University of Salahaddin College of Agricultural
Engineering Science
Secondary Thickening in monocots
Prepared by: Rezheen Abdullah M Sadeeq
Course: General Botany
Stage: 1st
Date: 23th Feb 2024
Supervised by: Dr. Sirwan T. Aldabbagh
Table of contents
Introduction............................................................................................................. 1
Physical characteristics......................................................................................... 2
Evolution.................................................................................................................. 3
Primary and secondary thickening meristems in monocots............................3
Vascular tissue........................................................................................................ 5
Xylem........................................................................................................................ 5
Phloem...................................................................................................................... 7
Summary.................................................................................................................. 8
References............................................................................................................... 9
List of figures
Figure 1: typical monocot plant with six petals................................................1
Figure 2: Germination of a monocot and a eudicot. (Top) In a corn seed
(monocot), nutrients are stored in the cotyledon and endosperm tissue.
The radicle and hypocotyl (region between the cotyledon and radicle) give
rise to the roots. The epicotyl (region above the cotyledon) gives rise to
the stem and leaves and is covered by a protective sheath (coleoptile).
(Bottom) In a bean seed (eudicot), all nutrients are stored in the enlarged
cotyledons. The radicle gives rise to the roots, the hypocotyl to the lower
stem, and the epicotyl to the leaves and upper stem......................................2
Figure 3: Yellow Cymbidium orchids...................................................................3
Figure 4: Diagram of monocot shoot apical organization showing PTM near
shoot apex................................................................................................................ 4
Figure 5: Monocot growth in thickness. Cordyline indivisa (monocot:
Asparagaceae), transverse section of stem, showing outer cortex and
inner region containing scattered vascular bundles. The STM has produced
secondary vascular bundles that are radially aligned. Scale = 100 µm........5
Figure 6: Vascular bundle. Lilium tigrinum (monocot: Liliaceae), transverse
section of stem vascular bundle with xylem (left) and phloem (right)
encircled by a ring of bundle sheath cells. bs = bundle sheath, c =
companion cell, mx = metaxylem vessel, px = protoxylem vessel, s = sieve
tube element. Scale = 50 µm................................................................................ 6
Introduction
Since cotyledons are part of a plant's embryo and seed; they are referred to
as seed leaves. They are the first leaves germinating from the plant, but they
are not actual leaves. Cotyledon is also referred to as a seed-bearing plant's
embryonic leaf, that is to say, the first leaf to emerge from a germinating
seed. Cotyledons are used to differentiate between angiosperms which are
flowering plants. So, are cotyledons the same in all plants? The answer to
this question is no because there are two types of cotyledons. The two types
are dicotyledon and monocotyledon. Some plants do not have cotyledons at
all. Monocotyledon is also known as monocots. Monocot definition is the
plant that contains one cotyledon or one embryonic leaf. Monocotyledons are
approximately 60,000 plant species. The existence of only one seed
leaf (cotyledon) identifies the majority of monocotyledons. Monocotyledons
have one seed, flower petals in multiples of 3 or 3, fibrous roots, long thin
leaves with parallel veins, lack secondary growth, and its vascular bundle is
scattered. Cotyledons are known as seed leaves since they are part of a
plant's embryo and seed. They are the plant's first leaves to germinate,
although they are not true leaves. Examples of monocot plants are garlic,
banana, onion, palm trees, and lilies. These plants are characterized by
having one cotyledon. The provided figure shows a typical monocot plant with six
petals.
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Figure 1: typical monocot plant with six petals.
Physical characteristics
Monocot plants are marked by seeds with a single cotyledon, parallel-
veined leaves, scattered vascular bundles in the stem, the absence of a
typical cambium, and an adventitious root system. Flower parts typically
come in multiples of three, and the pollen grains characteristically feature a
single aperture (or furrow). The roots of a monocot lack a vascular cambium
(the area of secondary xylem and phloem, or secondary vascular tissue,
development) and therefore have no means of secondary thickening. In
other structural respects, monocot roots are essentially similar to those of
eudicots. Many eudicots have a taproot or several strong roots, with several
orders of branch roots, all originating eventually from the
embryonic root (radicle). The taproot or primary roots in such a system have
a vascular cambium and are thickened by secondary growth. This kind of
root system is not available to monocots. Instead, the primary root that
originates from the radicle of the embryo soon aborts or is undeveloped so
that no primary root is produced. The root system of monocots is thus
wholly adventitious—i.e., the roots originate laterally from the stem or from
the hypocotyl (the region of transition between the root and the stem in the
embryo).Figure 2 shows the germination
of a monocot. The roots are all slender,
and the plant is said to be fibrous-rooted.
Flowers of monocots differ from those of
eudicots mainly in the number of parts of
each kind. Monocot flowers most often
have the parts in sets of three,
occasionally four, but almost never five.
The numbers are especially characteristic
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of the sepals and petals. The stamens and pistils may be numerous even
when the perianth is trimerous (in sets of three), or the single ovary may
have only two carpels instead of three. Often there are six stamens,
representing two whorls of three.
Figure 2: Germination of a monocot and a eudicot. (Top) In a corn seed (monocot),
nutrients are stored in the cotyledon and endosperm tissue. The radicle and hypocotyl
(region between the cotyledon and radicle) give rise to the roots. The epicotyl (region
above the cotyledon) gives rise to the stem and leaves and is covered by a protective
sheath (coleoptile). (Bottom) In a bean seed (eudicot), all nutrients are stored in the
enlarged cotyledons. The radicle gives rise to the roots, the hypocotyl to the lower stem,
and the epicotyl to the leaves and upper stem.
Evolution
Monocots form a monophyletic group, meaning that they share a common
evolutionary history. It is widely believed that the monocots
were derived from primitive eudicots. Given that the various physical
features of monocots are regarded as derived characteristics within the
angiosperms, any plant more primitive than the monocots in these several
respects would certainly be a eudicot. Some of the earliest known monocot
fossils are pollen grains dating to the Aptian Age of the Early Cretaceous
Epoch (125 million–113 million years ago). Molecular clock studies (which
employ differences in DNA to estimate when a group split from its ancestors)
suggest that monocots may have originated as early as 140 million years
ago. Evolutionary diversification among the monocotyledons appears to have
been constrained by a number of fundamental features of the group, most
notably the absence of a typical vascular cambium and the parallel-veined
rather than net-veined leaves. Within these constraints, the monocots show
a wide range of diversity of structure and habitat. They are cosmopolitan in
their distribution on land. They also grow in lakes, ponds, and rivers,
sometimes free-floating but more often rooted to the bottom. Some of them
grow in the intertidal zone along the seashore, and a few are submerged
marine plants rooted to the bottom in fairly shallow water along the shore
look at Figure 3.
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Figure 3: Yellow Cymbidium orchids
Primary and secondary thickening meristems in monocots
In monocots, which lack a vascular cambium, a limited degree of radial
growth is achieved by a primary thickening meristem (PTM) near the
vegetative shoot apex Figure 4. The PTM is especially extensive in species
with short internodes and crowded leaves, such as bulbous taxa. In
monocots, the PTM originates in ground tissue in the pericyclic region of the
stem. It is a tiered meristem, consisting of a zone of meristematic cells that
produces radial derivatives. The majority of its radial derivatives are
centripetal, consisting of both parenchyma and discrete vascular bundles,
though it also produces some parenchyma towards the outside. In addition to
primary stem thickening, the PTM forms linkages between roots, stem and
leaf vasculature, especially in relatively condensed bulbous plants. It
typically ceases activity at a short distance behind the apex, and subsequent
stem thickening is limited.
Figure 4: Diagram of monocot shoot apical organization showing PTM near shoot
apex
However, in some species, this region retains some meristematic potential
and resumes meristematic activity further down the stem; it represents the
site of adventitious root production in some monocots109. Althoug
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hmonocotsare predominantly herbaceous, some species achieve
considerable height and girth. Tree-forming palms possess an extensive PTM
that results in a large sunken shoot apex; in these species, further stem
thickening occurs by subsequent division and enlargement of ground
parenchyma cells, termed diffuse secondary growth. A PTM is not confined to
monocots; similar pericyclic meristems also occur in some eudicots with
thick stems (e.g. Cactaceae). In some woody monocots of the order
Asparagales (e.g. Agave, Aloe, Cordyline, Yucca), considerable increase in
stem thick ness is achieved by means of a secondary thickening meristem
(STM) Figure 5. The STM resembles the PTM in that it is radially located in the
pericyclic region of the stem within the cortex and it produces radial
derivatives, but it occurs further from the shoot apex; the two meristems are
best regarded as developmental phases of the same meristem. The STM
produces secondary vascular bundles that are mostly amphivasal and
radially elongated. In some woody monocots([Link] recurvata,
Cordyline terminalis), the PTM and STM are axially discontinuous, whereas in
others (e.g. Yucca whipplei) they are axially continuous.
Figure 5: Monocot growth in thickness. Cordyline indivisa (monocot:
Asparagaceae), transverse section of stem, showing outer cortex and inner region
containing scattered vascular bundles. The STM has produced secondary vascular
bundles that are radially aligned. Scale = 100 µm
The PTM and STM, although sometimes termed the monocot cambium, are
not homologous with the vascular cambium. The PTM and STM originate from
the pericyclic region, whereas the vascular cambium originates inside stem
vascular bundles (fascicular cambium) and subsequently spreads to inter
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fascicular regions. The PTM and STM produce discrete vascular bundles
centrifugally, whereas the vascular cambium is a bifacial meristem that
produces xylem centrifugally and phloem centripetally.
Vascular tissue
Vascular tissue consists of associated networks of cells that conduct water
(xylem) and nutrients (phloem). Primary vascular tissue is derived from the
procambium, which is itself produced by the apical meristems. Secondary
vascular tissue is derived from the vascular cambium in eudicots, and from
the secondary thickening meristem in a few monocotspecies. Both xylem
and phloemare complex tissues compose dofmany different cellt ypes.
Xylem
The primary function of xylem is transport of water throughout the plant,
from roots via the stems to the leaves, where water is combined with carbon
dioxide to make carbohydrates and oxygen during photosynthesis. Xylem is
composed of several distinct cell types, often including parenchyma and
fibres as well as vessels. Tracheids and vessel elements (collectively termed
tracheary elements) are the water-conducting cells; they lack contents at
maturity and are linked into cell chains to form vessels. Tracheary elements
are elongated cells with thickened lignified walls. In a stem vascular bundle
Figure 6, the first-formed (protoxylem) elements often possess wall
thickenings that are either helical or arranged in rings (annular). Later-
formed primary tracheary elements (metaxylem) and second arytracheary
elements possess bordered pits in their lateral walls. Bordered pits18 can be
oval, polygonal or elongated (scalariform); they can be organized in
transverse rows (opposite pitting) or tightly packed (alternate pitting). The
primary difference between tracheids and vessel elements is that vessel
elements possess large perforations in their adjoining end walls, whereas
perforations are absent from tracheids. Perforation plates are generally
either simple, with a single opening, or scalari form, with a ladder-like row of
openings divided by a series of parallel bars, or rarely a reticulate mesh.
Vessel elements differ considerably in diameter, not only between different
species but also sometimes across a single growth ring (e.g. in secondary
xylem of oak).
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Figure 6: Vascular bundle. Lilium tigrinum (monocot: Liliaceae), transverse
section of stem vascular bundle with xylem (left) and phloem (right) encircled by
a ring of bundle sheath cells. bs = bundle sheath, c = companion cell, mx =
metaxylem vessel, px = protoxylem vessel, s = sieve tube element. Scale = 50
µm.
Phloem
Phloem has complex roles in translocation of nutrients (sucrose and
electrolytes) and hormones throughout the plant39. Although commonly
associated with xylem, phloem can develop precociously in regions that
require a plentiful supply of nutrients, such as developing sporogenous
tissue. Phloem consists of conducting cells (sieve elements) and associated
specialized parenchyma cells (companion cells) Figure 6; these two closely
interdependent cell types are produced from a common parent cell
(meristemoid) that divides and develops asymmetrically to form a larger
sieve element and smaller companion cell. Most angiosperms possess sieve-
tube elements rather than the relatively unspecialized sieve cells. Plastids At
maturity, sieve elements lack nuclei and most organelles but retain and
phloem-specific proteins (P-proteins). Companion cells are densely
cytoplasmic, retaining nuclei and many active mitochondria. Sieve element
plastids and P-proteins occur in several morphological forms (amorphous,
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filamentous, tubular and crystalline) that are often highly characteristic for
particular plant families and are thus of systematic and evolutionary value.
Sieve-element plastids are classified according to their inclusions: starch (S-
type plastids), protein (P-type plastids), or both. Sieve elements are linked
axially to form sieve tubes via sieve plates. Slime plugs are formed when P-
protein accumulates on a sieve plate. The walls of sieve elements are thin
and possess characteristic regions (sieve areas) that connect adjacent sieve
elements; sieve areas consist of groups of pores and associated callose. In
sieve tube elements, the sieve areas are localized on the adjoining end walls,
forming sieve plates that are either simple or compound.
Summary
Since cotyledons are part of a plant's embryo and seed; they are referred to
as seed leaves. Cotyledon is also referred to as a seed-bearing plant's
embryonic leaf, that is to say, the first leaf to emerge from a germinating
seed. Monocot definition is the plant that contains one cotyledon or one
embryonic leaf. Cotyledons are known as seed leaves since they are part of a
plant's embryo and seed. They are the plant's first leaves to germinate,
although they are not true leaves. Monocot plants are marked by seeds with
a single cotyledon, parallel-veined leaves, scattered vascular bundles in the
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stem, the absence of a typical cambium, and an adventitious root system.
The roots of a monocot lack a vascular cambium and therefore have no
means of secondary thickening. In other structural respects, monocot roots
are essentially similar to those of eudicots. Many eudicots have a taproot or
several strong roots, with several orders of branch roots, all originating
eventually from the embryonic root Instead, the primary root that originates
from the radicle of the embryo soon aborts or is undeveloped so that no
primary root is produced. The root system of monocots is thus wholly
adventitious—i.e., the roots originate laterally from the stem or from the
hypocotyl.
References
1- BokeNH.1951. Histogenesis of the vegetative shoot in Echinocereus.
American Journal of Botany 38: 23–38.
2- DeMasonDA.1983. the primary thickening meristem: definition and
function in monocotyledons. American Journal of Botany 70: 955–962.
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3- Diggle PK and DA DeMason. 1983. The relationship between the
primary thickening meristem and the secondary thickening meristem
in Yucca whipplei Torr. II. Ontogenetic relationship within the
vegetative stem. American Journal of Botany 70: 1205–1216.
4- Evert RF. 2006. Meristems, cells and tissues of the plant body: their
structure, function and development. John Wiley & Sons, Inc., New
York, USA.
5- [Link] growth in
monocotyledons. Botanical Review 57: 150–163.
6- Stevenson DW. 1980. Radial growth in Beaucarnea recurvata.
American Journal of Botany 67: 476–489.
7- Stevenson DW and JB Fisher. 1980. The developmental relationship
between primary and secondary thickening growth in Cordyline
(Agavaceae). Botanical Gazette 141: 264–268.
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