5th Week Chap 2.

Plant Tissues

1. Classification of tissues

a. Meristematic Tissues, zones in apical meristem.

b. Dermal Tissues
Meristematic Tissue of Plants: Introduction, Types and Theories (With Diagram)

Introduction:

Meristematic tissue, commonly called meristem, is composed of cells which are immature, not
fully differentiated ones, and which possess the power of cell division.

The vascular plants exhibit an ‘open system’ of growth involving formation of new tissues and
organs throughout its life. It is possible due to presence of certain organised regions where the
cells are in perpetually immature condition and can form new cells throughout.

These organised regions are the meristems. Primitive plants have cells all similar, carrying on
different functions. With advance in the line of evolution growth has been restricted to certain
parts of the plant body, the function of cell division being confined to the meristematic cells.

The open system of growth in plants is in marked contrast to the development of an animal body.

The meristematic cells have generally some distinctive features (Fig. 527). The cells are usually
isodiametric in shape; they are compactly set without evident intercellular spaces; have dense
cytoplasm with small vacuoles and large prominent nuclei; ergastic matters are lacking; the
plastids are in proplastid stages; cell wall, made of celluose, is thin and homogeneous.

Though in general, meristematic cells possess the characters stated above, but departures cannot
be ruled out. The meristematic cells of vascular cambium are fusiform in shape (Fig. 527 B & C)
often with conspicuous vacuoles.

Comparatively thick walls with primary pit fields are found in some meristematic cells. Ergastic
matters like starch and tannins may also be present. Considering these departures some authors
are in favour of using the term eumeristem or true meristem for those having the general
characters stated above.

It has been stated that meristems are the formative regions where new cells are added to the
body. Besides, the meristems perpetuate themselves.

The cells which remain meristematic and thus continue cell division are called initiating cells;
whereas the cells formed by them, the so-called derivatives, gradually change their shape,
enlarge, lose the power of cell division and ultimately become mature cells with some definite
characters and functions.

These changes involving enlargement and specialisation represent the process which may be
referred to as differentiation. The differentiation of a particular tissue has been characterised as
“a progressive loss of embryonic features of meristematic cells and the progressive attainment of
the state of maturity” by a well-known authority.

The derivatives of the initiating cells gradually differentiate into mature cells and lose the power
of cell division, at least temporarily. These completely differentiated cells are called permanent
ones. Some permanent cells may get back the meristematic nature, a phenomenon which has
been referred to as dedifferentiation by some workers. That shows that though cells have attained
permanent form, but they have retained the power of cell division.
The living parenchyma cells and the epidermal cells are common reversible permanent cells. In
fact, all living cells retain the potentialities of division and growth, though they have become
permanent. Strictly speaking, permanent cells are those which have completely lost the capacity
of division, as for example, irreversibly specialised cells like the sieve tube elements and the
dead elements like tracheids and cork cells.

Various systems of classification of meristems have been proposed, based on characters like
origin and nature of initiating cells, stages of development, topography and function. No system
is exclusive and rigid.

Types:
The common important types of meristems according to their origin and development are
the following:
Promeristem or Primordial Meristem: Promeristem is the very foundation stage, the region of
formation of new organs and tissues. It may be called the earliest embryonic condition consisting
of young initials and their derivatives.
All the cells possess the characters of true meristematic cells, viz., diameters alike, dense
cytoplasm, large nuclei, proplastids, absence of ergastic matters and thin walls. Promeristem is
definitely of limited extent. As soon as the cells begin to show tendency of differentiation, they
have passed the earliest promeristematic condition.

Primary Meristem: Primary meristems are composed of cells which are direct descendants of
embryonic cells and which have all through retained meristematic nature. So primary meristem
may be called a later developmental stage.
Chief primary meristems are those located at the tips of stem, root, and appendages. Primary
meristems build up the primary body of the plants.

Secondary Meristem: Primary meristems gradually differentiate into permanent tissues. Some
living permanent cells may regain the power of cell division. They constitute the secondary
meristem, as they originate from permanent cells.
The cork cambium or phellogen, arising from epidermis, cortical and other cells during
secondary increase in thickness, is an example of secondary meristem.

It is to be noted that primary meristems are responsible for the building up of the primary body
of the plant; and the secondary meristems, formed later, add new cells to the primary body with
definite purposes like effective protection and repair.
The definitions given above are not always accurate. In case of adventitious organs the apical
meristem develops within permanent tissues secondarily though they are primary in structure and
function.

According to their position in the plant body meristems are put into three groups, viz., apical,
intercalary and lateral (Fig. 528).

Apical Meristem: Apical meristems occur at the apices of the stems, roots, main and lateral, of
the vascular plants. Growth in length of the axis is entirely due to their activities; so they are also
called growing points. The initiating cells may be solitary or in groups.
Solitary initiating cells go by the name apical cells, whereas those occurring in groups are called
apical initials. Solitary apical cells (Figs. 530 & 533A) occur in many pteridophytes like ferns
and horse-tails.

Other vascular plants possess apical initials which may be terminal or terminal and sub terminal
(Figs. 532 & 533). The apical initials may occur in one or more tiers.

In case of one tier all the cells of the plant body derive their origin from it; whereas if there is
more than one tier different parts of the plant body originate from different tiers of initials.

The derivatives of the apical meristem differentiate in course of time into permanent tissues
which together constitute the primary body of the plant.

Intercalary Meristem: These are the portions of apical meristems which are separated from the
apex during the growth of the axis and remain intercalated between permanent cells.

The position is such that apical meristems go ahead during development and a portion of it is left
behind which is inserted between permanent cells.

Intercalary meristems are found in the stem and leaf sheaths of many monocotyledons,
particularly grasses, and in the horse-tails. Here nodal regions are usually composed of
permanent cells and so intercalary meristems are internodal. These meristems are short-lived,
they very soon become permanent and merge with the tissues surrounding them.
Lateral Meristem: These meristems occur laterally in the axis, parallel to the sides of the organs
in dicotyledons and gymnosperms. They are composed of initials which divide periclinally. The
derivatives gradually differentiate into permanent tissues called secondary tissues.
These tissues are added to the existing ones and are responsible for increase in thickness. The
growth in thickness thus secured due to addition of secondary tissues is referred to as secondary
growth.

The cambium of the vascular bundles and the phellogen or cork cambium are lateral meristems.
It should be noted, however, that cambium is a primary meristems, whereas phellogen or cork
cambium is secondary in origin.

A classification of meristems chiefly based on function was followed in physiological anatomy.

This classification is helpful in tracing the three tissue systems of mature regions—epidermal,
ground or fundamental, and vascular, consisting of epidermis, ground tissues and vascular tissues
respectively. The advantage of this set of terminology from topographical point of view is
undeniable, though the interpretation of development of shoot and root from these zones is a
quite complex problem.

Meristems on the basis of planes of divisions are of three types, viz., mass meristem rib
meristem, and plate meristem. Mass meristems grow by dividing in all planes, so that the bodies
formed are either isodiametric or have no definite shape.

This pattern of growth is noticed in endosperm, young embryo and also in the formation of
spores and sperms. The rib meristems, also called file meristems, divide anticlinally to the long
axis and give rise to longitudinal files or rows of cells.

This growth pattern is clear in the development of cortex and pith. The plate meristems divide
chiefly anticlinally in two planes, so that new cells are formed but number of layers does not
increase.

The uniseriate epidermis and multiseriate flat blade of the leaf are illustrations of growth forms
by plate meristems.
Theories of Structural Development and Differentiation:
The classical concept of the apices of shoot and root had been that they represent the earliest
self-perpetuating promeristems consisting of homogeneous eumeristematic or truly meristematic
cells.

Modern workers are of opinion that a distinct zonation is exhibited by the immediate derivatives
of the promeristem. By zonation is meant the existence of distinct regions differing from one
another by characters like nature of cells, plane of cell division, position of the initiating cells
and rate of maturation of cells.

The growth and development of the apical meristems of shoot, root, and flower and their
differentiation have been most notable problems of plant anatomy.

Intensive studies on those problems have been going on since the nineteenth century and a few
theories have been proposed from time to time. A brief review of those theories is being given
here.

Apical Cell Theory: The solitary apical cells (Figs. 530 & 533A) were discovered in the
cryptogams—in algae, bryophytes and pteridophytes. This led early workers to believe that a
solitary apical cell was a constant unit of apical meristem governing the whole process of
growth.
They assumed that the same condition prevails in all higher plants. On that assumption apical
cell theory was advanced by Hofmeister in 1857 and supported by Nageli (1878) in the
nineteenth century.

Subsequent works revealed that complex apices of gymnosperms could not be interpreted in the
light of this theory. So it was not applicable to seed plants.
Histogen Theory: The older theory was replaced by histogen theory as an attempt towards
interpretation of the growing points of seed plants. Hanstein proposed
this theory in the nineteenth century and received support from Strasburger (1868).

According to this theory plant body develops from groups of initials forming a mass of meristem
of considerable depth, as opposed to superficial apical cells of former theory, and that three
distinct zones or strata can be recognised at the growing points of stem and root.

Every zone consisting of a group of initials was called a histogen or tissue builder. The histogens
arise from separate sets of initials and have different courses of development.

The three histogens (Fig. 531) suggested were (i) dermatogen, the outermost uniseriate layer; (ii)
plerome, the massive central core consisting of cells extended in longitudinal direction; and (iii)
periblem, the region composed of isodiametric cells lying between the dermatogen and the
plerome.

The dermatogen cells divide anticlinally and develop into uniseriate epidermis. The periblem
forms the cortex; and the plerome gives rise to the massive central cylinder or stele, consisting of
primary vascular tissues and ground tissues like pith, pith rays and pericycle.

Hanstein’s histogen theory as a basis of interpretation of the shoot and root apex held ground for
fairly long time. Though it is followed even now in case of root apex, it has been found to be
inadequate as regards shoot apex of angiosperms mainly for two reasons, viz. (i) sharp
distinction between dermatogen and periblem is absent; and (ii) origin of different regions from
the sharply defined histogens cannot be demonstrated.

Moreover in many gymnosperms and angiosperms shoot apices hardly show any distinction
between periblem and plerome.
Tunica-corpus Theory: As an interpretation of the apical growth in the shoot apex the third
theory—tunica-corpus theory—was proposed by Schmidt and supported by Foster and others in
the early part of the twentieth century.
According to this theory, two tissue zones occur at the apex. They are the tunica or cover,
consisting of one or more layers of cells forming the outer enveloping region, and the corpus, the
central core, a mass of cells surrounded by tunica (Fig. 532).

The two regions differ in structure and appearance due to varying rates and methods of growth.
The cells of tunica are smaller and lie in layers with planes of cell division predominantly
anticlinal.

So growth here is primarily in area. The corpus cells are larger and they divide in various planes,
so that a mass of irregularly arranged cells is formed. Increase of the mass here is thus in volume.

The tunica may be one-layered or many-layered with massive or slender corpus. The number of
initials is also variable. In fact, in lower vascular plants like pteridophytes and even in some
gymnosperms a sharp distinction between tunica and corpus cannot be traced.

In angiosperms, however, two sets of initials may be definitely located, the two sets giving rise
to tunica and corpus. Thus the two have independent origin.

In spite of the fact that many variations and fluctuations occur, the tunica-corpus theory has
given a renewed impetus in the interpretation of the apical growth in the stems of angiosperms.

As a result, the stages in the development of the primary body from the initiating cells are better
understood.
The Epidermal Tissue System of Plants (With Diagrams)

Epidermis: This system solely consists of the outermost skin or epidermis of all the plant
organs beginning from the underground roots to the fruits and seeds.
This layer represents the point of contact between the plants and the outer environment and, as
such exhibits diversities in structure.

It is primarily a protective tissue, which protects the internal tissues against excessive loss of
water by transpiration and mechanical injury. Subsidiary functions like storage of water,
mucilage, secretion and, though rarely, even photosynthesis, may also be carried on.

But for stomatal and lenticular openings the epidermis is a continuous layer. Normally it is
uniseriate—typically consisting of one layer of cells. It derives its origin from the protoderm of
the meristematic region.

The protoderm cells divide anticlinally and in course of time uniseriate epidermis is formed.
Many-layered or multiseriate epidermis, usually called multiple epidermis, is found in some
organs like roots of orchids, leaves of Ficus spp. (Fig. 507A), Nerium, Peperomia, etc.

Normally it may be assumed that these layers have originated from the protoderm by periclinai
divisions. The outermost layer of multiple epidermis is similar to ordinary uniseriate one. The
inner layers are different from other tissues in absence of chlorophyll.

There is room for doubts if all these layers belong to epidermis from ontogenetic point of view.
They may be outer layers of cortex originating from the ground meristem, but resemble the
epidermis both in structure and function.

The epidermal cells are living with lining layer of protoplast around large central vacuole. The
plastids are normally small and colourless. Chloroplasts are present only in the guard cells of the
stomata in case of organs exposed to sunshine, but they occur in the epidermal cells of aquatic
plants and plants growing in moist and shady situations. Mucilage, tannins and crystals may
occasionally be present.

Anthocyanins may occur in the cell-sap of the vacuoles. Epidermal cells retain the potentiality of
cell division. During normal course of development or due to external stimuli they may divide
and produce new cells.

Epidermal cells exhibit wide diversities as regards their size, shape and arrangement. But they
may be said to be essentially tabular in shape (Fig. 555 A & B) compactly set, so that a
continuous layer without intercellular spaces is formed. Only in the petals of some flowers
intercellular spaces are found, but they remain covered by outer cuticle. In surface view they are
more or less isodiametric in shape.

In the leaves and petals of flowers they may have irregular shapes, often with teeth and flanges
(Fig. 555 C & D) which remain peculiarly interlocked with one another. In monocotyledonous
stems and leaves with parallel venation the epidermal cells are rather elongated in the direction
of the long axis (Fig. 555 E), so much so that in extreme cases they may be fibre-like in
appearance.

Epidermal cells have unevenly thickened walls, the outer and radial walls being much more thick
than the inner walls. In some cases they may be so massive that the central lumen is almost
obliterated. The walls are strongly cutinised, what is very important for protection against
mechanical injuries and prevention of loss of water.

The fatty substance cutin is found in the wall—in interfibriller and intermicellar spaces of the
cellulose and forms the cuticle occurring all over the outer wall of the epidermal cells (Fig. 556
A&B).

It remains as a separate layer and in some cases it may be removed as a whole. The cuticle is
often found to project into the radial walls as peg-like bodies (Fig. 556C). Cuticle is absent only
in the epidermis of roots and some submerged aquatic plants.
The thickness of the outer walls of the epidermal cells depends on the environmental conditions
of the plants. It is quite thin in plants with adequate water supply, and it is unusually thick in
plants growing in dry situations.

The surface of the cuticle may be smooth or may possess ridges and cracks. The cutinised
portion of the walls, the portion lying beneath the cuticle, has been found to consist of alternating
layers of cutin and pectic materials.

Waxy matters are often deposited on the cuticle in form of rods and grarules (Fig. 556E). The so-
called ‘bloom’ of many fruits and glaucous characters of many stems and leaves are due to these
deposits. Lignification is rather rare in epidermal cells.

It occurs in the pine needles (Fig. 556D), in cycad, in grass leaves outside the sclerenchyma
patches and in a few dicotyledons. Deposition of silica is common in the epidermal cells of
horse-tails (Equisetum) and grasses.

It is really interesting to find long epidermal cells having corrugated margin (Fig. 555E)
associated with two kinds of short cells—the silica cells and cork cells in grasses. The silica cells
contain silicon oxide and cork cells with suberised walls contain organic materials.

In some dicotyledonous families like Malvaceae, Rutaceae, etc., the epidermal cells individually
or in groups undergo mucilaginous changes, particularly in the seeds. Special sac-like cells
remain scattered in the epidermis of some members of family Cruciferae.

These are idicblastic cells resembling the lati ciffers, but they contain an enzyme, myrosin, and
so they are called myrosin cells. It has been stated in a preceding chapter that many
dicotyledonous families like Urticaceae, Moraceae, possess cystoliths.

The cystolith-containing cells of epidermis are referred to a lithocysts. The epidermis is often
made up of a layer of sclereids, as found in the seed-coats of Pisum and Phaseolus of family
Leguminosae (Fig. 537D) and in the scales of garlic—Allium sativum of family Liliaceae (Fig.
537E).

Radial and inner walls of epidermal cells possess pit-fields. Plasmodesmata have also been
reported, those on the outer walls of leaves have also been called ectodesmata.
The origin of the shoot epidermis may be traced from the apical meristem. It arises from the
outer layers-of tunica, according to tunica-corpus theory, or from the dermatogen of Haustein or
protoderm, as suggested by Haberlandt, which may be called primordial epidermis.

But the root epidermis fundamentally differs from that of shoot in origin, structure as well as in
function. It has been pointed out in the previous chapter that epidermis of root is related to the
root-cap or the cortex from the developmental point of view.

The cells are tabular, lack in cutinisation of wall and their function is mainly absorption of water
and solutes. So the terms epiblema, piliferous layer or rhizodermis have been applied to it.

Epidermis, as a rule, persists as uniseriate layer throughout its life in the organs where distinct
secondary growth does not take place. In some monocotyledons, though secondary increase is
absent, a kind of periderm is formed, and thus the epidermis is destroyed.

In organs with distinct secondary growth in thickness epidermis continues till cork cells are
formed. In leaves, flowers and fruits, it persists as long as the organs do. In roots the epidermis
with a part of cortex becomes dead, lignified or suberised after the root hairs are destroyed.

Bulliform Cells: In the leaves of monocotyledons, excepting a few families, a peculiar type
of comparatively larger, highly vacuolate and thin-walled cells occur in the epidermis.
These are called bulliform (meaning, bubble-like) cells.
In transverse section they appear as a fan-like band because the median cell is usually the largest
in size (Figs. 557 & 557A). They may be present on both sides of a leaf, but are more common
on the upper side running parallel to the veins.

They either cover large areas or remain restricted to the grooves. These are mainly water-
containing cells with no chlorophyll. The walls are usually thin, but the outer walls may be thick
and cutinised like other epidermal cells, often filled with silica.

There are three views as regards the functions of bulliform cells. According to the first view they
are concerned with the unrolling of the developing leaves. It is suggested that these cells undergo
sudden and rapid expansion at a certain stage of leaf development and consequently bring about
unfolding of the leaves.

The second view is that they have a role to play in the hygroscopic opening and closing
movements of mature leaves, due to changes in turgor. They have also been called motor cells by
workers holding the above view. The third view is that they are simply concerned with water-
storage and have no other function.
Epidermal Outgrowths: Outgrowths of diverse forms, structures and functions develop
from the epidermis. All these appendages which are epidermal in origin, are referred to as
trichomes.
Thus they are different from the emergences like the prickles of roses, as the latter are formed by
epidermis and a part of cortex. Trichomes may occur on all parts of the plant body.

Some of them persist throughout the life of the organs, whereas many of them are ephemeral
bodies. They may remain alive or become dead and continue as such. Trichomes have been put
into a number of groups on the basis of their morphological characters.

(a) Hairs: Hairs constitute a very common type of trichome. They may be unicellular or
multicellular. Unicellular hairs are often simple unbranched elongated bodies or they may be
branched.

Some of them are very much elongated and twisted, so that they have woolly appearance (Fig.
564-C). Multicellular hairs may be formed of one row of cells (Fig. 564 A, D, E & F) or of many
layers as found in the base of petiole of Portulaca.

Often these hairs branch in very peculiar fashions; some of them assume dendroid or tree-like
appearance (Fig. 564 G & H), or the branches come out in one plane giving it stellate or star-like
shape.

They are also called stellate hairs (Fig. 564 I), A multicellular hair has usually two parts, the
basal part which remains embedded in the epidermis is the foot and the other which projects out
is the body. An initial cell divides periclinally into two parts, of which the outer one forms the
body and the inner one, the foot.

(b) Scales or Peltate hairs: These hairs consist of disc-like plate of cell (Fig. 564 J) put on a
short stalk or directly attached to the foot.
(c) Colleters: These are glandular trichomes. Some hairs have multicellular stalk and head, the
latter is composed of glandular cells (Fig. 565). Sticky exudations present on the surface of
certain leaves and buds are secreted by colleters.

Salt-secreting glands as found in Tamaricaceae and calcium- secreting glands of Plumbaginaceae

(d) Water vesicles or bladders: They form a very interesting type of trichome where some
epidermal cells become greatly distended and serve as water reservoir.

They occur in the so-called ‘ice-plant’ (Mesembryanthemum crystallinum of family Aizoaceae)

Those occurring in Artiplex, also called vesiculate hairs, dry up with maturity and persist as a
white layer on the leaf surface (Fig. 565A).
The walls of trichomes are commonly of cellulose covered by cuticle. They sometimes remain
impregnated with silica and calcium carbonate. Trichomes other than glandular ones have highly
vacuolated protoplast. The cotton fibres, which are really hairy outgrowths from the seeds, have
secondary walls of almost pure cellulose. The stinging hairs of nettle (Urtica dioica) possess a
peculiar type of wall structure for releasing the contents of the gland.

The hair (Fig. 244) resembles of fine capillary tube with silicified upper end and calcified lower
end. The base remains embedded in the epidermal cells. Coming in contact with the skin the tip
breaks at a predetermined point and the sharp edge penetrates into the skin when the contents
(histamine and acetycholine) are injected, so to say, to the wound.

Root-hairs: As already reported the root epidermis fundamentally differs from shoot
epidermis in origin and in absence of cuticle and stomata. But it bears hairs at a particular
zone. Unlike the hairs and trichomes discussed above, the root-hairs are not outgrowths or
appendages, but they are prolongations of the epidermal cells. During the formation of
root-hairs, growth in length of the epidermal cells is checked.
It has been found in some plants that root epidermis possesses two types of cells, short cells and
long cells due to unequal division, and the hairs are formed from the short ones (Fig. 567) which
are called trichoblasts.

It comes out as a protuberance, continues elongation and thus the hair is formed. It has vacuolate
protoplast and the nucleus moves on to the tip. The wall is thin, composed of cellulose and pectic
materials. Root-hairs are short-lived bodies.

During the growth of the root, old hairs are destroyed and replaced by new ones. They are
responsible for the absorption of water and mineral solutes from the soil.