Apical meristem:

delimitation of different
growth zones
• A meristem is the tissue in most plants containing undifferentiated cells
(meristematic cells), found in zones of the plant where growth can take place.
• Meristematic cells give rise to various organs of the plant and keep the plant
growing.
• The shoot apical meristem (SAM) gives rise to organs like the leaves and flowers,
while the root apical meristem (RAM) provides the meristematic cells for the
future root growth.
• SAM and RAM cells divide rapidly and are considered indeterminate, in that they
do not possess any defined end status.
• In that sense, the meristematic cells are frequently compared to the stem cells in
animals, which have an analogous behavior and function.
• The term meristem was first used in 1858 by Karl Wilhelm von Nägeli (1817–
1891) in his book Beiträge zur Wissenschaftlichen Botanik.
• It is derived from the Greek word merizein (μερίζειν), meaning to divide, in
recognition of its inherent function.
• In general, differentiated plant cells cannot divide or produce cells of a different
type.
• Therefore, cell division in the meristem is required to provide new cells for
expansion and differentiation of tissues and initiation of new organs, providing
the basic structure of the plant body.
• Meristematic cells are incompletely or not at all differentiated, and are capable of
continued cellular division (youthful).
• Furthermore, the cells are small and protoplasm fills the cell completely.
• The vacuoles are extremely small.
• The cytoplasm does not contain differentiated plastids (chloroplasts or
chromoplasts), although they are present in rudimentary form (proplastids).
• Meristematic cells are packed closely together without intercellular cavities. The
cell wall is a very thin primary cell wall.
• Maintenance of the cells requires a balance between two antagonistic processes:
organ initiation and stem cell population renewal.
• Apical meristems are the completely undifferentiated (indeterminate) meristems
in a plant.
• These differentiate into three kinds of primary meristems.
• The primary meristems in turn produce the two secondary meristem types.
• These secondary meristems are also known as lateral meristems because they are
involved in lateral growth.
• At the meristem summit, there is a small group of slowly dividing cells, which is
commonly called the central zone.
• Cells of this zone have a stem cell function and are essential for meristem
maintenance.
• The proliferation and growth rates at the meristem summit usually differ
considerably from those at the periphery.
Tunica-Corpus model of the apical meristem (growing tip). The epidermal (L1)
and subepidermal (L2) layers form the outer layers called the tunica. The inner L3
layer is called the corpus. Cells in the L1 and L2 layers divide in a sideways
fashion, which keeps these layers distinct, whereas the L3 layer divides in a more
• Shoot apical meristems
• The shoot apical meristem is the site of most of the embryogenesis in flowering
plants.
• Primordia of leaves, sepals, petals, stamens and ovaries are initiated here at the
rate of one every time interval, called a plastochron.
• It is where the first indications that flower development has been evoked are
manifested.
• The shoot apical meristem consists of 4 distinct cell groups:
• Stem cells
• The immediate daughter cells of the stem cells
• A subjacent organising centre
• Founder cells for organ initiation in surrounding regions
• The four distinct zones mentioned above are maintained by a complex signalling
pathway.
• In Arabidopsis thaliana, 3 interacting CLAVATA genes are required to regulate the
size of the stem cell reservoir in the shoot apical meristem by controlling the rate
of cell division.
• CLV1 and CLV2 are predicted to form a receptor complex (of the LRR receptor like
kinase family) to which CLV3 is a ligand.
• CLV3 shares some homology with the ESR proteins of maize, with a short 14
amino acid region being conserved between the proteins.
• Proteins that contain these conserved regions have been grouped into the CLE
family of proteins.
• CLV1 has been shown to interact with several cytoplasmic proteins that are most
likely involved in downstream signalling.
• For example, the CLV complex has been found to be associated with Rho/Rac
small GTPase-related proteins.
• These proteins may act as an intermediate between the CLV complex and a
mitogen-activated protein kinase (MAPK), which is often involved in signalling
cascades.
• KAPP is a kinase-associated protein phosphatase that has been shown to interact
with CLV1. KAPP is thought to act as a negative regulator of CLV1 by
dephosphorylating it.
• Another important gene in plant meristem maintenance is WUSCHEL (shortened
to WUS), which is a target of CLV signalling.
• WUS is expressed in the cells below the stem cells of the meristem and its
presence prevents the differentiation of the stem cells.
• CLV1 acts to promote cellular differentiation by repressing WUS activity outside of
the central zone containing the stem cells.
• STM also acts to prevent the differentiation of stem cells by repressing the
expression of Myb genes that are involved in cellular differentiation.
Organisation of an apical meristem (growing tip)
1 - Central zone
2 - Peripheral zone
3 - Medullary (i.e. central) meristem
4 - Medullary tissue
• Root apical meristem
• Unlike the shoot apical meristem, the root apical meristem produces cells in two
dimensions.
• It harbors two pools of stem cells around an organizing center called the quiescent
center (QC) cells and together produce most of the cells in an adult root.
• At the apex, root meristem is covered by the root cap, which protects and guide
its growth trajectory.
• Cells are continuously sloughed off the outer surface of the root cap.
• The QC cells are characterized by their low mitotic activity.
• Evidence suggests that the QC maintains the surrounding stem cells by preventing
their differentiation, via signal(s).
• This allows a constant supply of new cells in the meristem required for continuous
root growth.
• Recent findings indicate that QC may also act as a reservoir of stem cells to
replenish whatever is lost or damaged.
• (a) Root meristem anatomy
• Primary roots arise through controlled cell divisions in the apical meristem and
subsequent expansion and differentiation of these cells.
• Root formation (rhizogenesis) is usually extremely rapid: daughter cells exiting
the meristem may be found 24 h later in a fully differentiated structure (e.g.
phloem), even though further modifications to cell function are still possible (e.g.
formation of an exodermis).
• Critical steps in setting up dimensions and thickness of the root axis, and supply
of cells to the zone of elongation, are the rate and position of cell divisions in the
meristem.
• Divisions can be in any of three planes, either anticlinal (normal to the root axis),
periclinal (tangential to the root axis) or radial to the axis.
• These divisions will give rise, respectively, to increased root length, increased root
thickness (more layers of cells through the root), or increased root circumference.
Longitudinal section through a primary root
tip of radish (Raphanus sativus).
Files of cells extend forward from the centre
of the apex to form the root cap, and
backwards to form the main root tissues.
• Separate cell divisions at the leading edge of the root meristem generate a root
cap which extends forward as a protective structure.
• The central cells of the root cap are often oriented in longitudinal arrays
(columella) and are destined for rapid attrition as the ‘advancing’ soil particles
slough off the surface layers.
• These cells also fulfil a vital chemico-physical role by secreting a glycoprotein-rich
mucigel which reduces friction between root and soil matrix.
• Root caps advance at a dramatic speed: a root might elongate by 5 cm per day
and new root cap cells can be pushed in advance of the apex of the primary axis
at about the same rate.
• This means that the leading tip of a primary root, supplied with new root cap
cells, advances through the soil at up to 60 µm min–1.
• Another remarkable feature of root apices is the quiescent centre, a paradox at
the heart of the meristem.
• The quiescent centre is a zone of relatively inactive, slowly dividing cells,
numbering about 500–600 in a mature maize root.
• Its discovery (Clowes 1959) involved studies on mitotic frequencies, thymidine
incorporation into nuclear DNA, and ploidy after colchicine treatment.
• The quiescent centre functions as a reserve of cells which can survive stresses
and provide cells to a regenerating meristem.
• Recovery from surgical removal of parts of the meristem and irradiation to
destroy dividing cells supports this concept.
• Likewise, short determinate lateral roots often lack a quiescent centre, suggesting
it is closely tied to sustained indeterminate development.
Most root apices contain a quiescent centre of very slowly dividing cells. (a)
Diagram of longitudinal section through a maise (Zea mays) root tip. The quiescent
centre is shaded dark green. (b) Autoradiograph of transverse section through root
apex of Vicia faba (broad bean), fed for 24 h with radioactive [ 3H]thymidine which
specifically labels DNA in nuclei of dividing cells. The quescent centre has
significantly fewer labelled cells (dark silver grains).
• The changes which cells undergo in root meristems are profound; mitotic activity
is most rapid in the distal regions, but the mitotic cycle slows dramatically within
0.5–1.0 mm.
• Surprisingly, mitotic frequency in adjacent cell files can vary widely.
• The rate and plane of cell division and subsequent rate of cell elongation
determine the rate of delivery of new cells to mature root tissues.
• The coordination of cell flux is presumably under tight control, achieving the final
anatomical outcomes recognisable as mature roots — single layers of pericycle
and endodermal cells, long conducting vessels and epidermal cells are some
examples.
• The role of growth in cells exiting the meristem and the direction of expansion
are major factors in rhizogenesis, with a 30- to 150-fold volume expansion
required to generate the primary axis.
Cell division rates vary along different zones of the root apical meristem. Frequency
of observed cell division is represented by different densities of stippling (onion
root tip).