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apical meristem
Submitted to: Dr. Aisha Nazir
Submitted by: Shagufta Zaman
Roll no: BSBOT04F18
BS Botany 6th semester
University of the Punjab
May 03, 2021
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Table of Contents

Apical meristem...................................................................................................................................3
Types of apical meristems...............................................................................................................4
Delimitations of tissue system.........................................................................................................5
Evolution of the concepts of apical organization...............................................................................6
Shoot apical organization................................................................................................................6
Apical cell theory.............................................................................................................................6
Histogen theory................................................................................................................................7
Tunica corpus theory.......................................................................................................................8
Cytohistological zones of apical meristem.......................................................................................10
Inquires of initials and derivatives...................................................................................................11
Vegetative shoot apices......................................................................................................................13
Root apices.........................................................................................................................................14
Root apical organization...................................................................................................................15
Apical cell theory...........................................................................................................................16
Histogen theory..............................................................................................................................16
Korper kappe theory.....................................................................................................................16
Apical organization in roots..........................................................................................................17
Quiescent region in root apex.......................................................................................................20
Growth of root tip..........................................................................................................................21
Conclusive remarks...........................................................................................................................22
References..........................................................................................................................................23
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Apical meristem

“The term apical meristem refers to a group of meristematic cells at the apex of shoot and
root that by cell division lay the foundation of the primary plant body.’’

Meristems are composed of initials, which perpetuate the meristems, and their
derivatives. Also, the derivatives usually divide and produce one or more generations of cells
before the cytologic changes, denoting differentiation of specific cells and tissues, occur near
the tip of the shoot or root. The divisions continue at all levels where such changes are
already discernible. Therefore growth, in the sense of cell division, is not limited to the very
tip of shoot or root but extends to levels considerably removed from the region usually called
the apical meristem. Most commonly the term apical meristem is used in a wider sense than
merely the initials and their immediate derivatives; it also includes variable lengths of shoot
and root proximal to the apex.

The change from apical meristem to adult primary tissues is gradual and involves the
intergrading of the phenomena of cell division, cell enlargement, and cell differentiation, so
one cannot restrict the term meristem to the apex of shoot and root. The parts of shoot and
root where future tissues and organs are already partly determined but where cell division and
cell enlargement are still in progress are also meristematic.

It also includes variable lengths of shoot and root proximal to the apex. Shoot apex
and root apex often are employed as synonyms of apical meristem, although a distinction is
sometimes made between the shoot apical meristem and the shoot apex: the apical meristem
denotes only the part of the shoot lying distal to the youngest leaf primordium, whereas the
shoot apex includes the apical meristem together with the subapical region bearing young leaf
primordia (Cutter, 1965). When determinations of the dimensions of apices of shoots are
made, only the part above the youngest leaf primordium, or the youngest node, is measured.

There are detailed differences between the meristems at the apex of shoot and root of
monocotyledons, dicotyledons and gymnosperms. Three shoot apices are shown in Fig:
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Types of apical meristems

Apical meristems are divided into two types

1. Shoot apical meristem (SAM)

2. Root pical meristem (RAM)

Shoot apical meristem

These are regions of unlimited growth and development give rise to the leaves and
buds. These shoot apical growing centres form the primary plant body. It is the spot of the
embryogenesis in flowering plants. Leaf’s primordia, sepals, petals, stamens, and ovaries are
originated from here one on every interval of time called a plastochron.

Root apical meristem

In contrast to shoot apical meristem the root apical meristem produces cells in two
directions. Root tissues are originated from root apex. It consists of stems cells and quiescent
centre; both are involved in forming cells of mature roots. Root cap covered the apex of root
meristem mainly to protect and guide growth.
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Fig: Root and shoot apical meristem

Delimitations of tissue system
Apical meristem will be divided into two main regions

 the pro-meristem which comprises the apical initials and neighbouring cells
 the meristematic zone below it in which the three basic meristems (the protoderm,
procambium' and ground meristern) of the tissue systems can be distinguished

In descriptions of primary differentiation in shoot and root tips, the initiating cells and their
most recent derivatives are frequently distinguished from the partly differentiated but still
meristematic subjacent tissues under the name of promeristem (or protomeristem; Jackson,
1953). The subjacent meristematic tissues are classified according to the tissue systems that
are derived from them, namely into:

1. Protoderm, which differentiates into the epidermis.

2. Procambium (also called provascular tissue), which gives rise to the primary
vascular tissues.
3. The ground meristem, precursor of the fundamental or ground tissue system.

If the term meristem is used broadly, protoderm, procambium, and ground meristem are
referred to as the primary meristems (Haberlandt, 1914). In a more restricted sense of
meristem (combination of initials and immediate derivatives), these three tissues constitute
partly determined primary meristematic tissues.
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The terms protoderm, procambium, and ground meristem serve well for describing the
pattern of tissue differentiation in plant organs, and they are correlated with the equally
simple and convenient classification of mature tissues into the three tissue systems,
epidermal, vascular, and fundamental. It seems to be immaterial whether the protoderm,
procambium, and ground meristem are called meristems or meristematic tissues as long as it
is understood that the future development of these tissues is at least partly determined.

Fig: Delimitations of apical meristem

Evolution of the concepts of apical organization

In the nineteenth century research workers mainly dealt with the problem of the number of
the initials in the apices and the determination of the tissues that were derived from them.

Shoot apical organization

1. Apical cell theory
Apical Meristems Originally Were Envisioned as having a Single Initial Cell. Following
Wolff’s (1759) recognition of the shoot apex as an undeveloped region from which growth of
the plant proceeded and the discovery of a single, morphologically distinct initial cell at the
apex of seedless vascular plants, the idea developed that such cells exist in seed plants as
well. The apical cell was interpreted as a consistent structural and functional unit of apical
meristems governing the whole process of growth (the apical-cell theory). Subsequent
researchers refuted the assumption of a universal occurrence of single apical cells and
replaced it by a concept of independent origin of different parts of the plant body.
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2. Histogen theory
The Apical-Cell Theory was Superseded by the Histogen Theory. The histogen theory
was developed by Hanstein (1868, 1870) on the basis of extensive studies of angiosperm
shoot apices and embryos. According to this theory the main body of the plant arises not from
superficial cells but from a massive meristem of considerable depth comprising three parts,
the histogens, which may be distinguished by their origin and course of development. These
include:

 The dermatogen (from the Greek words meaning skin and to bring forth), is the
precursor of the epidermis.
 The second, the periblem (from the Greek, clothing), gives rise to the cortex.
 The third, the plerome (from the Greek, that which fills), constitutes the inner mass of
the axis.

The dermatogen, each layer of the periblem, and the plerome begin with one or several
initials distributed in superposed tiers in the most distal part of the apical meristem.

Hanstein’s “dermatogen” is not equivalent to Haberlandt’s (1914) “protoderm.”

Haberlandt’s protoderm refers to the outermost layer of the apical meristem regardless of
whether this layer arises from independent initials or not, and regardless of whether it gives
rise to the epidermis or to some subepidermal tissue as well. In many apices the epidermis
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does originate from an independent layer in the apical meristem, in such apices the protoderm
and dermatogen may coincide. The periblem and plerome in the sense of Hanstein are
discernible in many roots but are seldom delimited in shoots.

Thus, the subdivision into dermatogen, periblem, and plerome has no universal
application. But the fatal flaw of Hanstein’s histogen theory is its presumption that the
destinies of the different regions of the plant body are determined by the discrete origin of
these regions in the apical meristem.

3. Tunica corpus theory

The apical-cell and histogen theories were developed with reference to both root and shoot
apices. The third theory of apical structure, the tunica-corpus theory of A. Schmidt (1924),
was an outcome of observations on angiosperm shoot apices. It states that the initial region of
the apical meristem consists of:

 The tunica, one or more peripheral layers of cells that divide in planes perpendicular
to the surface of the meristem (anticlinal divisions).
 the corpus, a body of cells several layers deep in which the cells divide in various
planes.

Thus, whereas the corpus adds bulk to the apical meristem by increase in volume, the one
or more layers of tunica maintain their continuity over the enlarging mass by surface growth.
Each layer of tunica arises from a small group of separate initials, and the corpus has its own
initials located beneath those of the tunica. In other words, the number of tiers of initials is
equal to the number of tunica layers plus one, the tier of corpus initials.

In contrast to the histogen theory, the tunica-corpus theory does not imply any relation
between the configuration of cells at the apex and histogenesis below the apex. Although the
epidermis usually arises from the outermost layer of tunica, which thus coincides with
Hanstein’s dermatogen, the underlying tissues may have their origin in the tunica or corpus or
both, depending on the plant species and the number of tunica layers.
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As more plants came to be examined, the tunica corpus concept underwent some
modifications, especially with regard to the strictness of the definition of the tunica.
According to one view, tunica should include only those layers that never show any periclinal
divisions in the median position, that is, above the level of origin of leaf primordia (Jentsch,
1957). If the apex contains additional parallel layers that periodically divide periclinally,
these layers are assigned to the corpus and the latter is characterized as being stratified
(Sussex, 1955; Tolbert and Johnson, 1966).

Other workers treat the tunica more loosely and describe it as fluctuating in number of
layers: one or more of the inner layers of the tunica may divide periclinally and thus become
part of the corpus. Because of the differing usage of the term tunica, its usefulness in
accurately describing growth relations at the shoot apex was questioned by Popham (1951),
who proposed the term mantle to include “all layers at the summit of the apex in which
anticlinal divisions are sufficiently frequent to result in the perpetuation of definite cell
layers”; the mantle overarches a body of cells called the core. The term corpus was avoided.

The tunica-corpus concept, which was developed with reference to angiosperm shoot
apices, proved to be largely unsuitable for the characterization of the apical meristem of
gymnosperms. With few exceptions (Gnetum, Ephedra, and several species of conifers) the
gymnosperms do not show a tunica-corpus organization in the shoot apex; that is, they do not
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have stable surface layers dividing only anticlinally. The outermost layer of the apical
meristem undergoes periclinal and anticlinal divisions and contributes cells to the peripheral
and interior tissues of the shoot.

Cytohistological zones of apical meristem

Studies of gymnosperm apices led to the recognition of a zonation—termed a
cytohistological zonation—based not only on planes of division but also on cytologic and
histologic differentiation and the degree of meristematic activity of the component cell
complexes. Similar zonations, superimposed on a tunica-corpus organization, have since been
observed in most angiosperms. The cytologic zones that may be recognized in shoot apical
meristems vary in degree of differentiation and in details of grouping of cells. The zonation
may be characterized by dividing the apical meristem into:

 A central mother zones

 Rib/pith meristem
 Peripheral meristem

Central zones give rise to two other zones. The rib zone, or rib (pith) meristem, appears
directly below the central zone and is centrally located in the apex. It usually becomes the
pith after additional meristematic activity has occurred. The other, the peripheral zone, or
peripheral meristem, encircles the other zones.

The peripheral zone typically is the most meristematic of all three zones and has the
densest protoplasts and the smallest cell dimensions. It may be described as a eumeristem.
Leaf primordia and the procambium arise here, as well as the cortical ground tissue. In
species with a tunica-corpus organization, the central zone corresponds to the corpus and the
portion(s) of the tunica layer(s) overlying the corpus.
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Inquires of initials and derivatives

The next development in the interpretation of the shoot apical meristem resulted from
the efforts of French cytologists (Buvat, 1955a; Nougarède, 1967). Meristematic activity
drew the chief attention of this work. Counts of mitoses, and cytological, histochemical, and
ultrastructural studies served to formulate the theory that after the apical structure is
organized in the embryo, the central zone of cells becomes the waiting meristem méristème
d’attente.

The méristème d’attente stays in a quiescent state until the reproductive stage is
reached, and meristematic activity is resumed in the distal cells. During the vegetative stage,
meristematic activity is centred in the initiating ring (anneau initial), corresponding to the
peripheral zone, and in the medullary (pith) meristem (méristème medullaire). The concept of
the inactive central zone in the apical meristem was extended from the shoots of angiosperms
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to those of gymnosperms (Camefort, 1956, who called the central zone “zone apicale” and
the seedless vascular plants and to roots.

With regard to root apices, the occurrence of an inactive centre in the meristem found
confirmation in many studies, resulting in the development of the concept of quiescent centre
by Clowes (1961). The revision of the concept of apical initials by the French workers
served as a considerable stimulant for further research on apical meristems. Counts of mitoses
in different regions of the shoot apex, feeding root tips with radio labelled compounds to
detect the location and synthesis of DNA, RNA, and protein, histochemical tests,
experimental manipulations, and tracing of cell patterns in fixed and living shoot apices
provided data that, in essence, corroborated the postulate of the relative infrequency of
mitotic activity in the central zone.

Recognition of the relative infrequency of mitotic activity in the central zone has not
led to an abandonment of the concept that the most distal cells are the true initials and the
ultimate source of all body cells in the shoot. Considering the geometry of the apex, one can
deduce a priori that in view of the exponential growth of the derivatives of the apical
meristem, a few divisions in the distalmost cells would result in the propagation of any
distinctive genome characteristic of these cells through large populations of cells.

As noted earlier, the tunica-corpus theory postulated the presence of a small group of
initials in each layer of the apical meristem. Clonal analyses are often cited as providing
evidence for one to three initials in each. The relation between the initials and the immediate
derivative in the apical meristem is flexible. A cell functions as an initial not because of any
inherent properties but because of its position. At the time of division of an initial it is
impossible to predict which of the two daughter cells will “inherit” the initial function and
which will become the derivative. It is also known that a given initial may be replaced by a
cell that through prior history would be classified as a derivative of an initial.

Since no cells are permanent initials, Newman (1965) maintained that in order to
understand structure and functioning of a meristem, a distinction must be made between the
“continuing meristematic residue”— that is, the source of cellular structure that functions as
initials—and the “general meristem,” a region of elaboration. Emergence of new cells from
the continuing meristematic residue is a very slow, continuous process of long duration,
whereas the passage of cells in the cells in the general meristem is a very rapid, continuous
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process of only short duration. This concept is used in Newman’s classification of apical
meristems designed for all groups of vascular plants:

 Monopodial, as in ferns—the residue is in the superficial layer and any kind of division
contributes to growth in length and breadth.
 Simplex, as in gymnosperms—the residue is in a single, superficial layer and both
anticlinal and periclinal divisions are needed for bulk growth.
 Duplex, as in angiosperms—the residue occurs in at least two surface layers with two
contrasting modes of growth, anticlinal divisions near the surface and divisions in at least
two planes deeper in the apical meristem.

Vegetative shoot apex

The vegetative shoot apex is a dynamic structure that in addition to adding cells to the
primary plant body, repetitively produces units, or modules, called phytomeres. Each
phytomere consists of a node, with its attached leaf, a subjacent internode, and a bud at the
base of the internode. The bud is located in the axil of the leaf of the next lower phytomere
and may develop into a lateral shoot.

In seed plants the apical meristem of the first shoot is organized in the embryo before
or after the appearance of the cotyledon or cotyledons. Vegetative shoot apices vary in shape,
size, cytologic zonation, and meristematic activity. The shoot apices of conifers are
commonly relatively narrow and conical in form; in Ginkgo and in the cycads, they are rather
broad and flat. The apical meristem of some monocots (grasses) and eudicots is narrow and
elongated, with the distal portion much elevated above the youngest node.
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Fig: Vegetative shoot apex

Before the initiation of each leaf the apical meristem widens considerably and after
the appearance of the leaf primordium it again becomes narrow. This phenomenon is
rhythmic, i.e. it recurs with the initiation of each leaf or pair of leaves. Schmidt (1924)
introduced the terms minimal- and maximal-areas of the apex. For the period between the
successive initiations of two leaves or two pairs of leaves he suggested the use of the term
plastochron which had been used previously but with a much wider meaning. The shoot
apices of dicotyledons with opposite leaves (such as Lonicera, Coleus, Vinca, Ligustrum,
Syringa and others) are particularly suitable for the study of plastochronic changes.
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Fig: Diagram of a longitudinal section of a eudicot shoot tip. Activity of the apical meristem,
which repetitively produces leaf and bud primordia, results in a succession of repeated units
called phytomeres. Each phytomere consists of a node, its attached leaf, the internode below
that leaf, and the bud at the base of the internode. The boundaries of the phytomeres are
indicated by the dashed lines. Note that the internodes are increasing in length the farther they
are from the apical meristem. Internodal elongation accounts for most of the increase in
length of the stem.

Root apex
In contrast to the apical meristem of the shoot, that of the root produces cells not only
toward the axis but also away from it, for it initiates the root cap. Because of the presence of
the root cap the distal part of the apical meristem of the root is not terminal but subterminal in
position, in the sense that it is located beneath the root cap. The root apex further differs from
the shoot meristem in that it forms no lateral appendages comparable to the leaves and no
branches. The root branches are usually initiated beyond the region of most active growth and
arise endogenously. Because of the absence of leaves the root apex shows no periodic
changes in shape and structure such as commonly occur in shoot apices in relation to leaf
initiation. The root also produces no nodes and internodes and therefore grows more
uniformly in length than the shoot, in which the internodes elongate much more than the
nodes. The rib-meristem type of growth is characteristic of the elongating root cortex.

Fig: Root apical meristem

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Root apical organization

The architecture, or cellular configuration, of the apical meristems of roots has been
studied most often for the purpose of revealing the origin of the tissue systems and has served
for the establishment of the so-called types (Schüepp, 1926; Popham, 1966) and for
discussions of the trends in the evolution of root apical organization. By analysing the
patterns of cells in an apical meristem, it is possible to trace out planes of cell division and
the direction of growth.

In one type of analysis the differentiating tissues are followed to the apex of the root
in order to determine whether there are specific cells that appear to be the source of one or
more of the discrete tissues. Thus, the implication is made that a spatial correlation of tissues
with certain cells or groups of cells at the apex indicates an ontogenetic relation between the
two, in other words, that the apical cells function as initials.

1. Apical cell theory

This theory was proposed by Nageli in 1878. According to this theory the entire root
develops from, a single apical cell while in others, the Marattiaceae, a few initials are present.
When there is only a single apical cell it is tetrahedral, and it divides in such a way so as to
add new cells to the body of the root from its upper-three sides and to the root cap from its
base. Allen (1947), working on Pseudotsuga distinguished a central group of permanent
initials with three groups of temporary initials (mother cells) on its periphery. He observed
that the meristem of the vascular cylinder developed from the first group of temporary
initials, the meristem of the cortex. from the second, and the columella from the third.

The columella is a group of cells that forms the longitudinal axis of the root cap. In it
the cells are arranged in longitudinal rows. Cells are added to the root cap from the columella
by periclinal cell division on its periphery. "the protoderm was seen to develop from the
young cortex. According to Wilcox's work there appear to be two groups of temporary
initials, one of which gives rise to the central cylinder and the other to the columella, from
which the root cap and cortex develop.

2. Histogen theory
The analysis of origin of root tissues in terms of distinct initials at the apex
corresponds to the approach used by Hanstein (1868, 1870) when he formulated the histogen
theory. As discussed earlier, Hanstein considered the body of the plant to arise from a
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massive meristem comprising three precursors of tissue regions, the histogens, each
beginning with one to several initials at the apex arranged in superposed tiers.

The histogens are the dermatogen (precursor of epidermis), the plerome (precursor of
the central vascular cylinder), and the periblem (precursor of the cortex). Although the
subdivision into the three histogens does not have universal application—it is seldom
discernible in shoots, and many roots lack a dermatogen in the sense of Hanstein (1870), that
is, an independent layer that gives rise to the epidermis—it has often been used for
descriptions of tissue regions in the root.

3. Korper kappe theory

Another approach to an analysis of the relationship between cell patterns and growth
in root tips is that represented by the body-cap (Körper-Kappe) concept of Schüepp (1917),
which emphasizes the planes of those divisions that are responsible for the increase in
number of vertical cell files in the meristematic region of the root. Many of the files divide in
two, and where they do so, a cell divides transversely; then one of the two new cells divides
longitudinally, and each daughter cell of this division becomes the source of a new file. The
combination of the transverse and the longitudinal division results in an approximately T- (or
Y-) shaped wall pattern, and therefore such divisions have been named T divisions.

The direction of the top stroke (horizontal bar) of the T varies in different root parts.
In the cap it is directed toward the base of the root, in the body toward the apex. Whereas a
clear boundary exists between the body and the cap in some roots (those with separate root
cap initials), in others the boundary is not sharply delimited (e.g., in Fagus sylvatica where
the transition between body and cap is very gradual.
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Fig: Körper -kappe theory

Apical organization in roots
There are two types of root apical organization is found in different plants. Two kinds of
organization are classified as the open and the closed, respectively (Guttenberg, 1960).

Adapting Guttenberg's view the meristems of the different tissue systems can be traced, in the
root apex, at various distances from the central cells (i.e. the permanent initials). In some
species the initials (temporary) of the various tissue systems are already discrete immediately
adjacent to the central cells, i.e., closed type. These initials represent those of the vascular
cylinder, the cortex, and the common initials of the protoderm and root cap, e.g. as in
Brassica, or the separate initials of the protoderm and the root cap, e.g., as in Zea mays and
Triticum. The special initials of the root cap were termed calyptrogen. In other species the
meristems of the different tissue systems finally become distinct only some distance away
from the central cells, i.e., open type. In this type. common initials for the cortex meristem,
root cap and protoderm or for the meristems of all the tissue systems (e.g., Allium) appear on
the periphery of the central cells. The importance of the above types is queried as they have
been observed to occur in the roots of a single species.
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The distinction between open and closed meristems is not always clear-cut. Both
types of meristem have been reported to originate from the closed pattern in the embryonic
root or the primordium of a lateral or adventitious root. During later elongation of the root the
closed pattern may be retained or replaced by an open one. In pea (Pisum sativum) both
embryonic and adult roots have open meristems.

The tetrahedral apical cell of the Equisetum root contributes both to the main body of
the root and to the root cap, but early root development in Equisetum is markedly different
from that of most ferns. Inasmuch as the root cap in Azolla is discrete from the rest of the
root, the apical meristem of the Azolla root is classified as closed. Conversely, the root apices
of Equisetum and ferns with apical cells that cut off cells from all four faces are classified as
open (Clowes, 1984).

The two types of apical organization in angiosperms, the closed and the open, require
separate consideration. The closed pattern is often characterized by the presence of three tiers
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or layers of initials. One tier appears at the apex of the central cylinder, the second terminates
the cortex, and the third gives rise to the root cap. The three-tiered meristems may be grouped
according to the origin of the epidermis. In one group, the epidermis has common origin with
the root cap and becomes distinct as such after a series of T divisions along the periphery of
the root.

In the second, the epidermis and cortex have common initials, whereas the root cap
arises from its own initials that constitute the root cap meristem, or (from the Greek calyptra,
veil, and Tenos, offspring; Janczewski, 1874). If the root cap and the epidermis have common
origin, the cell layer concerned is called dermatocalyptrogen (Roots with a
dermatocalyptrogen are common in eudicots (representatives of Rosaceae, Solanaceae,
Brassicaceae, Scrophulariaceae, and Asteraceae.

Groot et al. (2004) distinguish between two types of open root apical meristems in
eudicots, basic-open and intermediate-open. In the basic-open meristem, the cell files
terminate apically in a relatively large region of initials, and the fate of the initials’
derivatives are not immediately evident. In the intermediate-open meristem, the initial
region is much shorter than in the basic-open type, so the fate of a derivative is usually
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evident immediately after it has been cut off its initial cell. The cell files in intermediate-open
meristems appear to converge on the initial region, but the initials are shared between the root
cap and both the cortex and the vascular cylinder. Mapping root apical meristem organization
on a phylogenetic tree, Groot et al. (2004) determined that the intermediate-open meristem is
ancestral and the basic-open and closed types are derived

Quiescent region in root apex

The discovery by Clowes (1954, 1956) of a quiescent centre in the root apex brought
about a fundamental change of view about the behaviour of root meristems. Extensive
research on normally developing roots and on those treated surgically, has shown that, as a
general phenomenon, the initials that are responsible for the original cell pattern- the minimal
constructional centre largely cease to be mitotically active during the later growth of the root.
They are supplanted in this activity by cells on the margin of the relatively inactive region, or
quiescent centre. A quiescent centre arises twice in primary roots, first during embryogeny
and then again during the early stages of germination of the seed.

Fig: Quiescent centre concept

The relatively inactive state of the quiescent centre cells does not mean that they have
become permanently non-functional. Quiescent centre cells do divide occasionally and serve
to renew the more actively dividing regions around them, the cells of which are unstable and
displaced from time to time. At the height of the growing season, they exhibit a well-
developed quiescent centre, but during reactivation of growth early in the growing season, a
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quiescent centre is not discernible. When the root cap is removed, the cells of the quiescent
centre begin to grow and undergo a controlled sequence of divisions that regenerate the root
cap.

Growth of root tip

The region of actively dividing cells, the apical meristem extends for a considerable
distance basipetally from the apex, that is, toward the older part of the root. At one level of
organization both the root cap and the root proper may be envisioned as consisting of files of
cells that emanate from the promeristem.

Relatively close to the promeristem some files divide longitudinally either radially or
periclinally by T divisions to regions but also in the different cell files of the same tissue
region, and even in individual cells. Typically, the meristematic cortex vacuolates and
develops intercellular spaces close to the apex, where the central-cylinder meristem
(procambium) still appears dense. In the central cylinder, the precursors of the innermost
xylem vessels (metaxylem vessels) cease dividing, enlarge, and vacuolate considerably in
advance of the other precursors, and the first sieve tubes commonly mature in the part of the
root where cell division is still in progress. In individual cells, division, elongation, and
vacuolation are combined.

It is now clearly established that cell division continues well into the region where cell
length increases. Thus, a transition zone apparently exists in the basal part of the meristem
and the region where cells expand rapidly or, to be more exacting, “where cells are
undergoing their final division as well as expanding rapidly” (Beemster and Baskin, 1998). It
has been hypothesized that the division and elongation regions are coupled and may actually
constitute one developmental zone.

The tip of the root does not grow continuously at the same rate, especially in
perennial plants (the roots show periodic deceleration of growth and have periods of
dormancy (Wilcox, 1954). Dormancy is preceded by lignification of cell walls and by a
deposition of suberin—a double process called metacutization—in the cortex and root cap
throughout a layer of cells that is continuous with the endodermis and completely covers the
apical meristem. The latter thus becomes sealed off by a protective layer on all sides except
toward the base of the root.
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Fig: Growth of root tip

Conclusive remarks
Apical meristems are continuously growing tissue of the plants. These are responsible for the
primary growth of the plants. It mainly has two types shoot apical meristem and root apical
meristem. Different papers are published on the inquiries of apical initials and derivatives.
Apical meristem is delimited to three tissues which include protoderm, ground meristem and
procambium that forms the future mature tissue system of plants. Various attempts have been
made on the apical organization of shoot and roots. Some are comparative and aim at
conclusions of taxonomic significance, but others, and probably the more useful, are
concerned with the development of the particular plants under study. Proper developmental
studies demand a high degree of competence and are vital to an understanding of mature
plant.

References

Esau's Plant Anatomy: Meristems, Cells, and Tissues of the Plant Body: Their

Structure, Function, and Development, 3rd Edition by Ray F. Evert

Plant Anatomy -An applied approach by David F. Cutler

Plant Anatomy by A. Fahn

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https://courses.lumenlearning.com/boundless-biology/chapter/plant-development/

#:~:text=Apical%20meristems%20are%20organized%20into,meristem%20has%20a

%20particular%20function.

https://www.britannica.com/science/apical-meristem