EQUISETUM

Sushmitha K M
I MSc.Botany
 EQUISETUM
• SYSTEMATIC POSISTION:
Division - Sphenophyta
Class - Sphenopsida
Order - Equisetales
Family - Equisetaceae
Genus - Equisetum
 DISTRIBUTION AND HABITAT
• Equisetum is almost world-wide in distribution with 15 species and confined to N.
Temperate regions, though some are met with in Tropics also, except Australia.
• From India four species are known E. arvense, E. diffusum, E. palustre and E.
ramosissimum. Species of Equisetum are usually known as ―horse tails or pipes or
scouring rushes.
• Plants are small to large, terrestrial; usually grow in wet or marshy places or in open,
sunny sand banks along rivers and margins of lakes.
• Many species in this genus prefer wet sandy soils, though some are semi-
aquatic and others are adapted to wet clay soils.
 STRUCTURE OF EQUISETUM
• External features: The plant body of Equisetum is differentiated into stem, leaves
and roots.
• Rhizome: The Equisetum sporophyte develops a usually perennial, much branched
underground rhizome. The rhizome produces true roots. The rhizome divided into
nodes and internodes; scales present at nodes are fused to form a sheath.
• Roots: The roots are adventitious except the primary root. They arise in whorls from
the base of the branch primordia at each node of rhizome. The roots are slender and
fibrous, but sometime branched.
• 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.
• 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.
•
• In majority of the species, all the shoots are alike and chlorophyllous and some of
them bear strobili at their apices (e.g., E. ramosissimum, E. debile). Sometimes
shoot shows dimorphism (two types of shoots i.e., vegetative and fertile) e.g., E.
arvense.
• 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. These are schizolysigenous canals extending the entire length of internodes
and form a distinct aerating system.
• The stele is ectophloic siphonestele which is surrounded by an outer endodermal layer.
An inner endodermis is also present in some species of Equisetum (e.g., E.
sylvaticum). The endoder­mis is followed by a single-layered pericycle.
• 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, protoxylem is disorganised to
form a carinal cavity which lies opposite to the ridges.
• The metaxylem tracheids (scalariform or reticulate) are present on both sides of the
phloem. In some species vessels with reticulate perforations are reported. The central
part of the internode of aerial shoot is occupied by a large pith cavity which is formed
due to rapid elongation of the intermodal region.
 INTERNAL STRUCTURE OF ROOT
• 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. The
stele is protostelic where the xylem is triarch or tetrarch, or, in smaller roots, may be
diarch.
• 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.
•
 REPRODUCTION
• Equisetum reproduces vegetatively and by means of spores.
Vegetative Reproduction:
• The subterranean rhizomes of some species (e.g., E. telmateia, E. arvense) form
tubers which, on separation from the parent plant, germinate to produce new
sporophytic plants. The tubers develop due to irregular growth of some buds at the
nodes of the rhizomes.

Reproduction by Spores:
• Spores are produced within the sporangia. The sporangia are borne on the
sporangiophores which are aggregated into a compact structure termed strobilus or
cone or sporangiferous spike.
•
Strobilus:
• The strobilus are terminal in position and generally are borne terminally on the chloro­phyllous
vegetative shoot. However, they may be borne terminally on a strictly non- chlorophyllous axis
(e.g., E. arvense).
• The stro­bilus is composed of an axis with whorls of spo­rangiophores. Each sporangio­phore is a
stalked structure bearing a hexagonal peltate disc at its distal end. On the under surface of the
sporangiophore disc 5-10 elongate, cylindrical hanging sporangia are borne near the periphery
in a ring.
• The flattened tips of the sporangiophores fit closely together which provide protection to the
developing spo­rangia. The axis bears a ring-like outgrowth, the so-called annulus immediately
below the whorls of sporangiophores which provide additional protection during early
development.
• The annu­lus has been interpreted as a rudimentary leaf sheath by some botanists, whereas others
con­sider it to be sporangiophoric in nature as occa­sionally it bears small sporangia.
Equisetum. Structure of Strobilus: A. fertile shoot, B. Sporangiophore with sporangia,
C. T.S. of strobilus, D. L.S. of strobilus, E. T.S. of a spore showing various wall layers, F-G.
spores with elaters, H. Spore with free elaters.
 DEVELOPMENT OF SPORANGIUM
• Development of Sporangium:
• The mode of development of sporangium is eusporangiate, as it is not entirely
formed from a single initial. Superficial cells adjacent to the original initial may
also take part in the develop­ment of sporangium.
• Sporangia are initiated in single superficial cell around the rim of the young
sporangiophore. The periclinal division of the sporangium initial forms an inner and
an outer cell. The inner cell, by further divisions in various planes, gives rise to
sporogenous tissue.
• The outer cell, by periclinal and anticlinal divi­sions, gives rise to irregular tiers of
cells, the inner tiers of which may transform into sporoge­nous tissue and the outer
tiers become the future sporangial wall cells.
•
• The innermost layer of the sporangial wall differentiates as the tapetum. The sporogenous
cells separate from each other, round off and eventually transform into spore mother cell.
All but the two outermost wall layers disorganise to form periplasmodial fluid.
• However, not all of the sporogenous cells func­tion as spore mother cells. Many of them
degene­rates to form a multinucleate nourishing sub­stance for the spore mother cells.
Each spore mother cell undergoes meiotic division (reductional division) and produces
spore tetrad. All spores in a sporangium are of same size and shape i.e., homosporous.
• Structure of Mature Sporangium:
• The mature sporangium is an elongated sac­like structure, attached to the inner side of the
peltate disc of the sporangiophore. It is surrounded by a jacket layer which is com­posed
of two layers of cells. The inner layer is generally compressed and the cells of the outer
layer have helical thickenings which are involved in sporangial dehiscence.
•
• Dehiscence of Sporangium:
• At maturity, the strobilar axis elongates, as a result the sporangiophores become separated and
exposed. Then the sporangium splits open by a longitudinal line due to the differential hygroscopic
response of the wall cells.
• Spores:
• The spores are spherical and filled with densely packed chloroplasts. The spore wall is laminated and
shows four concentrate layers. The innermost is the delicate intine, followed by thick exine, the
middle cuticular layer and the outermost epispore or perispore. The intine (endospore) and exine
(exospore) are the true walls of the spore.
• The outer two layers i.e., cuti­cular layer and epispore are derived due to the disintegration of the
nonfunctional spore mother cells and tapetal cells. At maturity, the epispore (the outermost layer)
splits to produce four ribbon like bands or strips with flat spoon-like tips.
• These bands are free from the spore wall except for a common point of attachment and remain tightly
coiled around the spore wall until the sporangium is fully matured.
•
• These are called elaters. The elaters are hygroscopic in nature. The spores remain
moist at early stages of develop­ment, thus the elaters are spirally coiled round the
spore. The spores dry out at maturity and conse­quently the elaters become
uncoiled.
• These uncoiled elaters become entangled with the elaters of other spores. Through
these actions the elaters help in the dehiscence process and also the dispersal of
spores in large groups from the spo­rangium.
• The elaters of Equisetum are different from those of the bryophytes.
•
 GAMETOPHYTIC GENERATION
• Equisetum is a homosporous pteridophyte. The haploid spores germinate to form gameto­phyte. The
germination takes place immediately if the spores land on a suitable substratum. If the spores do not
germinate immediately, their via­bility decrease significantly.
• The spores swell up by absorbing water and shed their exine. The first division of the spore results in
two unequal cells: a small and a large cell. The smaller cell elongates and forms the first rhizoid. The
larger cell divides irregularly to produce the prothallus. The prevailing environmental conditions
deter­mine the size and shape of the prothallus. If a large number of spores are developed together
within a limited space, then the prothalli formed are of thin filamentous type. But a relatively thick
and cushion-shaped prothalli are formed from sparsely germinating spores. Mature gametophy­tic
plants may range in size from a few milli­meters up to 3 centimeters e.g., E. debile) in diameter.
• They are dorsiventral and consist of a basal non-chlorophyllous cushion-like portion from which a
number of erect chlorophyllous muticellular lobes develop upwards. Unicellular rhizoids are formed
from the basal cells of cushion.
• The prothallus bears sex organs and repro­duces by means of sexual method.
• 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.
Each antherozoid has about 120 flagella attached to the anterior end.
•
• 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. The archegonia are confined to cushion region in- between the aerial
lobes.
ANTHERIDIUM ARCHEGONIUM
 FERTILISATION
• Water is essential for fertilisation. 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 fertilised in a pro­thallus.
 EMBRYO
• 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 i.e., towards the neck of the archegoni­um) in polarity.
• 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 two 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 remai­ning cells of one quadrant of the epibasal hemi­sphere.
• One cell of the epibasal quadrant and a portion of the adjacent quadrant of the hypobasal region
contribute to the development of root. The first root develops from one of the epibasal quadrants and
a portion of the adjacent hypobasal quadrant. The shoot grows rapidly.
• 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 fertilised
LIFE CYCLE
 ECONOMIC IMPORTANCE
• Supports bone health
• Acts as a natural diuretic
• Promotes wound healing and nail health
• Promotes hair growth
• It was used traditionally to stop bleeding, heal ulcers and wounds, and treat
tuberculosis and kidney problems
• Cures diabetes.
• Boosts immune system
• Equisetum stems have been used in scouring (cleaning of utensils) and
polishing of metals.
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