants. The quantitative differences in the number of cells and the size of
the elements that exist between the species of a single genus make it possible
to identify individual species on the basis of secondary xylem alone.

Fig. 6.11: Structure of secondary xylem in three dimensional diagram of a cube

120 of Pinus.
Unit 6 Plant Anatomy and Embryology
6.5.1 Basic Structure of Secondary Xylem
Secondary xylem is characterised by the existence of two systems of elements
which differ in the orientation of their longitudinal axis. One system is
horizontal and other is vertical. The horizontal system is made up of xylem
rays (see Fig. 6.11) and the vertical or axial system consists of tracheary
elements, fibres and wood parenchyma. The living cells of the rays and of the
vertical system are usually interconnected and a continuous system of living
cells is formed, which in turn is connected with the living cells of the pith,
phloem and cortex.

6.5.2 Wood Parenchyma

Two types of parenchyma are found in secondary xylem: The axial
parenchyma and the ray parenchyma. The relatively short, special cambial
initials give rise to ray parenchyma, whereas fusiform initials from axial
parenchyma. The axial parenchyma cells may be as long as the fusiform
initials or much shorter. It is more common to find shorter parenchyma cells
(Fig. 6.12).

Fig. 6.12: Early and late wood vessels.

The ray parenchyma may be of different kinds, the most common forms are:
The color distinction
i) in those which longest axis of the cell is radial, between sapwood and
heartwood may be
ii) in which it is vertical.
sharp as noted in
The number of xylem rays increases with the expansion of stem girth. The Pinus roxburghii,
Dalbergia sisso,
length, width and height of each ray can be measured by cross sections and
Albizzia lebbek or
tangential sections respectively, when the ray is one cell wide it is called
gradual (Shorea
uniseriate ray; when two cells wide it is biseriate and when more than two robusta, Adina
cells wide, it is multiseriate. cordifolia). No color
distinction has been
All the cells of ray parenchyma may have primary wall or only secondary walls
noted in species such
are found. The secondary walls may develop pitpairs that are to be simple, as Abies pindrow,
half bordered, and sometimes even bordered. These secondary cell walls are Picea smithiana.
commonly characterised by the presence of depressions or cavities varying in
size, depth and structure. Such cavities are termed as pits .The parenchyma
cells of the xylem serve to store reserve food materials such as starch and
121
Block 2 Secondary Growth and Adaptive Features
fats. Tannin, crystals, silica bodies and other substances are also deposited in
these cells (Fig. 6.13 a). The ray parenchyma is the main route of radial
symplasmic transport between xylem and phloem.

In several plants wood parenchyma cells form protuberances which penetrate

into the vessels through the pits after they become inactive, or later injured.
These outgrowths are termed tylosis (- singular: tylose) (Fig. 6.13 b). The
nucleus and part of cytoplasm of the parenchyma cells may enter the tylosis.
Tylosis may also divide. Although the formation of tylosis is considered a
natural phenomenon, in many species it has been reported to result from
mechanical injury or diseases.

(a) (b)
Fig. 6.13 (a-b): Tylosis in wood.

6.5.3 Heartwood and Sapwood

The outer part of the secondary xylem contains living cells and at least one or
In some plants
two outermost rings participate in the conduction of water. The outer part of
heartwood contains
important pigments secondary xylem with living parenchyma is named as Sapwood or alburnum
such as hematoxylin (Fig. 6.14).
(Hematotoxylon
campechianuum),
brazilin (Cesalpinia
sappan) and santalin
(Pterocarpus
santalinus).

122 Fig. 6.14: Heartwood and sapwood in Cross Section of a tree.

Unit 6 Plant Anatomy and Embryology
In nearly all the trees the central portion of the xylem consists of dead
parenchyma which ceases to conduct water. This is called heartwood or Heartwood may
sometimes develop
duramen. The events that occur in the formation of heart wood include
as the result of
disintegration of protoplast, loss of cell sap and hydrolysis of reserve material pathological
stored and formation of tylosis. In such species in which the tylosis are formed conditions
the inner portion of the cell is totally blocked by the tylosis. Oil, gums, resins,
tanins, aromatic compounds and coloured substances which develop in the
cells are accumulated in the heartwood. The amount of heartwood and
sapwood varies in different species. These differences are also influenced by
genetic and environmental conditions.

In the timber trade wood of dicotyledons is known as hardwood and that of

gymnosperm as softwood. These terms do not accurately express the degree
of hardness, as in both groups wood with both hard and soft structure can be
noted. However, there is an important difference in the wood of dictoyledons
and gymnosperms. The former have vessels and the later lack them. If we see
the histological structure of the wood of dicotyledons and that of gymnosperms
there are fundamental differences.

6.5.4 Porous and non-porous wood

The wood is categorised on the basis of presence or absence of vessels. The
woody dicots that possess vessels in xylem are known as porous wood or
hard wood while the wood of gymnosperms that lack vessels is called non-
porous or soft wood.

Fig 6.15 (a to c): Ring porous and diffuse porous wood.

For example wood of Pinus is non-porous wood while Cedrus is referred as

soft wood. The hard wood looks more complex than the soft wood. In a
transverse section, the vessels appear in the form of circular or oval pores in
the hard wood. The arrangement of the pores varies for species and has been 123
Block 2 Secondary Growth and Adaptive Features
considered a diagnostic feature of timber. On the basis of distribution of pores
within the growth ring, the wood can be classified as ring-porous and diffuse
porous wood (Fig. 6.15). In ring porous wood, the pores of the early wood are
larger than those of late wood and arranged in the form of conspicuous ring. In
diffuse porous wood, the pores of the early and late wood do not show any
size difference in the pores and pores are uniformly distributed throughout the
growth ring.

6.5.5 Economic Importance of Wood and

its Characteristics
Wood has much importance in day to day life in form of furniture, paper gum,
resin and several other industrial purposes. We will discuss some of the
properties of wood by which wood quality is judged.

i) Weight : The wood may be either light or heavy. Differences in weight

are due to variations in the proportions of wall substance and of lumen
space, when the lumen is small the wood is dense and heavy. The
abundance of slender, thick walled fibres make wood heavy. Extremely
light woods such as Ochroma sp. is also found. The majority of well
known commercially important wood range from 0.35 to 0.65 in specific
gravity. Balsa wood (Ochroma lagopus) and sola (Acshynomene sp.)
have abundant of parenchyma and very few fibres.

Light woods have limited economic importance. Balsa (Ochroma sp.)

with a specific gravity of 0.12 to 0.35, is used extensively in insulation,
as material for modelling (by architects) and for life rafts. The heavy
woods are used in construction of wagons, carts, railway sleepers and
furniture etc.

If a large proportion of the wood is made up of fibers or fiber tracheids, it

tends to be strong. Thus the dense and heavy woods are of greater
strength. Strong woods are used for building, structural works and
furniture etc.

ii) Durability : The ability of wood to withstand decay by the action of fungi
and bacteria is largely dependent upon the chemical nature of the wood
and is referred as its durability. The presence of tylosis and other natural
constituents of wood such as tannin, resin and oils largely determine the
durability of wood. Both light and heavy woods can be durable. The
durable woods are used for ship building, boats, masts, ships, carts,
bodies of railway compartments, railway wagons, in construction,
bridges, and railway sleepers.

Some other important uses of Wood

Teak, sisham and rose wood are used as decorative timbers for panelling
furniture, cabinet, boxes, for carving idols, inlay works and various aspects of
art etc.

i) Pines : They are chiefly used for doors, windows, pattern making in
cabinets, boxes and matches. In India P. roxburghii is used for building
houses, packing cases, matches, music instruments, railway sleepers
124 etc.
Unit 6 Plant Anatomy and Embryology
ii) Sandal Wood : Sandal wood is smooth and tends itself to exquisite
used for carving as well as for extraction of the oil. The sandalwood oil is
fragrant and finds its use in perfumes, cosmetics, soaps, and incense
sticks etc.

SAQ 5
a) List the two types of systems and various cells found in secondary
xylem.

b) How do uniseriate, biseriate and multiseriate rays differ from one

another?

c) Distinguish between hardwood and softwood.

d) What structural features give strength to wood?

e) List as many independent uses of wood as you can think.

6.6 SECONDARY PHLOEM

In the secondary phloem there are also two systems- the vertical and
horizontal products from those cambial initials as in the case of xylem. Though
these two tissues secondary xylem and secondary phloem differ in ontogeny
and structure at maturity .The important components of the vertical system are
sieve elements, phloem parenchyma and phloem fibers (Fig.6.16).

Fig 6.16: Basic structure of phloem.

The horizontal system consists axial and ray phloem which is made up of
parenchymal cells. As in the xylem, the arrangement of the tissue in phloem is
primarily determined by the nature of cambium i.e. whether it is storied or not.
Secondly it depends upon the extent of elongation of various elements of
vertical system during the differentiation of the cells. 125
Block 2 Secondary Growth and Adaptive Features
In several species of dicotyledonous trees, growth rings can be observed in
the secondary phloem but they are less distinct than in secondary xylem. This
is because cells which are produced at the beginning of the season are
extended radially while in the end of the season they are flattened. After some
growth the arrangement of growth ring becomes obscured due to the
obliteration of the sieve elements. The primary reason is also because of
absence of lignin that is why they cannot be used as an indication of the age
of secondary phloem.

As you have already studied, that the ray initials in the cambium produce cells
towards both xylem and phloem. Thus xylem and phloem rays are continuous.
In the vicinity of the cambium the xylem and phloem rays are equal in size but
in many plants the mature outer portions of phloem rays are wider. The
widening of the phloem ray may be accomplished solely by the lateral
expansion of the existing cells or as is more common by an increase in the
number of cells on the periphery by radial divisions.

In the dicotyledons the functional secondary phloem is restricted generally to

the produced in the last growing season. In some cases when the cambium
starts producing new phloem, almost all the previously produced sieve tubes
cease to function. However, in Tilia sp. the sieve tubes are active for several
years including winter.

6.6.1 Economic Value of Secondary Phloem

The secondary phloem may be quite rich in secretory tissues because of its
role of protecting the plant. The phloem may contain well developed duct
systems in some species. It is the system of lactifers in the bark of Hevea that
is tapped to obtain rubber and the resin canals of the bark of conifers that are
harvested for pine resin which is further distilled to make turpentine and resins.

Bast Fibers: These are sclerenchyma fibers associated with the phloem of
certain stems of plants. They are rather easy to separate from underlying
woody tissues. They arise with primary tissues from the apical meristem or
with secondary tissues produced by the lateral meristem, the cambium.
Important sources of bast fibres are flax, jute, sunnhemp etc.

SAQ 6
Write T in front of true statement and F for false statement in the bracket
provided.

a) In phloem there are two systems the vertical and horizontal. [ ]

b) We can judge the precise age of tree by counting the number of rings of
phloem. [ ]

c) In several dicotyledonous species we can see more distinct ring in

secondary phloem than in secondary xylem. [ ]

6.7 SECONDARY GROWTH IN MONOCOT STEM

Normally in a monocot stem, no secondary growth takes place, as the
126 vascular bundles are closed i.e. cambium is absent. But in some herbaceous
Unit 6 Plant Anatomy and Embryology
and treelike woody monocotyledons plants belonging to families Liliaceae,
Agavaceae etc., the increase in thickness is through secondary cambium
(Fig 6.17) (which is entirely a secondary meristem).

Fig. 6.17: Portion of Dracaena stem showing secondary growth.

At the time of secondary growth some of the innermost parenchymatous cell

becomes meristematic; the cells divide tangentially forming a band of
secondary cambium, a few layers in thickness. The secondary cambium is
made up of rectangular fusiform cells. They do not produce secondary phloem
or xylem outside and inside respectively as in dicotyledonous stems and roots.
Instead, the secondary cambium cuts of secondary tissues on the inner side
first and then a small amount of new tissues on the outerside. These newly
formed inner secondary tissues directly differentiate into oval shaped collateral
vascular bundles and radially arranged parenchyma cells called conjunctive
tissue. Thus the vascular bundles remains embedded in the conjunctive
tissue.

In a monocot stem a periderm is absent but some storied cork cells, forming a
protective tissue with suberisation (Dracaena) are present.

6.8 PERIDERM
Before dealing with periderm formation we must know what periderm is? The
term periderm is collectively given to the protective tissues phellem and
phelloderm and the meristem that lies between them and gives rise to them.
This meristem, the phellogen, similar to vascular cambium is a uniseriate
layer of initial.

Cells of phellogen by periclinal divisions give off derivatives from their adaxial
and abaxial faces. A phellogen produces phelloderm adaxially and phellem
abaxially (Fig. 6.18). Phellem, a major constituent of periderm, is generally
suberised and inhibits translocation of water and solutes to tissue abaxial to it,
as a result, these tissues senescence and die. The cork tissue forms a
protective layer of the tree after the epidermis dies and is shed. Cork is 127
Block 2 Secondary Growth and Adaptive Features
generally formed in the stem and root of dicotyledons which have a continuous
and pronounced secondary thickening. Cork is not formed in leaves with the
exception scales of winter buds of certain plants. Cork is an important part of
secondary tissue which is termed periderm (Fig. 6.18). Peridem is usually
divided into three parts :

i) The phellogen- cork cambium

ii) The phellem-cork which is produced centrifugally by the phellogen.
iii) The phelloderm-a parenchymatous tissue in some species, and
produced centripetally by the phellogen.

6.8.1 Structure
Phellogen is a lateral meristem consisting of a single layer of initial cells. The
cells are uniform, rectangular in cross section with their shorter axis in the
radial direction. In a TLS they appear as regular polygons. The protoplasts of
phellogen cells contain vacoules of various sizes and may contain chloroplasts
and tannins. Phellogen has no intercellular spaces except in the regions of
lenticels.

The phellogen has distinct periods of activity and nonactivity. The activity
period may or may not coincide with that of the cambium. However, in some
plants two periods of phellogen activity have been noticed in a single annual
period of cambial activity.

Fig 6.18 (a-b): Formation of Periderm.

6.8.2 Phellem
Phellem or cork arises from the abaxial derivatives of phellogen. The cells of
phellem or cork are usually polygonal and radially flattened and divide
tangentially in a cross section. Cells are devoid of intercellular spaces except
in lenticular region. The cells of phellem divide tangentially. Cork cells are
dead. They may contain crystal containing cell, they may be sclerieds, or even
nonsuberised. In certain species the cork cell's primary walls are suberised
and contain a thick suberin layer interior to the primary wall called the suberin
lamella. This substance suberin is highly impervious to gases, water and resist
the action of acid. This phenomenon of impregnation of walls with suberin is
known as suberisation.

6.8.3 Phelloderm
The phelloderm cells are living cells with nonsuberised walls. They are similar
128 to the parenchyma cells of the cortex but, if the phelloderm is multiseriate, they
Unit 6 Plant Anatomy and Embryology
are usually arranged in radial rows. In some plants cells of the phelloderm
have chloroplasts and are photosynthetic.

6.8.4 Origin and Development of Periderm

The phellogen may differentiate in a living epidermal (Fig.6.19), collenchyma
or a parenchyma cell. Just before the onset of meristmatic activity the cell
loses the central vacuoles, the volume of cytoplasm increases and it
undergoes a periclinal division. Following the first periclinal division, two
similar cells are formed, of which the inner ceases to divide further. The outer
divides periclinally. The outer cell differentiate into the cork cell and the inner
constitutes the phellogen initial and continues to divide.

Fig. 6.19: Fully developed periderm.

The initials of phellogen cells occasionally undergo anticlinal divisions, to keep

pace with the increase in the circumference of the cork cylinder. The number
of phellem layers is usually greater than the number of phelloderm layers. If
the first formed periderm remains on the axial organ for many years, the other
layers of cork show cracks and are shed off. Thus the thickness of cork on a
plant remains constant.

6.8.5 Bark
The first formed periderm may be replaced by newer periderm. When such
replacement takes place, the last periderm is always produced inner to the
earlier (last formed) one.

This later produced periderm may have its origin in cortex, primary phloem or
even in secondary phloem. Generally, two types of formation of subsequent
periderms may be distinguished :

i) Plants where first formed periderm is developed in inner tissue, the later
formed periderm usually form an entire cylinder similar to the first formed
periderm. Such plants produce ring bark.

ii) Plants in which the first formed periderm originates in epidermis or outer
layers of cortex, the later periderms develop in the form of scales or
shells. The concave side of these scales is directed outwards. Such
plants produce scaly bark. 129
Block 2 Secondary Growth and Adaptive Features
Rhytidome : Whenever a new periderm is formed inner to the one already
present, the tissues exterior to it become cut off from the inner tissues. The
nutrition and water supply to them is cut off and such cells eventually die. A
hard outer crust develops out of such tissues. This crust increase in thickness
due to the additional cork tissue is being produced from beneath. All such cork
layer together with cortical/phloem tissues exterior to the inner most phellogen
are termed as rhytidome (outer bark) (Fig. 6.20).

All tissues exterior to the vascular cambium are included in the term bark. The
living part of the bank inside the rhytidome is often referred to as inner bark.

Fig. 6.20: Formation of periderm and rhytidome following the secondary activity
in a stem.

6.8.6 Commercial Cork

The source of commercial cork is. Quercus suber which occur naturally in the
The two types of cells
countries boarding the Mediterranean sea. In this species phellogen arises in
of cork may found in
one species e.g. the epidermis. It may remain on the plant for an indefinite period of time. For
Arbutus and Betula, commercial purpose the first formed periderm is removed when the tree is
where they occur in about 20 years old and about 40 cm in diameter. The exposed cells of the
alternating layers in phelloderm and cortex dry out and die, and a new phellogen is formed a few
Betula this feature millimeters within the cortex. The subsequent phellogen produces cork more
causes an interesting rapidly and in about 6 years a sufficient thickness to be of commercial value.
feature and cork is The stripped cork shows a rough outer surface and a smooth inner surface.
peeled off like sheets
of papers. The cork is valuable because it is impervious to gases and liquid, nonreactive
and has strength, elasticity and lightness. It is used for insulation, sound
proofing and in the manufacture of sports goods. Cork is unparallel as a
material for making stoppers for wine and Champange bottles.

You may like to know why commercial cork is to be cut in a particular plane.
130 Cork is several centimeters thick and the lenticels remain active for a long time
Unit 6 Plant Anatomy and Embryology
and result in the formation of cylinders of complementary tissue which extend
from the phellogen to the surface of the phellem. This complementary tissue
forms the patches of dark brown crumbling tissue found in commercial cork.
Because of the radial orientation of the tissues, bottle corks to be in a direction
parellel to the surface of the trunk. This way the cylindrical lenticels extend
transversely through them. Sheets of cork from the tree are rarely more than 3
cm. thick. Cork with a diameter greater than that cannot be obtained by cutting
in the usual manner. Large corks are usually cut from sheets of ground and
compressed cork or from "multiple sheets" composed of layers cemented
together. These kinds of cork are of low quality.

SAQ 7
Complete the sentences from section A with those given in section B.

Section A Section B

a) Periderm is usually i. consists only one type of initial cells

divided into

b) In some plants phellogen ii. three parts

has
a) Phellogen
b) Phellem
c) Phelloderm

c) Histologically phellogen iii. is impermeable to water gases and

can withstand the action of acid

d) Cork iv. alternating periods of activity and

inactivity

6.9 DISTRIBUTION OF LENTICELS

You must also know about lenticels. These are restricted area of relatively
loosely arranged cells; in the periderm. Lenticel protrudes above the periderm
because of their larger size and loose arrangement of numerous cells. There
is continuity of intercellular spaces of lenticels with tissue in the axial organs.

Lenticels are highly differentiated lens shaped areas of periderm. They are
usually found on stem and roots and appear on young branches or other
organs as rough dark patches. Due to many intercellular spaces, lenticels
have a loose structure. Mostly scattered over the entire surface of stem (Fig.
6.21).If we examine lenticel it usually looks like a convex lens both internally
and externally.

6.9.1 Development and Structure of Lenticels

Lenticels originate from localized regions in the phellogen that become
continuous with the nonlenticular phellogen. Lenticular phellogen has more
intercellular spaces and produces derivatives at a higher state than do
nonlenticular phellogen. The first formed lenticels generally appear below a 131
Block 2 Secondary Growth and Adaptive Features
stoma or group of stomata. Cells below the stomata begin to divide in different
Only a few plants e.g.
Philadephus directions and the chlorophyll in them disappear so that a loose colourless
Anabasis Haloxylon, tissue is formed. The cells that are derived from the divisions become more
Campsis radicans and more periclinal until the phellogen of the lenticels is formed (Fig. 6.21 a).
,Vitis and some other
species many of The cells which are derived from the divisions of the substomatal cells, as well
which are climbers do as those produced towards the exterior by the phellogen of the lenticels are
not possess lenticels termed complementary cells. After division the number of cells increases and
masses of complementary cells are pushed out and rise above the surface of
the organ (Fig. 6.21 b).

Fig. 6.21: a) Well formed Lenticel in L.S.; b) Lenticels on stem.

In the temperate regions lenticels become closed by the end of autumn

season by a closing layer. Whereas in some plants lenticels are formed
relatively early in the life of the plant and are shed together with the bark, in
others they may remain active for several years.

SAQ 8
Define the following in two or three sentences:

Lenticels and Complementary cells.

6.10 CAMBIAL VARIANTS

In this unit you have studied the function of vascular cambium and cork
cambium in dicotyledonous and monocotyledonous stems accounting for
development of secondary tissues. Cambium shows variation in its activity
132 giving rise to conditions which are rather typical. Various terms such as
Unit 6 Plant Anatomy and Embryology
cambial variants, anomalous or aberrant secondary growth has been used to
describe these instances (Fig. 6.22). As the variants are of quite regular
occurrence in certain plants, anatomists have discouraged the use of the term
anomalous secondary growth. Instead cambial variant has been
recommended for usage.

In normal conditions, in most of the dicot stems the vascular bundles are
arranged in a ring whereas in monocots they are scattered in the ground
tissue. The division of lateral meristem i.e. vascular cambium increases the
thickness of the stem. The division of the cambium results in formation of
secondary xylem towards inner side and secondary phloem towards outer side
or periphery. The cambium remains functional/ active throughout the life of the
plant and hence is referred as normal secondary growth. This occurs in most
of the dicot stems. In some dicots and monocots the structures appear
different from normal. Cambium shows variation in its activity. The terms used
to describe such condition is referred as cambial variants, anomalous (less
common) growth or aberrant secondary growth has been used to describe
this.

The anomalies may be present in the primary structure of the stem i.e. cortical
bundles, medullary bundles, and xylem are described as primary anomalous
structures. Some structures also develop during secondary growth such as
included phloem, accessory cambial strips. They are referred as secondary
anomalous structures.

Fig. 6.22: Cambial Variants a) Bignonia; b) Serjania; c) Bauhinia;

d) Boerhaavia.

6.10.1 In Stems
First we will study about the cambial variants found in stem.

I. The cambium is persistent and normal in position. Its products

show unusual arrangement and proportion.

a) In Bignonia (Fig.6.23) and some other members of the Bignoniaceae

family the cambium produces secondary xylem and secondary phloem
in different amounts. Thus in some part of the plant the amount of xylem 133
Block 2 Secondary Growth and Adaptive Features
is much greater than phloem while in the other, phloem is much more
abundant than xylem. This feature results a characteristically ridged and
furrowed xylem cylinder. Phloem can be identified by the presence of
wedges. There, as usually, four such wedges symmetrically arranged
and corresponding in position to the larger primary vascular bundles.

Fig 6.23: T.S. of Stem of Bignonia Showing anomalus secondary growth.

b) In some climbing species of genus Vitis (grapes) Clematis, Aristolochia

(Fig. 6.24) Tinospora etc. a complete ring of cambium is formed. The
fascicular cambium functions normally but the interfascicular cambium
produces only ray like parenchyma cells. As a result, broad and long
medullary rays and fluted vascular cylinders are formed.

134 Fig 6.24: T.S. of stem of Aristolochia.

Unit 6 Plant Anatomy and Embryology
II. The Cambium is abnormal in position but normal in activities

This situation is noted in some climbing species of Serjania (Fig 6.25). During
development, the cylinder of primary vascular bundles become notched at
certain points, so that groups of bundles are formed. Thus bundles become
constricted to form the cylinder. They may be cut off even at the procambial
stage. These groups of bundles function as independent cylinders and give
rise to separate cambia, each of which functions normally and independently.
Thus the stem appears as if it is made up of several discrete woody cylinders,
each of which has its own periderm. Sometimes the woody cylinder is only
lobed.

Fig 6.25 : T.S. of stem of Serjania

III. Anomaly due to the formation of accessory cambium and its

activity

a) In some species of Chenopodium and members of the Amaranthaceae,

the anomalous secondary growth results from accessory cambia. A
hollow cylinder of vascular tissue or a ring of irregularly arranged
bundles. The bundles are of secondary nature but their cambial activity
soon ceases. Just outside the bundles a new secondary cambium arises
in the pericycle. In some species the cambium forms tissues
centripetally, consisting of bundles embedded in nonvascular tissue. The
bundles formed in this way may be arranged irregularly or in defenite
concentric rings.

b) In Tecoma sp. secondary xylem and phloem are produced in the

beginning by the activity of a normal cambium ring. At later stage an
accessory secondary cambium arises in two arcs on the inner side of the
normal wood or towards the pith. This accessory cambium cuts off xylem
and phloem in an inverse order i.e. xylem towards the periphery and
phloem on the inside. This newly formed phloem is intraxylary phloem
and is secondary in origin. And the secondary xylem merges gradually
with the previously formed secondary xylem (Fig. 6.26). 135
Block 2 Secondary Growth and Adaptive Features

Fig.6.26: T. S. of stem of Tecoma diagrammatic and cellular.

IV. Anomaly due to the formation of interxylary (included) phloem as a

result of aberrant activity and position of cambium

Secondary phloem patches are sometimes embedded in secondary xylem in

the form of strands. Those extra patches of secondary phloem present inside
the secondary xylem are known as interxylary or included phloem.

a) In certain plants there are strands of secondary phloem within the

secondary xylem, e.g., in Avicennia, Thunbergia, Bougainvillea.
Salvadora (Fig. 6.27) and in the families of Amaranthaceae and
Chenopodiaceae. In these plants cambium differentiates outside the
primary vascular bundles, in the pericycle or in the inner cortical layers.
Later a series of vascular cambia arise successively outward, each of
which cambium produces xylem toward the inside and phloem towards
the outside until a new cambium develops from parenchyma cells on the
136 outside of the phloem.
Unit 6 Plant Anatomy and Embryology

Fig 6.27: T.S. of Salvadora stem diagrammatic and a portion of stem (cellular).

b) In Chenopodiaceae the successive cambia can be seen in the form of

long or short arches. They produce irregularly or spirally arranged
phloem strands (Fig.6.28) frequently the additional cambia in this family
form more or less entire rings.

Fig 6.28: Diagrammatic representation stem and a portion of the stem (cellular) of Chenopodium. 137
Block 2 Secondary Growth and Adaptive Features
6.10.2 In Roots
Cambial variants are also found in the roots of some plants, especially those
that serve a storage function :

a) In the beet roots Beta vulgaris first the cambium ring develops near the
primary xylem patches which in turn produces secondary xylem towards
inside and secondary phloem toward the outside. Soon its activity
ceases and then from the cells of pericycle and phloem a secondary
cambium ring is formed. This is followed by the formation of several
concentric cambia. All the cambial layers continue to function and
produce a large amount of storage parenchyma and strands of xylem
and phloem. Although cambia are in continuous rings they produce
separate bundles which are surrounded by conjuctive tissue. The
bundles are separated from one another by wide radial panels of
parenchyma formed due to the activity of newly formed cambium. Thus
alternate bands of proliferated pericycle and vascular bundles are
formed which can be seen as dark coloured and light coloured rings
respectively. The bundles are themselves largely parenchymatous with a
few lignified elements in xylem (Fig.6.29).

Fig 6.29: T.S. of root of sugar beet.

b) In the sweet potato root of Ipomoea batatas (Fig. 6.30) (Convolvulaceae)

the secondary growth is unique. The xylem, has an adundant amount of
parenchyma .Secondary cambia develop in the parenchyma around the
Individual vessels or vessel groups. The cambia produce tracheary
elements towards the vessels and sieve tubes away from the vessels; A
considerable amount of storage parenchyma is produced in both the
directions. Thus the phloem appears to be a portion of root that originally
138 differentiated as xylem.
Unit 6 Plant Anatomy and Embryology

Fig 6.30: T. S. of root of sweet potato.

SAQ 9
List various types of cambial variants and describe any one variant from root
and one from stem along with labeled diagram.

6.11 SUMMARY
• In general the plants having only primary growth are limited in size and
longevity. Secondary growth helps the perennial gymnosperms and
dicotyledons to increase the diameter and support the height and growth
size.

• After the procambium strand become differentiated into primary vascular

tissue, active meristematic regions lie between primary xylem and
primary phloem from which continued, addition of fresh tissues is
possible. These meristematic cells constitute the fascicular cambium.

• An interfascicular cambium arises from ray parenchyma cells between

vascular cambia.

• The fascicular and interfascicular cambium joins together and forms a

complete cylinder of vascular cambium. The divisions in cambium are
longitudinal, so that the stem now increases only in girth.

• In many plants, phellogen differentiates near the surface of the stem.

This gives rise to phellem and phelloderm. Cork cells have suberin in
their wall, which makes cell imprevious to gas and liquids. Lenticels in
the bark facilitate gas exchange.

• Cork cambium may originate in successively deeper tissues: epidermis,

cortex and phloem.

• Woody dicotyledons have most of their secondary tissues arranged in

concentric layers. The most conspicuous tissue is wood (secondary
xylem). The early wood usually has relatively large vessel elements,
while late wood has smaller vessels and/or a predominance of tracheids. 139
Block 2 Secondary Growth and Adaptive Features
• Generally an annual ring comprises one year's growth of xylem. The age
of a tree and other aspects of its ecological history can be determined by
studying histological details of the annual rings. Older wood ceases to
function and in gradually accumulated in the centre to form the
heartwood or dead core which becomes plugged with tylosis. The
younger, more peripherally located or living wood is called sap wood.
The scientist have shown that only one or two recently formed annual
rings or wood are actually involved in the ascent of sap.

• Some monocots such as Agave and Dracaena have true secondary

growth, with a type of cambium that produces secondary vascular
bundles and parenchyma.

• Certain dicotyledonous stem have cambial variants that contribute to

unusual secondary growth. Cambial variants may arise from the
following situations/conditions: i) The cambium is persistent and normal
in position. Its products show unusual arrangement and proportions. ii)
The cambium is abnormal in position but normal in activities. iii)
Formation of acessory cambium and its activity. iv) The formation of
interxylary phloem because of abnormal activity and position of
cambium.

• Cambial variants are also noted in some roots such as Ipomoea batatas,
Beta vulgaris etc.

6.12 TERMINAL QUESTIONS

1) What is secondary thickening (growth)? Explain the mode of secondary
growth in dicotyledons and monocotyledons. List the similarities and
differences in their secondary structures.

2) If a nail is driven into the trunk of a tree say at breast height, will it
remain at the same distance from the ground and more up or down in
the course of the next 6 years. Explain your answer.

3) What is periderm? Explain how it divides and the tissue it produces.

4) What are tylosis? How are they formed?

5) How is cork formed? Explain the structure, properties and uses of

commercial cork.

6) What are lenticels? How they are formed? What are their functions?

7) List various types of cambial variants .Describe the structural anomaly

arising as a result of formation of accessory cambium and its activity
provide suitable diagram?

8) Explain how dark coloured and light coloured rings are formed in the
beet roots.

140 9) Explain briefly the main features of unusual secondary growth in roots.
Unit 6 Plant Anatomy and Embryology

6.13 ANSWERS
Self-Assessment Questions
1. a) Secondary growth is growth in diameter due to addition of new
tissues due to activities of lateral meristems.

b) Secondary growth occurs both in intrastelar amd extrastelar

regions.

2. a) ray initials, fusiform initials

b) stress

c) resin

d) smaller

e) cambium

3. a) Fascicular cambium and interfascicular cambium.

b) Vascular cambium consist two types of cell: Fusiform initials and

ray Initials The fusiform Initials: are elongated cell with tapering
ends. They are found in tracheary elements, fibres, xylem and
phloem parenchyma and sieve elements. Ray Initials: are much
smaller than fusiform initials and are almost isodiameteric. They
have intense vacuolation, the cell wall possess primary pit fields
with plasmodesmata. The radial walls are thicker than the
tangential wall.

c) See section 6.3.2

4. a) False b) True c) True.

5. a) Secondary xylem can be classified in two systems: i) One system

is horizontal and ii) Other is vertical.

The horizontal system is made up of xylem rays and the vertical

consists of tracheary elements, fibres and wood parenchyma. The
living cells of rays are usually interconnected and a continuous
system of living cells is formed

b) i) Uniseriate Ray; when the ray is one cell wide it is called

uniseriate ray.

ii) Biseriate Ray: When two cell wide the ray is called biseriate
rays. On stem they are longitudinally or horizontally arranged
but most of the time they are scattered all over the entire
surface. Usually itlooks like a convex lens both internally and
externally.

iii) Multi seriate: When it is more than two cells wide.

c) In timber trade wood of dicotyledonous is known as hardwood and

the wood of gymnosperms as soft wood. But these words do not
express the degree of hardness. But histologically the
dicotyledonous woods have vessels while gymnospermous wood
lack them. 141
Block 2 Secondary Growth and Adaptive Features
d) The large part of the wood is made up of fibers or fiber
tracheids and they give strength to wood. These woods are dense
and heavy.

e) Woods have several uses which are listed below:

i) building material for houses in windows, doors, cabinets,

boxes, furniture.

ii) Building boats, ships, masts.

iii) Bodies of automobiles, railway wagons, railway sleepers,

bridges.

iv) Electric poles.

v) Various aspects of art are also made up of wood.

vi) Wood is a good material for carving idols, inlay work.

vii) Packing cases, matches.

viii) Several music instruments.

ix) Cosmetics, soaps, perfumes and incense sticks are made

from oil derived from woods.

6. a) True; b) False; c) False.

7. a) ii; b) iv; c) i; d) iii.

8. Lenticels: Restricted areas of relatively loosely arranged cells, in the

periderm. They protrude above the periderm because of their loose
arrangement, larger size of the numerous component cells. Lenticels
usually found on young branches of stem and roots. They are involved in
exchange of gases between internal tissues and the atmosphere
through periderm.

Complementary Cells: The cells which are derived from the divisions
of the substomatal cells as well as those produced towards the exterior
by the phellogen of the lenticels. As the division progresses, masses of
complementary cells are pushed out and rise above the surface of an
organ.

9. List the types of cambial variants from section 6.10 and give an example
from subsection 6.10.1 and one example from 6.10.2 along with diagrams.

TERMINAL QUESTIONS
1. In the dicotyledons the stem generally increases in girth due to the
activity of the vascular cambium. The growth is known as secondary
growth or secondary thickening. Also see section 6.2 and Fig.6.1.

2. Stem increases in height due to the activity of shoot apical meristem. So

if a nail is driven in trunk. It will always remain at the same height above
the ground. The nail may eventually become embedded as the stem
increases in girth and you can see the differences between the activities
142 of the apical meristem and of the vascular cambium.
Unit 6 Plant Anatomy and Embryology
3. Refer to section 6.8 and Subsection 6.8.1 and 6.8.2.

4. The outer part of the secondary xylem with living parenchyma is named
sapwood. In nearly all the trees the central portion consists of dead
parenchyma which ceases to conduct water. This is known as
heartwood. Heartwood is formed after disintegration of the protoplast,
loss of cell sap, hydrolysis of stored reserve materials and formation of
tyloses. Tyloses totally block the cell. Oils, gum, resins, tannins are
accumulated in the heartwood making it extremely durable. Thus
heartwood is preferred to sapwood for making furniture

5. Cork is an important part of secondary tissue which is termed as

peridem. It is divided into three parts. i) The phellogen cork cambium.
ii) The phellem-parenchymatous tissues in some species, produced by
the phellogen. iii) The phelloderm-parenchymatous tissues in some
species, produced by the phellogen.

Phellogen abaxially cuts off derivatives as phellem. The phellem or cork

cells are usually polygonal, and radially flattened. The cells are arranged
in compact radial rows which are devoid of intercellular spaces. Cork
cells are dead. Some cells are hollow and thin walled, and somewhat
radially rounded; some others are thick walled and radially flattened. The
later type of cells may be often filled with dark resiniferous or
tanninferous substance. Most of the commercial cork comes from
Quercus suber. In this species phellogen arises in the epidermis. The
first formed periderm is removed when tree is about 20 years old and 40
cm in diameter. A new phellogen differentiates a few millimeters within
the cortex. This cambium divides rapidly and in about 10 years it forms
sufficiently thick cork for commercial uses.

Cork is of commercial value because it is impervious to gases and

liquids and has strength, elasticity and lightness and also see
Subsection 6.8.2.

6. Refer to Section 6.9 and Fig. 6.21.

7. Refer to section 6.10. Sometime unusual secondary growth occurs due

to the formation of accessory cambia and their activity. There are two
types of vascular tissues i) a hollow cylinder; or ii) irregularly arranged
bundles. Their cambial activity ceases and outside a bundle in the
pericycle a new secondary cambium arises. The cambium forms tissues
centripetally, consisting of bundles embedded in nonvascular tissue. The
bundles may be arranged in concentric rings or irregularly.

8. In the beet roots first formed cambium activity ceases very soon. Then
there is formation of several concentric cambia. All the cambial layers
continue to function and produce a large amount of storage parenchyma
and strands of xylem and phloem. Though the cambia are continuous
they produce separate bundles which are surrounded by conjuctive
tissue. Bundles are also separated from one another by radial panels of
parenchyma. Thus alternate bands of proliferated pericycle and vascular
bundles are formed which is seen as dark coloured and light coloured
rings respectively Fig. 6.29.

9. Refer to Subsection 6.10.2, Fig. 6.29 and Fig. 6.30. 143

Block 2 Secondary Growth and Adaptive Features
Acknowledgements for Figures
Fig 6.17 : https://i.pinimg.com/736x/81/59/19/8159195fc8da6d027ff095
78e8ad27d8.jpg

Fig 6.25 : https://lh3.googleusercontent.com/proxy/W7Cel0pOZO4If_

http://virtualplant.ru.ac.za/Main/ANATOMY/serjania-
stem1.jpg

Fig. 6.27 : https://encrypted-

tbn0.gstatic.com/images?q=tbn%3AANd9GcRCPOylvfY3-
jJxjNpuxurToj3pBxPUvOVieQ&usqp=CAU

144