Meristems and Differentiation

• Embryonic tissue regions, primarily concerned

with formation of new cells, are the meristems.
• Greek merismos, division cell division activity as
a characteristic of a meristematic tissue
• Those cells that maintain the meristem as a
continuing source of new cells are referred to as
the initiating cells, or meristematic initials, or
simply initials.
• Their products, which after a variable number of
cell divisions give rise to the body cells, are the
derivatives of the initials.
• So, a dividing cell is a precursor of derivatives.
• A given initial in a meristem is a precursor of
two derivatives, one of which is a new initial
and the other a precursor of body cells.
• Initials are not inherently different from their
derivatives. The apical meristems of shoot and
root certain cells in the meristems act as
initials mainly because they occupy the proper
position for such activity.
• Initials are totipotent (from the Latin totus,
entire); they have the capacity to produce the
entire spectrum of cell types, even to develop
into complete plants. the development and
organization of a plant may be characterized
as having plasticity
 Classification of Meristems
• A Common Classification of Meristems Is
Based on Their Position in the Plant Body
• There are apical meristems, that is, meristems
located at the apices of main and lateral
shoots and roots, and lateral meristems, that
is, meristems arranged parallel with the sides
of the axis, usually that of stem and root. The
vascular and cork cambia are lateral
meristems.
• The third term based on the position of a
meristem is intercalary meristem. This term
refers to meristematic tissue derived from the
apical meristem and continuing meristematic
activity some distance from that meristem.
• The word intercalary implies that the meristem is
inserted (intercalated) between tissues that are
no longer meristematic. The best known
examples of intercalary meristems are those in
internodes and leaf sheaths of monocots,
particularly grasses.
• 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
The subjacent meristematic tissues are classified
according to the tissue systems that are derived from
them, namely into protoderm, which differentiates
into the epidermis; procambium (also called
provascular tissue), which gives rise to the primary
vascular tissues; and the ground meristem, precursor
of the fundamental or ground tissue system primary
meristems
• The terms protoderm, procambium, and
ground meristem correlated with the equally
simple and convenient classification of mature
tissues into the three tissue systems,
epidermal, vascular, and fundamental
 Meristems Are Also Classified
According to the Nature of Cells That
Give Origin to Their Initial Cells
• If the initials are direct descendants of
embryonic cells that never ceased to be
concerned with meristematic activity, the
resulting meristem is called primary. If,
however, the initials originate from cells that
had differentiated, then resumed
meristematic
 Characteristics of Meristematic Cells
• Meristematic cells are fundamentally similar to
young parenchyma cells. During cell division the
cells at the shoot apices are relatively thin-
walled, rather poor in storage materials, and
their plastids are in proplastid stages. The
endoplasmic reticulum is small in amount and
mitochondria have few cristae. Golgi bodies and
microtubules are present as is characteristic of
cells with growing cell walls. The vacuoles are
small and dispersed. Deeper layers of apical
meristems may be more highly vacuolated and
contain starch
• During periods of cell division, the vascular cambium
cells are highly vacuolated, with one or two large
vacuoles limiting the dense cytoplasm to a thin parietal
layer , which contains rough endoplasmic reticulum
and other cell components. ratio of cell size to nuclear
size—the cytonuclear ratio—varies considerably The
nuclei show characteristic structural variations during
changes in mitotic activitymeristematic cells vary in
size, shape, and cytoplasmic features. In recognition of
this variability, the term eumeristem (true meristem)
has been suggestsed for designating a meristem
composed of small, approximately isodiametric cells
with thin walls and rich in cytoplasm
 Growth Patterns in Meristems
• Meristems and meristematic tissues show varied arrangements of
cells resulting from differences in patterns of cell division and
enlargement. Apical meristems having only one initiating cell
occupying the very apex (Equisetum and many ferns) have an
orderly distribution of the recently formed, still meristematic cells
In seed plants the pattern of cell division appears less precise. It is
not random, however, for an apical meristem grows as an
organized whole and the divisions and enlargement of individual
cells are related to the internal distribution of growth and to the
external form of the apex. These correlative influences bring about
a differentiation of distinctive zones in the meristems. In some
parts of the meristem, the cells may divide sluggishly and attain
considerable dimensions; in others they may divide frequently and
remain small.
• Some cell complexes divide in various planes (volume growth), and
others only by walls at right angles to the surface of the meristem
(anticlinal divisions, surface growth). The lateral meristems are
particularly distinguished by divisions parallel with the nearest
surface of the organ (periclinal divisions), which result in
establishment of rows of cells parallel with the radii of the axes
(radial seriation or alignment) and an increase in thickness of the
organ. In cylindrical bodies, such as stems and roots, the term
tangential division (or tangential longitudinal) is commonly used
instead of periclinal division. If the anticlinal division occurs parallel
with the radius of the cylinder, it is referred to as a radial anticlinal
(or radial longitudinal) division . If the new wall is laid down at right
angles to the longitudinal axis of the cylinder, the anticlinal division
is transverse.
• Organs arising at the same apical meristem
may assume varied forms because the still-
meristematic derivatives of the apical
meristem (primary meristems) often exhibit
distinct patterns of growth. Some of these
patterns are so characteristic that the
meristematic tissues showing them have
received special names. These are mass
meristem (or block meristem), rib meristem
(or file meristem), and plate meristem.
• The mass meristem grows by divisions in all
planes and produces bodies that are
isodiametric or spheroidal or have no definite
shape. Such growth occurs, for example,
during the formation of spores, sperms (in
seedless vascular plants), and endosperm.
Divisions in various planes are associated with
the spheroidal shape of many angiosperm
embryos at a certain stage of development.
• The rib meristem gives rise to a complex of
parallel longitudinal fi les (“ribs”) of cells by
divisions at right angles to the longitudinal
axis of the cell row. This pattern of growth
occurs in elongating cylindrical plant parts as
illustrated by the cortex of roots and pith and
cortex of stems.
• The plate meristem shows chiefly anticlinal divisions
so that the number of layers originally established in
the young organ does not increase any further, and a
plate-like structure is produced. The fl at blades of
angiosperm leaves exemplify the result of growth by a
plate meristem (Fig. 5.5). The plate meristem and the
rib meristem are growth forms that occur mainly in the
ground meristem. They are associated with the two
basic forms of the plant body, the thin spreading
lamina (blade) of the leaf-like organs and the
elongated cylindrical plant parts such as root, stem,
and petiole.
 Meristematic Activity and Plant
Growth
• Meristems and their meristematic derivatives are
concerned with growth in the broad sense of the
term, meaning irreversible increase in size,
including volume and surface.
• In multicellular plants growth is based on two
processes, cell division and cell enlargement. The
recent derivatives of meristematic initials
produce other derivatives by cell division, and the
successive generations of derivatives increase in
size.
• Cell enlargement becomes dominant over cell division
and, in time, replaces it entirely.
• When the derivatives farthest from the meristem
initials cease to divide and to expand, they acquire the
characteristic specific for the tissues in which they are
located; that is, the cells differentiate and eventually
mature. Although cell division as such does not
contribute to the volume of the growing entity
• Cell division and cell expansion are different stages of
the growth process, in which cell enlargement
determines the final size of the plant and its parts.
• In ontogeny (development of an individual
entity), the enlargement of a cell serves as the
transition between the stages of division and
maturation. Cell division rarely occurs without
being accompanied by cell enlargement.
• The primary growth of the plant, initiated in the
apical meristems, expands the plant body,
determines its height, and through increase in its
surface, the area of contact with air and soil.
Eventually primary growth also gives rise to the
reproductive organs.
• The secondary growth resulting from the activity
of the cambia increases the volume of conducting
tissues and forms supportive and protective
tissues. Usually not all of the apical meristems
present on a plant are active simultaneously.
Suppression of lateral bud growth while the
terminal shoot is actively growing (apical
dominance).
• Then the activity of cambia varies in intensity,
may show seasonal fluctuations in meristematic
activity.
 DIFFERENTIATION

• Differentiation refers to the succession of

changes in form, structure, and function of
progenies of meristematic derivatives, and
their organization into tissues and organs. One
may speak of differentiation of a single cell, a
tissue (histogenesis), an organ
(organogenesis), and the plant as a whole.
• Differentiation also refers to processes by which
a fertilized egg cell produces a progeny of cells
showing increasing heterogeneity, specialization,
and patterned organization.
• The degree of differentiation and specialization
(structural adaptation to a particular function)
vary greatly.
• Some cells diverge relatively little from their
meristematic precursors and retain the power to
divide (various parenchyma cells).
• Others are more profoundly modified and lose most,
or all, of their meristematic potentialities (sieve
elements, laticifers, tracheary elements, various
sclereids). Thus cells differentiating in a multicellular
body become different from their meristematic
precursors, as well as from cells in other tissues of the
same plant. Mature cells with complete protoplasts
may resume meristematic activity when properly
stimulated. The stimulation can be induced by
accidental wounding, invasion by parasites, and
infection with agents of diseases on resumption of
meristematic activity by nonmeristematic cells the
terms dedifferentiation— loss of previously acquired
characteristics—and redifferentiation— acquisition of
new characteristics—are often used. The entire
process is referred to as transdifferentiation.
• During its development, a plant assumes a
specific form. Thus the plant undergoes
morphogenesis (from Greek words for shape
and origin). The term commonly is used with
reference to both external form and internal
organization, and just like differentiation,
morphogenesis is revealed at all levels of
organization, from cell components to whole
plants.
Senescence (Programmed Cell Death)
• The natural termination of life of a plant as a
result of senescence may be regarded as a
normal stage in plant development, a sequel
to the events of differentiation and
maturation.
• The term senescence specifically refers to the
series of changes in a living organism that lead
to its death.
• Senescence may affect the whole organism or some of
its organs, tissues, or cells. Annual plants that bloom
only once in their lifetime (monocarpy: fruiting once
only) senesce within one season.
• In deciduous trees the leaves commonly senesce at the
end of seasonal growth. Since senescence occurs in
orderly sequences in the life of the plant and is an
active degenerative process, it is considered to be
genetically controlled, or programmed—a process of
programmed cell death.
Cellular Changes in Differentiation
• During differentiation, histologic diversity results from
changes in the characteristics of individual cells and
from alterations in intercellular relationships. The
common features of more or less differentiated cells,
including the structure and function of individual cell
components,
• The changes in cell wall structure during cell
differentiation
• Differential increase in thickness of primary and
secondary cell walls, changes in cell wall texture and
chemistry, and development of special sculptural
patterns introduce differences among cells.
• Endopolyploidy refers to a condition that
arises from a DNA replication cycle within the
nuclear envelope and without spindle
formation. Hence the newly formed strands of
DNA remain in the same nucleus, and the
nucleus becomes polyploid. This type of DNA
replication cycle is called an endocycle
 One of the Early Visible Changes in
Differentiating Tissues Is the Unequal
Increase in Cell Size
• The increase in cell size may be relatively
uniform along all diameters, but frequently
the cell enlarges more in one direction than in
others. Such cells may become strikingly
different in shape from their meristematic
precursors.
 Intercellular Adjustment in
Differentiating Tissue Involves
Coordinated and Intrusive Growth
• Enlargement and change in shape of cells in a
differentiating tissue are accompanied by
more or less profound changes in the spatial
relations between cells.
• With regard to the growth of cell walls during tissue
differentiation, two kinds of intercellular adjustments
are visualized: (1) contiguous growing wall layers
belonging to two neighboring cells do not separate but
expand jointly; (2) contiguous wall layers do separate
and the growing cells intrude into the resulting spaces.
The fi rst method of growth, originally named
symplastic Growth The intercellular adjustment that
involves an intrusion of cells among others is called
intrusive growth or interpositional growthThe
elongating cells grow at their apices (apical intrusive
growth), usually at both ends.
CAUSAL FACTORS IN DIFFERENTIATION
• Use of growth-regulating chemicals, surgical procedures,
exposure to radiation, confinement to controlled
temperatures and light, interference with normal gravity
effects, and growth under selected day length conditions.
• Polarity refers to the orientation of activities in space. It is
an essential component of biological pattern formation.
• Polarity manifests itself early in the life of the plant. It is
displayed in the egg cell, in which the nucleus is located in
the chalazal end and a large vacuole in the micropylar end,
and in the bipolar development of the embryo from the
zygote. It is later expressed in the organization of the plant
into root and shoot and is also evident in various
phenomena at the cellular level.
• Polarity is related to the phenomenon of
gradients, since the differences between the
two poles of the plant axis appear in graded
series. There are physiological gradients, for
example, those expressed in the rates of
metabolic processes, in concentration of
auxins, and in concentration of sugar in the
conducting system; there are also gradients in
anatomical differentiation and in the
development of the external features
Plant Cells Differentiate According to
Position
• PLANT HORMONES from the Greek horman,
meaning to set in motion Tissues may require
different amounts of hormones. Such
differences are referred to as differences in
sensitivity