CHAPTER 5

Tissue System
5.1 Definition: Combination of one or more tissues performing the
in the
COmmon physiological function regardless of position or continuity
Plant body is known as tissue system, which is structurally and functionally
organised into a unit.
5.2 Classification and the origin of tissue system: Various anatomists
have segregated the tissues ofthe plant body into tissue systems on thebasis
oftopographical continuity or physiological similarity" (Foster, 1949). Sachs
(1875), on the basis of the topographical continuity of tissues, recognised
three principal systems of tissues in the plant body, viz. (a) the epidermal
tissue system which includes epidermal and cork layers, (6) the fascicular
1.e. vascular tissue system which contains conducting tissues like xylem and
phloem of vascular bundles and (c) fundamental tissue system which
includes all the tissues other than fascicular (i.e. xylem and phloem) and
epidermal.
On the basis of physiological function Haberlandt (1914) in his
Pbysiological Plant Anatomy" treated tissue systems as the (a) mechanical
System, 6) absorbing system and (c) storage system. Here, in agreement
with Sachs, the main tissues of a vascular plant are grouped into three tissue
Systems, e.g. (a) epidermal tissue system which includes the outer covering
of the plant body in primary state i.e. epidermis, (b) fundamental or ground
tissue system which includes tissues forming the ground substance of the
plant body i.e., tissues other than epidermal and vascular tissues and
Cvascular tissue system which includes complex tissues like xylem and
phloem of vascular bundles only.
The origin of ditferent tissue systems and their relations between apical
meristems and primary permanent tissues are shown as follows
PRIMARY APICAL PRIMARY PERMANENT
MERISTEM TISSUE
TISSUE SYSTEM
--PROTODERM EPDERMIS- EPIDERMAL
TISS
SYSTEM

CORTEX
GROUND
PROMERISTEM MERISTEM PERICYCLE
MEDULLARY
RAYS
GROUND TISSUE
SYSTEM
PITH
STELE

-PROCAMBIUM VASCULAR-
BUNDLE VASCULAR TISSUE
SYSTEM
53 Epidermal tissue system: This tissue system is formed
by the
enidermis and its associated structures. Epidermis constitutes the outermost
1aver or layers of cells of all parts of the primary plant body such as
stems, roots, leaves, i1owers, fruits and seeds. Epidermis forms a continuous
laver or layers except interrupted by stomata or lenticel openings.
 TISSUB SYSTEM 257

The epidermis of stems, leaves and floral parts originates from the surface
laver of shoot apical meristem (protoderm) and that of the root originates
Crom a layer of cells in the root apical meristem which is covered by the
from
root cap. Sometimes, in different plants, the epidermis may have a common
rigia with the cortex or with the root cap..
The epidermis usually consists of only one layer of cells in thickness.
But in some cases, cells of this layer may divide further periclinally to give
or
rise to a several-layered structure-such a structure is called multiple
multiseriate epidermis. Multiple epidermis is found in the leaves of Ficus sp.
Nerium sp. (Apocynaceæ), members of Piperaceæ etc.
(Moraceae),

CUTICLE

a leaf in surface (A)

epidermal cell1s. A-B-Ordinary epidermal cells from
Flg. S.1-Types
of
views (B). C--Epidermal cellsfrom
the leaf of Solanum tuberosum
and sectional cells from the petal of Viola sp.
in surface view. D-Epidermal
of aerial roots of epiphytic
The velamen, special absorbing tissue as it arises from the tangential
a
a multiple epidermis
orchids, is practically or derma BULLIFORM CELLS
divisions of protoderm be-
later all the cells
togen cells; protoplasts shrivel
come dead, the air.
the cells contain
up and dry, orchid roots
The velamen of such
forms the first covering layer
and
is to absorb
the function of which the
in atmos-
water vapour present of
i.e. such type
phere. Velamen present not
multiple epidermis isin
orchids but the acrial
only in members of
Araceae.
roots of many
velamen looks like a
Bxternally
spongy whitish coat. showing
leaf bulli-
of the
Fig. 5.2-T.s. in grass
upper epidermis.
According to DeBary (1877), form cells
layer of tissue.
the cells of the innermost
of leaves usually
function as a water-storing
the multiple epidermis oval and closely
cells are tabular or still
epidermal
Though normallycontinuous layer without any intercellular spaces,
and shoot,
Compact, forming a In both root
in their size, shape and arrangement.
they vary
258 STUDIES IN BOTANY

surfaco of organ tothe gIvo

at an anglo to the
r n a l cells may elongate plant parts liko stems,
potioles eto. eplderat
S toarehairs. In elongated longitudinal axis of sueh parts.

els elongated and parallel in relation to have wavy walls. Intercellular

and leaves, the epidermal cells some flowers, but they ar
n uower petals epidermal cells of petals of in
found in anapo
Opcesare cells are isodiametri in
COvered by outer cuticle, Epidermal of flowers they are irregular cino m shapo
but in leaves and petals
of silica is
mon
and ICW,
and often provided with teetb and flanges. Deposition
LONO CELL CORN CELLS
LONG CELLSILICACELL
HAIR
SILICA CELL CORK CELL BRISTLE STOMA

A B
Eis. 5.3-Epidermal cells of Saccharum oficinarum(sugarcane) in surface view.
A-Epidermis ofstem; B-Lower epidermis from a leaf blade.
t h e epidermal cells of grasses (Gramineæ) and horsetails (Equisetum
in the epidc"Gramineæ, long
In membets epidermal cells are associated with two types
silica cells and cork cells (Fig. .3, A). These two types of
often occur together in pairs. The silica cells are almost filled
short cells of various shapes.
with SiO, which solidiles into structures organi Snapes. The co c d
suberised walls and often contain solid
have suDether
have kind of enlarged, thin-walea and highly vacuolated cells oceur
grass cells-these. s occur
or motor cells (Fig. 5.2) whi
(b) hygroscopic movements of r helpful
cells are in (a) 1
as bulliform
ofdeveloping
leaves,
water storage tissue. mem aves and C) 8in
become elonoa 1n some
Ome membersrs of
Sac. of Cruciferae.
the e p i d e r m a l cells which c o r e d and
s o m e of cells cells,
as myrosin nzyme myro
they a r e known cells are made up of sclereide 8. epidermOSin, So:
times epidermal epidermal cells may also cont rystals, e. ells of Pis
seed coat. Some are formed in the epidermis
cystoliths
in which
In many grasses, the specialised cells in the root epidermis which
trichobla5
rise to
root hairs i.e.
cytoplasm and othes
give
from the adjacent
cells by
their
dense
epidermal living,
the cells have
a
productscentral
mal cells.contain vacuole
metabolism and
various
like peripheral
p r o t o p l a s t ; cells may of
mucilage
 TISSUE SYSTEM 259
tannins and crystals of various salts.
Leucoplastids
chloroplastids are present only in guard cells
cells stomata andgenerally
of stom.re
are generally present,
in the epidermal
of
cells of some pteridophytes. hThe Buard
cell sap contains anthocyanin pigments, e.g.
petals of
petals of flowers.

cUTINISED LAYER
CUTICLE PEG
WAX

A
CELLULOSE LAYERS
CUTICLE

Fig. 5.4-Types of epidermal cells: from leaf blade

of Aloe sp. (A) ;
sp. (B);from leal of Allium cepa (C); from needle of Pinusfrom leaf blade of
sp. (D); from leaf
Musa
blade of Saccharum sp. (E).

The epidermal cell walls vary in thickness either in

or in different plants ; different plant parts
some cells are thin-walled while some cells have thick
outer walis. Primary pit fields are
often found in the walls, specially on the
radial and inner walls of the epidermal
cells. In the outer epidermal wall of
many leaves, e.g. Chrysanthemum, Antirrhinum, Hexine, Allium etc. plasmodes
mata are present, such
plasmodesmata of leaves are called ectodesmata.
outer exposed walls of the epidermal
cells of aerial plant parts is thick The
to the deposition of a fatty substance called cutin. due
This substance i.e. cutin
remains deposited in the outermost cell wall layer,
special layer called cuticle (Fig. 5.4, A). The cuticle and
thus constitutes a
is often found to project
into the radial walls as peg-like bodies
of several lamellae. The lamellae (Fig. 54, B). The cuticle layer consists
to cutin. Deposition of
contain cellulose and pectin in addition
wax in the form of granules or rods
or as continuous
layer iS often found on the surface of the(Fig. 5.4, B, E),
cuticle. The
Surface of the cuticle may be smooth,
Uniformly thick-walled epidermis is seen in the rough,
leaves of
ridged or furrowed.
conifers, here the
cells are lignified (Fig. 5:4, D) and the
obliterated. Thin-walled and non-cuticled
lumens of the cells are practically
non-aerial plant parts like roots and in epidermal cells are seen in
submerged
thin-walled epidermis of roots developing from the root aquatic plants-such
covered by root cap (not from apical meristema
epiblema protoderm as
in case of stem) is called
or piliferous layer. Lignification i.c.,
epidermal cell walls is also noted in some grassdeposition
of lignin in the
leaves and in
Quercus, Nerium etc. species of
The epidermis may persist as uniseriate layer
plant where distinct secondary growth does not take throughout the life of the
place, or in plants which
undergo secondary growth, it may be sloughed off along with the underlying
tissues after the formation of the periderm.
260
STUDIRS IN BOTANY
internal
Functions Epidermis is mainly a protectvercessive it protectswater
the protects

internal tissues against mechanical injury and checkstissue,

excessive
ininrOective
loss of water
during transpira st mechanical inju mis
echanical
in water storage,
Secretion, absorption
absation
secondarily
(epiblema of root) and serves
photosynthesis in some cases,
e.g. o n ,

A. EPIDERMAL OPBNINGS
U Stomata (Fig. 5.5, A-B)-The continuity of the epidermis is
terrupted by minute pores openings call e specially on leaves
or
).
tomata occur on most of the green
aerial plant Panoists of a pore bounded
and young stems; roots lack stomata. A
o two
specialised epidermal cells called guardstoma och by their changes
cells, W

SUBSIDIARY CELLS
CEPIDERMAL CELLS) oUTER LEDGE GUARD CELLS
SUBSIDIARY CELLS

STOMATAL
APERTURE
-GUARD JNNER LEDGE
CELLS
SUBSTOMATAL
CHAMBER

CHLOROPLASTS
NUCLEUS

Fig. 5.5-Structure of stomata in surface (A) and sectional

views (B).
in shape bring closing and opening of stoma.
about
is the stomatal aperture The
pore of
the stoma
guard
through which gaseous
cells together with the stomatal exchange takes place. The
aperture
constitute the stoma proper. Thee epidermal cells i.e. pore between them
often differ morphologically from the rest adjoining the guard cells
such cells are called subsiaiary of the typical epidermal cells,
or..accessory cells.
stoma together with the subsidiary cells 1s oftenThe whole structure i.e.
In most dicots, the guard termed stomatal complex.
attached to each other ceils are semilunar, crescent-shaped
Dy tne curved, ends of their concavecells and are
Slit-like opening. nmany nonocots sides leaving
the guard cells are dumbbell-shaped
having a thick wall ad Darrow pore 1g.
Aistinct single nucieus, uneoUs 3.7, A). The guard cells contain
toChondria,
reticulum, Golgi bodics, VCuo5 0 various sizes, elements of endoplasmic
grains occur in plastids,. Just beneath each
grains chloroplastids
stoma, a prominentetc.-starch
nace known as sub-stomatal cavity or
chamber is
communicates to the exterior by stomatal pore,
present-this chamber
Stomata occur abundantly on
In leaves; in
to land plants,
to the lower surface of leaves. they
many plants stomata areconined
leaves off are
h the surfaces. n
ioating aquatic plants stomata are
upper TOelow the levelSometimes confined present
of outer stomata are sunken (Fig. 56,
to the
epidermal cells e.g. A-B) 1.e.
sp.), sp. (Apocynaceae), Dasylirion serratifolium gymnosperms (Cycas
(Liliaceae) etc.
 TISSUB SYSTEM 261
In many xerophytes also, some of the epidermal cells with accompanying
stomata lie deep in the concavity forming a socket-like structure, so that the
stomata are sunken i.e. within the epidermal depression which prevents too
much loss of water by transpiration. Stomata raised above the epidermal
layer are found in the fower-peduncle and leaf-petiole of Cucurbita sp.mch
stomata are called raised stomata (Fig. 5'6, C).

STOMATA
OUTER
CHAMBER

INNER
CHAMBER

STOMATAL PIT B MULTIPLE

EPIDERMIS

B-Sunkena
Fig. 5.6-Types of stomata in sectional view. A-Sunken stooma of Hakea sp.
stomata of Nerium sp. C-Raised stoma in the peduncle of Cucurbita sp.

leaves
Arrangement of stomata is of two types. In monocotyledonous
having parallel venation, stomata are arranged in parallel rows while in
thickness of walls of guard cells
dicotyledons they are irregularly scattered. The and thin on the sides
varies. Usually the walls are thick towards the aperture
extensible. The guard
away from the aperture-the thinner parts are more continues on that wall
cells are usually covered by cuticle which generally
also reaches the cells abutting on
which faces the stomatal pore and which

SUBSIDIARY
CELLS
k
GUARD CELLS

C
B in dicots. A-Anomo-
the leaf of Fig 5.8-Different stomatal types
type. C-Paracytic type
Fig.5.7-Stomata from cytic type. B-Anisocytic
and sec
(A) dumbbell- D-Diacytic type. 1950)
Zea mays in surface
tional views (B) showing (After Matcalfe and Chalk,
shaped guardcels.
262 STUDIES IN BOTANY

pteridophytes, 8ymnos
- s t o m a t a l chamber, The guard cells of many are related
lignified wall thickenings that
t hd
tOtThe
some angiosperms possess .
7, Boulter 1970).
opening mechanism of the stomata (Kaufman and in
cell walls of guard cells
their size varies the
ne thickness of the
Gramineae and Cyperaceae),
In grass type (noted in regions with
guard cellepes,
r a cells are dumbbell-shaped (Fig. 5.7) having narrow middlewith subsidiary
cells are associated
; the guard
a n d inner walls found in both
dicotyledons
cells, one on In Amaryllis type,
each long side. opening are
monocotyledons, the cell walls of guard cells around each ventral
found in bryophytes
ckened. In another i.e. Mnium type,
type,
wall of guard cells is thinner than dorsal wall.
Metcalfe
the of the subsidiary cell neighbouring the guard cells,
n e basishave of classified four stomatal types among dicotyledons (Fi8. S.8),
arrangement VizZi
and Chalk
(1950)
or irregular-celled type-Here
actual subsidiary cells are absent;
peA., Anomocytic number of cells which do not differ
in size and
Cn stoma is irregularly surrounded by a certainalso known as RanunculuS type as
seen
1n
Snape Irom other epidermal cells. This type is type is found in speciesS
also
embers of Ranunculaceae (Ranunculus, sp., Clematistosp.)-this
Families Malvaceae, Scrophularia
in
or Cucurbit a, Vitis, and many other plants belonging
ceae, Papaveraceae, Geraniaceae etcC.
Type B Anisocytic or unequal-celled 1ype-Here each, stoma is surrounded by three
two. This type is
subsidiary cells, one cell being considerably smaller or larger than the other
seen in members of Cruciferae (Brassica sp., lberis sp.) and thus known as Cruciferous type.
This type is also found in species of Nicotiana, Solanum, Petunia etc, of the Family Solanaceae,
and in species of Sedum (Crassulaceae).
Type C. Paracytic or parallel-
celled type-Here each stoma is accom
panied on either side by one or more
(often two) subsidiary cells, the longitu-
dinal axes of which are parallel to that of
the guard cells and pore. This type is
also known as Rubiaceous type as found
in members of Rubiaceae (species of
Gardenia, lxora etc). This type is also
seen in species of Phaseolus, Arachis,
Vigna etc. of the Family Leguminosae;
in species of Magnolia. Michelia etc.
of the Family
Magnoliaceae.
A B Type D. Diacytic or cros-celled
type-In this type each stoma is
surroun
dedby two subsidiary cells and their
common wall is at right angles to the
longitudinal axis of the guard cells. This
type is also called Caryophyllaceous
type as it is seen among the
the Family Caryophyllaceae members of
sp.). This type is also noted in(Diant hus
Hygro
phila sp. of Acanthaceae.
Stebbins and Khush (1961) have
classiied the stomatal complex of mono
cotyledonous
(Fig. 5.9): leaves into following types
Type1. In this
type the guard
cells are surrounded by 4-6 subsidiary
cells. This type is found in members
D Araceae, Musaceae, Cannaceae. ot
Jc beraceae and Commelinaceae
Commelina, Rhoeo etc.).
Zingl
(species or
Bia. 5.9-Different stomatal types in monocots. Type 2 Here the guard cells are
2 surrounded by 4-6 subsidiary cells of
A-Type 1 (Commelina Vpe).B-Type type) which are smaller and rounded
2
(Pandanus type). . p e Alisma SteAADe). the
rest and are situated than
the ends or
D-Type 4
(ristype).After
and KhAsh, 1961. guard cells. This type isatfound in
species of Palmae, Pandanaceae manY
danus sp.) and (Pat
Cyclanthaceae.
 TISSUB SYSTEM 263
Type 3. In this type guard cells are surrounded laterally by 2 subsidiary cells, one on
each side. This type is noted in many species of Alismaceae (Alisma sp.), Cyperaceae (Cyperus
sp.), Gramineae (Oryza sp.), Pontederiaceae, Juncaceae eto.
Type 4. Inthis type guard cells are not surrounded by any subsidiary cells. This type is
seen in many species of Orchidaceae (Vanda sp.), Amaryllidaceae, Iridaceac (Iris sp.), Dioscorea-
ceac etc.
Pant (1965), on the basis of ontogeny, has classificd stomata into the following types
()Mesogenous-In this type the subsidiary cells have a common origin with the guard
cells i.e. both are developing from the same meristemoid.
(b) Perigenous-Here the subsidiary cells do not have a common origin with the guard
cells, instead subsidiary cells are formed by cells Iying around the meristemoid that divides to
form the guard cells.
(c)Mesoperigenous-In this type, at least one of the subsidiary cells has a common
origin with the guard cells, but othes do not have common origin.
The opening and closing of stomata are mainly regulated by the changes in the turgor
pressure of the guard cells. Generally an increase in turgor of the guard cells results in the
widening of the stomatal aperture. As a result of increase in turgor, the thinner walls are
stretched more than thicker walls of the guard cell. In grass type, opening is regulated by the
swelling of the ends of guard cells (for detail refer Studies in Botany, volume 2, article 6.4
chapter 6, Plant Physiology section).
Origin: The stomata originate from the protoderm. Protodermal cell divides unequally
into two cells-of these two cells, the smaller one acts as themother cell or the precursor of the
guard cells. The mother cell divides into two cells-these resultant cells take the characteristic
shape of the guard cells through differential expansion. Due to the accumulation and swelling
of intercellular substance between the guard cells, the connection between the cells becomes weak
and thus they (i.e. guard cells) split in their median parts resulting in the formation of stomatal
opening or pore. Development of subsidiary cellstakes place from protoderm cells lying close
to stoma mother cell. In gymnosperms subsidiary cells may not be related to guard cells in
their origin.
Functions of Stomata : Stomata are very important organs from the
physiological point of view (i) To take part in gaseous exchange between
plant organs and atmosphere during respiration and photosynthesis. (i) To
eliminate excess of water absorbed by root system by the process transpi-
ration. (ii) To carry out photosynthesis in the guard cells owing to the
presence of chloroplastids.
2. Water stomata-Refer article 4.7, 3 (iv), page 249
B. EPIDERMAL OUTGROWTHS-Epidermal cells give rise to various types
of multicellular and unicellular appendages or outgrowths differing in shape,
size and function-all these are designated by the term trichome. Trichomes
include glandular or nonglandular hairs, scales, papillae, absorbing root hairs
etc. Other structures such as warts, spines, some secretory structures etc.
which consist of epidermal as well as subepidermal tissues are called
emergences,
1. Hairs-Hairs or trichomes may be unicellular or multicellular,
glandular or nonglandular, branched or simple i.e. unbranched. Trichomes
are formed on all parts of the plants including stamens e.g. Tradescantia spP:
and seeds e.g. Gossyptum sp. In Gossypium sp. the fibres used in commercial
purpose constitute epidermal hairs which are located on the seed coat.
Stem hairs are multicellular, branched or unbranched and either glandular or
nonglandular. Multicellular hairs may be composed of a single row ofcells
(Fig. 5.10, D) or of several rows of cells. Multicellular hairs are usually
provided with two parts viz., (a) basal part embedded in epidermis known as
branched
foot and (b) projected part known as body. The shape of
multicellular hairs are diverse, sometimes they look like star-shaped structure
be tree-like in structure
known as stellate hair (e.g. Styrax sp.) or they mayPlatanus and Verbascum).
of
(Fig. 5.10, G-H), called dendroid (e.g. species
unbranched and elongated forming
Unicellular hairs are often simple,