Unit : 2

(a) Shoot apical meristem (SAM)

(b) Root apical meristem (RAM)

LESSON STRUCTURE
2.0 Objective
2.1 Introduction
2.2 Shoot Apical Meristem (SAM)
2.3 Origin and Evolution of Meristematic Tissue Organisation
2.4 Root Apical Meristem (RAM)
2.5 Root Meristematic Tissue Organisation And Example of
(RAM)
2.6 Questions for Exercise
2.7 Suggested Readings

2.0 OBJECTIVE

Initiation of permanent tissues from meristematic tissue is a very common but important
natural phenomenon by which a plant elongates and obtains its optimum length and width
according to their genetic set up.

2.1 INTRODUCTION

Meristem - divided (divisible) Meristematic tissue are made up of those cells which
have the capacity of division or they retain the dividing power. These cells are either spherical
oval or polygonal in shape without inter cellular spaces. Thin walled homogeneous active
and abundant protoplasm with large nuclei and vacuole almost very small or absent.
According to Eames and MacDaniels (1947) meristem is also known as meristematic
tissue meaning divisible. Meristem can be defined as the localized region where the cells

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 (a) Shoot apical meristem (SAM) (b) Root apical meristem (RAM)

are more or less orderly arranged and divide to produce new cells which undergo
differentiation to form mature tissues.
Though Nageli (1844) coined the term meristem to designate the dividing cells, but in
botanical literature, this term is applied to the undifferentiated, partly differentiated and mature
cells, which has regained the power of cell division. The transformation of zygote into embryo
and its subsequent growth and development leads to the formation of multicellural mature
plant. These events occur because of multiple mitosis. In mature plants, the mitotically active
centres are localized and are called meristem.
Meristem occurs in the root tip, stem tip, in the axil of leaf at the bases of internodes, in
the flower buds, in the vascular bundle as fascicular cambium, at the peripheral side of cortex
as cork cambium (Phellogen) and between the vascular bundles as inter fascicular cambium
at the time of secondary growth.
The meristematic tissues give rise to permanent tissues. The permanent tissue system
of plants comprise (1) The dermal tissue system, which makes up epidermis and periderm or
protective tissues (2) The vascular tissue system, which includes xylem and phloem or
conducting tissue and (3) The ground tissue system, which consists of all tissues except
dermal and vascular tissues.

Characteristic features
(1) Shape Isodiametric
(2) Intercellular space more or less absent.
(3) Thin walled
(4) Protoplasm contains food, proplastids, crystals, large nucleus, vacuoles and plastids.
The region of undifferentiated meristem constitutes promeristem and these regions are
the growth centres where the formation of new tissues are initiated. These initiating regions
are sometimes referred to as embryonic meristem or primordial meristem. Promeristem is
situated at the apical most part of the organ with undifferentiated cells. As soon as differntiation
starts, they are no longer a part of promeristem.

Meristems are classified on the basis of certain factors

(a) According to their origin and development.
(b) According to their position.
(c) According to their function.
(A) According to their origin and development
(a) Promeristem or primordial meristem.
(b) Primary meristem.
(c) Secondary meristem.

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 (a) Shoot apical meristem (SAM) (b) Root apical meristem (RAM)

(B) According to their position in the plant body

(a) Apical
(b) Intercalary
(c) Lateral
(C) According to their function
(a) Proto dermal
(b) Procambium
(c) Ground or fundamental meristem.

(A-a) Promeristem
Promeristem consists of a groups of meristematic cells representing the earliest or
youngest stage of a growing organ and from this stage the differentiation of later meristem
and finally of permanent tissues takes place. It occupies a small area at the tip of the stem
and the root. The promeristem by cell divisions gives rise to the primary meristem.

(A-b) Primary meristem

Primary meristem is derived from the promeristem and still fully retains its meristematic
activity. Its cell divide rapidly and became differentiated into distinct tissues. The primary
permanent tissues which make up the fundamental structure of plant body mainly growing
apical region of the root and stem, cambium of the stem give rise to primary meristem as well
as secondary permanent tissues. The cambial cells divide mainly in single tangential plane
while those of the primary meristem divide into three or more planes.

(A-c)Secondary meristem
Secondary meristem appears later at a certain stage of development of an organ of a
plant. It is always lateral lying along the side of the stem and the root. It is seen that some of
the primary permanent tissues become meristematic and they aquire the power of division
then constitute the secondary meristem. The cambium of the root, the interfascicular cambium
of the stem as well as cork-cambium are the best examples. All lateral meristems give rise to
the secondary permanent tissues which are responsible for growth in thickness of the plant
body.

B. According to the position in the plant body

(B-a) Apical meristem
On the basis of position the apical meristem lies at the apex of the stem and the root
representing their growing regions and is of varying lengths. Usually ranging from a few
millimetres to a few centimetres. It includes the promeristem and the primary meristem which
give rise to the primary permanent tissues and is responsible for growth in length of the plant
body. It is observed that the promeristem consists of a group of meristematic cells in higher
plants where as in lower plant or in pteridophytes it is represented by a single cell.

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 (a) Shoot apical meristem (SAM) (b) Root apical meristem (RAM)

(B-b) Intercalary meristem

Intercalary lies between masses of permanent tissues either at the base of the leaf or at
the base of the internode or some times below the node. It is a detached portion of the apical
meristem separated from the later due to growth of the organ. Intercalary meristem also gives
rise to the primary permanent tissues. It is generally short lived disappearing soon or becoming
transformed into permanent tissues.

(B-c) Lateral meristem

Cambium of the stem lies laterally in strips of elongated cells extending from the apical
meristem as in the stem of dicotyledons and gymnosperms it divides mainly in the tangential
direction, give rise to the secondary permanent tissues to the inside and outside of it and is
responsible for growth in thickness of the plant body.

(C) According to their function

(C-a) Protoderm
It is observed by Nageli (1858) that the apical meristem consists of a single apical cell
in all plants and supported by Hofmiester what ever he observed that after a sequence of cell
division it is responsible for the formation of different membranes of plant body, protodermal
meristems are often common is lower plants which gives rise to the epidermal tissue system.

(C-b) Procambium
Before the intiation of cambium meristematic cells are known as procambium. Activation
of procamibium differentiates the growing layers of plant body. Procambium can be absorbed
in almost all the vascular plants which gives rise to the vascular tissue system.

(C-c) Ground of fundamental meristem

Fundamental or ground meristem gives rise to the ground tissue system and responsible
for the thickness of the plant body. Haberlandt (1914) explained it on the basis of physiological
activities.

2.2 SHOOT APICAL MERISTEM (SAM)

A meridian T.S. and L.S. through the apex of a stem, when examined under the
microscope, shows that the apical meristem or growing region is composed of a small mass
of usually rounded or polygonal cells which are all essentially alike and are in a state of division,
these meristematic cells constitute the promeristem. The cells of the promeristem soon
differentiate into three regions dermatogens, periblem and plerome. The cells of these three
regions grow and give rise to primary permanent tissues in the mature portion of the stem.
The section further shows on either side a number of out growth which arch over the growing
apex; these are the young leaves of the bud, which cover and protect the tender growing apex
of the stem. Theories regarding apical meristem were formulated by Nageli (1858), Hanstein
(1870) and Schmidt (1924) are known as:

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 (a) Shoot apical meristem (SAM) (b) Root apical meristem (RAM)

(a) Apical cell theory

(b) Histogen theory
(c) Tunica corpus theory
(a) Apical cell theory
Nageli in (1844-1859) proposed this theory. He first coined the term meristem and said
that the apical meristem consists of a single apical cell in plants, and that the sequence of cell
divisions is responsible for the formation of different members of the plant body. Nageli
believed that a single apical cell is present in the root and shoot apices of cryptogams. This
apical initial may be lens or wedge shaped. Prismatic or tetrahedral. One side of the tetrahedral
cells faces towards the apex and the other three sides face inner and peripheral sides. Nageli
assumed that all the tissues of root and shoot are derived from this single apical cell, which is
the basis of apical cell theory and considered as the structural and functional unit of apical
meristem. Nageli’s single cell theory is no doubt true of the thallophytes and vascular
cryptogams but his assumption that this is applicable to all cryptogams and phanerogams
has proved to be wrong. Though this single theory was supported by Hofmiester but he
expressed doubts regarding its applicability to all cases, particularly phanerogams. After
extensive investigation it is found that solitary apical cell theory is not universally supported
and it was soon superseded by other theories.
(b) Histogen theory
Hanstein (1868-70) put forward this theory. The shoot and root apices consist of a group
of homogeneous meristematic cells, which are promeristem. This theory postulates the
existence of three cell initiating regions or histogens or tissue builder in the promeristem.
According to Hanstein, the three regions originate from independent group or separate set of
intials. These regions are histogen and termed as dermatogen.

PERIBLEM AND PLEROME

Dermatogen (meaning skin)
The outermost histogen of promeristem is
designated as dermatogen. It forms a mandle like layer
surrounding the periblem. It is single layered epidermis
of root and shoot develops from this meristem.

Periblem (meaning clothing)

The intermediate zone (histogen) internal to
dermatogen of the apical meristem is the Periblem. It
remains covering the plerome. At the topmost apices
of apical meristem, it is single layered and gradually it
becomes multilayered and this histogen gives rise to
cortex of root and shoot. The inner tissues of the leaf Fig. 1 Stem apex in median L.S.,
at the shoot apex are derived from periblem. Longitudinal Section.

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 (a) Shoot apical meristem (SAM) (b) Root apical meristem (RAM)

Plerome (Greek word means-fills)

The innermost histogen which is
situated at the central region of the apical
meristem in the plerome. It remains
surrounded by periblem. The cells of the
histogen are thin walled and isodimetric.
Plerome gives rise to entire vascular cylinder
including pith. The pericycle and medullary
rays also develop from this meristem.
Precambian is formed from plerome at later
stages, procambium gives rise to primary
vascular bundles.
Homstein’s histogen theory dominated
for a long time to interpret the origin of tissues
of an organ. On the basis, Haberlandt (1914)
designated the histogens as protoderm.
Procambium and ground meristem though
these terms are quite distinct and not
Fig. 2 : Apical Meristem Showing V.S.
synonymous to Hanstein’s dermatogen,
periblem and plerome. Now a days this
theory is of little use due to (1) there exists
no distinction between histogens in an
apex (ii) The distinction between periblem
and plerome is not prominent in many
gymnosperms and angiosperms. (iii) The
derivatives of histogen is not constant. In
some plants plerome forms pith only in
other it gives rise to part of cortex and in
still other the entire vascular cylinder. On
the other hand, the periblem gives rise to
cortex and peripheral layers of stellar
tissue or part of the cortex.

(c) Tunica-Corpus Theory

Tunica-corpus theory was proposed
by Schmidt (1924). In contrast to apical
cell theory and Histogen theory, this theory
is applicable only to the apical meristem Fig. 3 : Apical Meristem (Redrawn after Meyar et al)
of shoot and not to the root. This theory Pr.d.-Protoderm, G.M.-Ground Meristem, Pr.C.-Procambium,
E-Epidermis, C-cortex, Pr.ph.-Primary phloem, Pr.X-Primary
recognized two regions or zones the Xylem, Camel-Cambium Ring.Sc.Ph.-Secondary phloem,
tunica and the corpus in the shoot apex. Sc.X-Secondary Xylem.

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 (a) Shoot apical meristem (SAM) (b) Root apical meristem (RAM)

Tunica : It is the surface layer of shoot apical meristem. It consists of one or more
peripheral layers of cells. Which are not always constant and usually range from one to nine.
One single layer, called as monostrotose is probably most common, followed by two to four,
known as multistrotose. The number of layers may even be more or varies in families, genera
and species at different stages of development. The cells to tunica are compactly set and
regularly oblong in longitudinal section. The cell division in this layer is predominantly anticlinal,
it means walls are laid perpendicular to surface especially at the point of origin of leaf and
exillary bud. Therefore, these layers have only surface growth and thus shoot apex grows in
surface area.
There are separate initials for each tunica layer, the number of initials corresponds to
the number of layer of tunica present in the shoot apex.
There are two zones can be distinguished in tunica:-
(a) Peripheral zone
(b) Central zone
Peripheral zone surrounds Central zone where as Central zone consists of a single or
few initials. The cells of the central zone are larger in size and possess larger nuclei and
vacuoles than the cells of peripheral zone. The peripheral zone lies between the control apical
zone leaf primordia. In the peripheral zone sometimes periclinal divisions are observed in
addition to anticlinal divisions which is the chief mode of division of tunica. Periclinal division
also occurs in tunica of some monocotyledons. In general the outermost layer of tunica
differentiates into epidermis. The other zones like cortex and stele may also arise from tunica
and it varies in different plants.
CORPUS : This meristematic zone lies beneath the tunica. The cells are larger than the
cells of tunica. The cells of corpus divide in all direction and form the central core of the shoot
apex, generally corpus originates from a single tier of initial cell situated below the tunica.
This initial layer becomes several layers by periclinal division and then divides in all planes to
increase the volume of the shoot apex. In most cases corpus gives rise to cortex and stele.
It is the position and plane of cell division that differentiates tunica and corpus, the tunica
is more homogeneous. Recent electron microscope study reveals that the ultrastructural
differences between these two layers are mainly quantitative.
The tunica-corpus concept is applicable to angiosperm shoot apex only. The two-zone
tunica and corpus cannot be distinguished in cryptogames and Gymnosperms except Gnetum
which shows tunica-corpus pattern of growth in the shoot apices. The number of layers, plane
of cell division and the destiny of tunica and corpus is not constant and varies from families,
genera and species and even at different stages of development. There is no precise definition
of tunica and corpus. This theory is purely descriptive and served well to describe the shoot
apex of angiosperm.

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 (a) Shoot apical meristem (SAM) (b) Root apical meristem (RAM)

2.3 ORIGIN AND EVOLUATION OF MERISTEMATIC TISSU

ORGANISATION
Since the origin of the cellular organisation there was a tendency of each and every
individual to prove it self to be the fittest one in the existing environmental conditions. As
different authers differ widely in their lines of thinking with regard to the origin and evolution of
meristematic tissue. It is felt necessary to review those possible lines instead of expressing
one’s personal opinion in this aspect of meristematic tissue. This will give every one a wide
scope for independent thinking after giving due consideration of every body’s line of thought.
This discussion is primarily based on the solitary or single layered meristematic cells upto
differentiation zones.
The difficulties associated with the interpretation are mainly wide variation in the
differentiation zone due to the biochemical and metabolic activities. As well as variable
pigmentation in meristematic zones of plant body.
It is too difficult to say that when and how the meristematic initial tissue was originated
but some principles, theories and hypothesis are leading to show the possible path of this
natural evolution in the absence of the evidences, only on the basis of same assumptions and
postulates of some renouned thinkers philosophers as well as the fossil history and foot prints
which are shown below for disclosing this mystery.
There are two lines of evolution which are recognised and supported by various scientists
(a) Gametophytic generation : (Haploid cells and tissues as well as Haploid thallus
reorganisation) usually the apical initial meristem single cell was observed there)
Prominent in cryptogamic flora.
(b) Sporophytic generation : (Diploid cells and tissues) where the apical initial
meristematic cells were observed.
Prominant in phanerogames.
Similarly there are two contradictory views of evolution of stem apical meristem.:
(1) Up grade or Progressive evolution theories:
These supporters are:
Nageli (1858-60)
Hanstein (1870)
Lotsy (1909), Campbell (1940),
Smith (1955), Calakovsky (1874)
Feldmann (1952)
(2) Down-grade or Regressive evolution theory: supporters are - Haberlandt (1914)
Takhtazan and Zimmermann (1966), Mehra (1953), Evans (1939), Wettstein (1903-
1908), Harris (1938), Church (1919), Bold (1938) and Fritsch (1945).
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 (a) Shoot apical meristem (SAM) (b) Root apical meristem (RAM)

It is assumed that the evolution started from, volvoxlike colony and forwarded like.
Volvox - Coleochaete - Chara - Nitella - Laminaria - Fucus - Polysiphonia - Peziza -
Agaricus - Sphagnum - Funaria - Lycopodium - Selaginella - Marsilea - Pteris - Cycas - Pinus
- Gnetum and then Angiosperms growing regions.
Successive cuts of cells in apical pyramidal cell, it supports apical cell theory of (SAM)
development of stele in pteridophytes, development of antheridium and archegonium in
gymnosperms and finally in every growing stem regions of angiosperms are the examples of
the (SAM). Now some diagrammatic representation of cyto-histological zonation in the
promeristem of vegetative shoot apices,
1. Pteridophyte type
2. Selaginella type
3. Cycas type
4. Ginkgo type
5. Cryptomeria - Abies type.
6. Opuntia
7. Usual Angiosperm type
(Redrawn of Fahn 1997)

Internode

Fig. 4 (Redrawn after Lyndon, 1988)

A.M.-Apical Meristem, L.M.-Lateral Meristem,
L.S.-Longitudinal Section, T.S.-Transverse Section, Fig. 5 (Redrawn after Lyndon, 1990)
I.M.-Intercalary Meristem, S-Seath, A.M.-Axillary Meristem. A Typical plant showing Meristematic zones.

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 (a) Shoot apical meristem (SAM) (b) Root apical meristem (RAM)

Buvat (1955) advocated the idea of existence of quiescent centure in the shoot apex.
Buvat (1955) divided the shoot apical meristem into
(a) Waiting meristem.
(b) Initiating ring.
(a) Waiting meristem is apical in position and it rarely divides during vegetative
development, they become active when the vegetative apex become reproductive at the time
of formation of terminal flower or inflorescence. It is postulated that all reproductive growth is
due to the activity of waiting meristem. Buvat (1955) recognised four meristematic zones in
the vegetative shoot apex.
(1) Initial
(2) Sporogenous promeristem
(3) Receptacular promeristem
(4) Pith meristem

Fig. 6 (Digram showing C.M.C. and arrows showing the direction of cell displacement from the cmc.
Redraw after Newman (1965).
A Typical plant showing Meristematic zones.

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 (a) Shoot apical meristem (SAM) (b) Root apical meristem (RAM)

Initial and sporogenous correspond to tunica layer and remaining two represent the
corpus. Another interpretation was put forward by Newman (1965) regarding apical meristem.
Where no cell is a permanent cell. According to this view the apical meristem is continuing
meristematic residue (c.m.r.) on the basis of (c.m.r.) Newman (1965) proposed three basic
types of shootapex in vascular plants.
(1) Monoplex apex.
(2) Simplex apex.
(3) Duplex apex.
(1) Monoplex apex: It is found in mostly Algae and Fern. Continuing meristematic
residue may consists of a single cell, two cells with a common wall or three or more cells
which have a common angle. During division cell wall is formed parallel to inclined wal. This
type of division contributes to increase in both length and breadth. This also provides for
increase in both length and breadth as well as increase in bulk growth.
(2) Simplex apex: In gymnosperm (c.m.r.) lie on the superficial layer only. It may consist
of a single cell more than one cell that have a common wall and three or more cells which have
a common angle. The cells of c.m.r. are parallel with each other.
Division of walls occur in two ways:
(a) Perpendicular to outer face.
(b) Parallel with outer wall.
or
(a) Anticlinal divi-sion
(b) Periclinal divi-sion
Anticlinal division causes the increase in breadth, where as periclinal division contributes
in the increase of length. Peri and anticlinal-these two divisions are necessary for bulk growth.
These division are restricted to a single layer and is named as simplex.
(3) Duplex apex: In Angiospermic flora the cells of (c.m.r.) appear to be paralled with
each other, at least in two successive layers, Peripheral being the surface layer and the
innermost layer behaves as simpex apex. Periclinal and anticlinal divisions occur in the
innermost layer only. Which causes the increase in length and breadth and thus the bulk
growth. The place where the anticlinal division occurs is responsible for the increase in surface
or breadth only, therefore duplex apex, shows two modes of growth.
(i) Outer anticlinal division causing surface growth.
(ii) Inner cells have both anticlinal and periclinal divisions. As a result, increase in
length and breadth occurs. Thus causing bulk growth.
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 (a) Shoot apical meristem (SAM) (b) Root apical meristem (RAM)

Difference between root apical meristem and shoot apical meristem

ROOT APICAL MERISTEM SHOOT APICAL MERISTEM
1. It is less complicated than the shoot 1. It is more complicated than the root apical
apical meristem. It produces lateral roots meristem. It has lateral organs like leaves
only. It functions in a constant manner. or floral organs and cyclic. The position
of new organs is predicatable of those
already present.
2. It is sub-terminal in position. It remains 2. It is terminal in position. It remains over-
covered by vaculated tissues of root cap. arched by leaf primordia.
3. Due to the absence of lateral primordia, 3. There is a regular rhythmic change due
there is no rhythmic change in size and to presence of leaf primordia.
form.
4. Phenomenon of ageing is prominent in 4. Phenomenon of ageing is combated
excised roots. when grown in light.

2.4 ROOT APICAL MERISTEM (RAM)

A median longitudinal section through the apex of the root shows that it is covered over
and protected by a many layered tissue
which constitutes the root cap. The apical
meristem or growing region lies behind the
root cap. The promeristem as in the stem,
early differentiates into three regions.
(1) Dermatogen
(2) Periblem
(3) Plerome
At the embryonic stage the root apical
meristem is organized opposite to the shoot
apex. In contrast to shoot, It is simpler in
gross structure but is complicated at apex
due to root cap, which develops from the
root apical meristem, thus the meristematic
region remains well protected. The root
apex may consist of a single cell or a group
of cells. The apical cell theory of Nageli in
1978 and the Histogen Theory of Hanstein
(1868) is applicable to root apex but Tunica Fig. 7 Diagram showing cyto-histological zone.
corpus theory is not applicable. (Redrawn after Fahn (1997))

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 (a) Shoot apical meristem (SAM) (b) Root apical meristem (RAM)

(1) Dermatogen
As in the stem, this is also single layered, but at the apex it merges into the periblem;
many new cells cuts off just out side the dermatogen and forms calyptrogen means cap
producing small celled tissue. These small celled tissues are responsible for the formation
and replanishing or renewing the root caps under the hard soil. In some plants the dermatogen
directly gives rise to the root cap without the intervention of the calyptrogen. The walls of outer
cells of the root cap may be modified into mucilage which helps the root to push forward in the
soil more easily. Sometimes in dicotyledons generally the dermatogen is exhausted in the
formation of the root cap so that the outermost layer of the root is derived from the outermost
layer of the periblem.

(2) Periblem
As in the stem, this is also single-
layered at the apex and many layered
higher up. In monocotyledons generally the
outermost layer of the periblem forms the
outermost layer of the root forms the middle
region or cortex of the root.

(3) Plerome
The plerome’s structure and function
are practically the same as those of the
stem. But here some procambial strands
give rise to bundles of vessels and others
to bundles of sievetubes or phloem in an Fig. 8 Diagram Showing
alternating manner. Root-apex in median longitudinal section.

2.5 ROOT MERISTEMATIC TISSUE ORGANISATION AND EXAMPLEX OF (RAM)

At the embryonic stage the root apical meristem is organised opposite to the shoot
apex. In contrast to shoot, it is simpler in gross structure but is complicated at apex due to the
presence of root cap, which develops from the root apical meristem.
Now same examples of root meristematic tissue organisation.

(1) Root of Pteridophyta

The apical cell theory of Nageli holds for the root apex of vascular cryptograms. The root
apex of Equisetum, Dryopteris, Ophioglossum and other members of pteriodophytes possess
a single tetrahedral apical cell in their root apices. This single initial cell has four cutting faces,
which on repeated division contributes to growth in root cap, epidermis cortex and stele. The
upper three sides of the initiating cell forms epidermis, cortex and stele and the root cap
develops from the lower side.

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 (a) Shoot apical meristem (SAM) (b) Root apical meristem (RAM)

All cells of the root can be traced back to a single apical initial. The immediate derivatives
of this apical initial are also mitotically active. Therefore, the root apex appears to constitute
a multi cellular meristem. The idea of an inactive root apical meristem or quiescent was
applied to the root tips of many vascular cryptogams. However recent investigation reveals
that entire root tip is mitotically active

(2) Root apex of Gymnosperm

Allen (1974) described the root apex in Pseudotsuga. It consists of four zones one
permanent initials and three groups of temporary initials. The first group of temporary initial
gives rise to meristem of vascular cylinder that form stele. The cortex is formed from the
second group of temporary initials, which form the meristem of cortex the peripheral layer of
cortex gives rise to protoderm from where epidermis develops. The third group of temporary
initial is the collumella mother cell and it produces columella. Wilcox (1954) reported the
presence of two groups of temporary initials in Abies procera. One group of temporary initial
gives rise to the vascular cylinder, and the columella develops from the other group. The
columella by cell division contributes to growth in both root cap and cortex.
Schematic diagram showing the zones of a root apical meristem.
(Redrawn after Mauseth 1988)

(3) Root apex of Angiosperm

Gutten Berg (1960) distinguish three groups of temporary initials and a central group of
permanent initials in the root apex of angiosperm. The derivatives of temporary initials may
vary from plant to plant, as for example in Brassica, the meristem of vascular cylinder develops
from one group of temporary initial. Other group give rise to meristem of cortex, protoderm
and root cap originates from the last group of histogen or temporary initials. Protoderm gives
rise to epidermis. The root cap and epidermis have common histogen and this special
temporary initial is called dermatocalyptrogen (Gutten Berg, 1960). Dermotocalyptrogen is
also observed in other members of Solanaceae, Rosaceae and Asteraceae etc. However in
Zea mays, protoderm and cortex originate from a common temporary initial. The vascular
cylinder develops and cortex originate from a common temporary initial. The vascular cylinder
develops from another temporary initial. The root cap originates from a separate histogen or
temporary initial. This special temporary initial or meristem of root cap was termed as
calyptrogen by Janczewski in 1874, some member of Arecaceae and Zingiberaceae also
possess calyptrogen.
Eames and Mac Daniels (1947) described the root apex of Cryptogams, Gymnosperms
and Angiosperms. Diagram demonstrating the type of root apex in (A) Cryptogams
(B) Gymnosperm (C) Dicotyledon (D) Monocolyledon based on concept of Eames and
Macdaniels (1947).

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 (a) Shoot apical meristem (SAM) (b) Root apical meristem (RAM)

Q.C. - Quiescent Centre

Theories of Root Apical Meristem

1. Apical cell theory by Negeli (1844)
2. Histogen Theory by Hanstein (1865)
3. Korper - Kappe theory by Schuepp (1917)
Apical cell theory by Nageli (1844) put forward this theory on the root apex of cryptogams
where as Histogen theory of Hanstein (1865) proposed this theory to prove and show the
three distinct cell initials or histogen in the apical meristem of root of angio-sperms. Recent
investigations discarded
the histogen theory still
several authors
described the root apex
based on histogen
concept. Korperkappe
thory Schuepp (1917) put
forward this theory
Korper and Kappe
means body and cap
respectively. The body-
cap concept is based
solely on planes of cell
division and deals with
different aspects of
apical activity.
Newman (1965)
proposed five types of
root apex based on the
types described by Fig. 9 : Diagrams showing types apical meristem. Arrows showing
Eames and Macdaniels direction of cell displacement from CMC.
(1947) and Esau (1953) Redrawn after Newman 1965
V.C. - Vascular Cylinder, E-Epidermis
the amalgamated the O.C. - Outer Cortex, I.C. - Inner Cortex
types as follows. R. C. - Root cap

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 (a) Shoot apical meristem (SAM) (b) Root apical meristem (RAM)

(Redrawn after Newman (1965)

Diagrams demonstrating types of root apical meristem in longitudinal section arrows
show the direction of cell displacement from the continuing meristematic residue.

Fig. 10 : Diagram showing Root apex.

Based on Concept of Eames and MacDaniels (1947)

2.6 QUESTIONS FOR EXERCISE

1. Give an account of organisation of root apical meristem.

2. Give a detailed account of shoot apical meristem.
3. What are the main differences in organisation of root apical meristem and
shoot apical meristem.
4. Write short notes on the following :–
(a) (RAM) (b) (SAM)
(c) Dermatogen (d) Periblem
(e) Plerome (f) Apical cell theory
(g) Histogen theory (h) Protoderm
(i) Procambium (j) Cambium
(k) Fundamental meristem
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 (a) Shoot apical meristem (SAM) (b) Root apical meristem (RAM)

5. Give a detailed account of meristem.

6. Discuss the theories related to the development of (SAM).
7. Discuss the tissue organisation of (RAM).

2.7 SUGGESTED READINGS

1. Collage Botany - Kar

2. Botany for Degree student - A. C. Dutta.
3. Botany - J. Ben. Hill
4. Plant Anatomy - Piyush Ray.
5. Plant Anatomy - E. G. Cutter.
6. Plant Anatomy - K. Esaw.
7. Plant Anatomy - A. Fahn.

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