Equisetum 180922130237

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EQUISETUM

Structure and Life Cycle


Habitat of Equisetum:
• The plant body of Equisetum has an aerial part and an underground
rhizome part. The rhizome is perennial, horizontal, branched and
creeping in nature. The aerial part is herbaceous and usually annual.
Majority of the species are small with a size range in between 15
and 60 cm in height and 2.0 cm in diameter.
• Equisetums generally grow in wet or damp habitats and are
particularly common along the banks of streams or irrigation
canals (E. debile, E. palustre). However, some species are adapted
to xeric condition (e.g., Equisetum arvense).
Some species of Equisetum are indicators of the mineral content of the

soil in which they grow. Some species accumulate gold (about 4.5

ounce per ton of dry wt.), thus they are considered as ‘gold indicator

plants.
Structure of Equisetum:
The Sporophyte:
The sporophytic plant body of Equisetum is differentiated into
 Stem
 Roots
 Leaves

 Stem:
The stem of Equisetum has two parts: perennial, underground, much-branched
rhizome and an erect, usually annual aerial shoot. The branching is monopodial,
shoots are differentiated into nodes and internodes.
• Some shoots are profusely branched, green (chlorophyllous) and purely vegetative. The

others are fertile, unbranched, brownish in colour (achlorophyllous) and have

terminal strobili.

• The underground rhizome and the aerial axis appear to be articulated or jointed due to the

presence of distinct nodes and internodes. Externally, the internodes have longitudinal ridges

and furrows and, internally, they are hollow, tube-like structures. The ridges of the successive

internodes alternate with each other and the leaves are normally of the same number as the

ridges on the stem.


Internal Features of Stem:

 In T.S., the stem of Equisetum appears wavy in outline with ridges and furrows.

The epidermal cell walls are thick, cuticularised and have a deposition of

siliceous material.

 Stomata are distributed only in the furrows between the ridges. A hypodermal

sclerenchymatous zone is present below each ridge which may extend up to

stele in E. giganteum. The cortex is differentiated into outer and inner regions.
 The outer cortex is chlorenchymatous, while the inner cortex is made up of

thin-walled parenchymatous cells. There is a large air cavity in the inner

cortex corresponding to each furrow and alternating with the ridges, known

as vallecular canal.

 New leaves and branches of Equisetum are produced by the apical

meristem, however, most of the length of the stem are due to the activity of

intercalary meristem located just above each node. The activity of

intercalary meristem causes rapid elongation of the inter- nodal region.


 The vascular bundles are arranged in a ring which lies opposite to the ridges in

position and alternate with the vallecular canals of the cortex. Vascular bundles

are conjoint, collateral and closed. In the mature vascular bundle, protoxylum is

disorganised to form a carinal cavity which lies opposite to the ridges.

 The vascular bundles remain unbranched until they reach the level of node. At

the nodal region, each vascular bandle trifurcates (divided into three parts).
The xerophytic features are:
(i) Ridges and furrows in the stem,

(ii) Deposition of silica in the epidermal cells,

(iii) Sunken stomata,

(iv) Sclerenchymatous hypodermis,

(v) Reduced and scaly leaves, and

(vi) photosynthetic tissue in the stem.


The hydrophytic characteristics on the other hand
are:
(i)Well-developed aerating system like carinal
canal, vallecular canal and central pith cavity.

(ii) Reduced vascular elements.


Root:

 The primary root is ephemeral. The slender adventitious roots arise endogenously at
the

nodes of the stems. In T.S., the root shows epidermis, cortex and stele from periphery to the

centre. The epidermis consists of elongated cells, with or without root hairs.

 The cortex is extensive; cells of the outer cortex often have thick walls (sclerenchymatous)

and those of the inner cortex are thinner parenchymatous.


 A large metaxylem element is present in the centre of the stele

and the protoxylem strands lie around it. The space between

the protoxylem groups is filled with phloem. There is no pith.


Leaves:
The leaves of Equisetum are small, simple, scale-like and isophyllous; they are
attached at each node, united at least for a part of the length and thus form a
sheath around the stem. The sheath has free and pointed teeth-like tips.

The number of leaves per node varies according to the species. The species
with narrow stems have few leaves (e.g., 2-3 leaves in E. scirpoides) and those
with thick stem have many leaves (e.g., up to 40 leaves in E. schaffneri).
The number of leaves at a node corresponds to the number of ridges

on the internode below. The leaves do not perform any

photosynthetic function and their main function is to provide

protection to young buds at the node.


Life cycle:
• Sex Organs of Equisetum:

• i. Antheridium:

• In monoecious species, antheridia develop later than archegonia. They are of

two types — projecting type and embedded type. Antheridia first appear on

the lobes of the gametophyte. The periclinal division of the superficial

antheridial initial gives rise to jacket initial and an androgonial cell.


The jacket initial divides anticlinally to form a single-layered jacket.

The repeated divisions of androgonial cells form numerous

cells which, on metamorphosis, produce spermatids/antherozoids.

The antherozoids escape through a pore created by the separation

of the apical jacket cell. The apical part of the antherozoid is spirally

coiled, whereas the lower part is, to some extent, expanded.


ii. Archegonium:

Any superficial cell in the marginal meristem acts as an archegonial initial which
undergoes

periclinal division to form a primary cover cell and an inner central cell. The cover cell,

by two vertical divisions at right angle to each other, forms a neck. The central cell

divides transversely to form a primary neck canal cell and a venter cell.
Two neck canal cells are produced from the primary neck canal cell. While,

the venter cell, by a transverse division, forms the ventral canal cell and

an egg.

At maturity, an archegonium has a projecting neck comprising of three to

four tiers of neck cells arranged in four rows, two neck canal cells of

unequal size, a ventral canal cell, and an egg at the base of the embedded

venter.
Fertilization:

Water is essential for fertilization. The gametophyte must be covered with a thin layer of

water in which the motile antherozoides swim to the archegonia. The neck canal cells

and ventral canal cell of the archegonia disintegrate to form a passage for the entry of

antherozoids.

Many antherozoids pass through the canal of the archegonium but only one of them

fuses with the egg. Thus diploid zygote is formed. Generally more than one archegonia

are fertilized in a prothallus.


Embryo (The New Sporophyte):

The embryo is the mother cell of the next sporophytic generation. Unlike most

pteridophytes, several sporophytes develop on the same prothallus. The first

division of the zygote is transverse. This results in an upper epibasal cell and

lower hypobasal cell. The embryo is therefore exoscopic (where the apical cell is

duacted outward. No suspensor is formed in Equisetum.


The epibasal and hypobasal cells then divide at right angles to the

oogonial wall, and as a result a tour-celled quadrant stage is established.

All the four cells of the quadrant are of different size and shape.

The four-celled embryo undergoes subsequent divisions and the future

shoot apex originate from the largest cell and leaf initials from the remai-

ning cells of one quadrant of the epibasal hemisphere.


Later the root grows directly downward and penetrate the gametophytic tissue

to reach the soil or substratum. A number of such sporophytes may

develop from a large mature gametophyte if more than one egg is fertilized

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