Ephedra

Division: Gnetophyta

Class: Gnetopsida

Order: Ephedrales

Family: Ephedraceae

Genus: Ephedra

Distribution of Ephedra:

Ephedra (commonly known as joint pine, joint fir) is the only genus in family Ephedraceae and order
Ephedrales. It is represented by 65 species.

These species grow in dry climate over wide areas of the Northern hemisphere including North
America,Europe, North Africa, and South west and central Asia. Eight species of Ephedra are known from
India. Some of the common Indian species are E. intermedia, E. gerardiana, E. sexatilis, E.foliata etc.

Morphological Features of Ephedra:

The plant body is sporophytic and shows xerophytic characters. Mostly the plants are woody shrubs (Fig. 1
A), a very few species are lianas and some species grow into a small tree. E. compacta reaches 30 cm in
height E. triandra is a tree. Its height is several meters. Plant body can be differentiated into three parts –
root, stem and leaves.

1. Root:

There is a prominent underground tap root system. Later on the adventitious roots develop. Many root hairs
are present but there is no mycorrhiza.

2. Stem:

Like Equisetum, the stem is green, ribbed, branched, fluted and differentiated into nodes and internodes
(Fig.1B). It is distinctly jointed fir) (therefore, commonly known as jointed fir). It performs the function of
photosynthesis and may be called as phylloclade. The branches arise from the axillary buds and are,
therefore, in pairs of threes or fours according to the number of the scaly leaves at the nodes in different
species.

The branches are also green and differentiated into nodes and internodes. The branching starts early at the
cotyledonary stage. The apical meristem is having well marked tunica layer but the growth of internode is
independent due to the presence of the meristemetic zone at its base. This zone dries up at the end of each
growing season. It results in the brittleness and shedding of the branches. These branches are again replaced
in next season by new axillary branches.

3. Leaves:

Leaves are small scaly, present in pairs at the nodes and are arranged in opposite decussate manner. (Fig. 1
C, D). These leaves unite at the base to form a basal sheath. Each leaf contains two unbranched, parallel
veins. They are so minute that they are of no use i. e., unable to perform photosynthesis. The function of
photosynthesis is carried by green stem. In the axil of each leaf is present a bud for the branch. True foliage
leaves are absent.
 Reproduction in Ephedra:

Ephedra is heterosporous (produces two

types of spores: microspores and
macrospores) and dioecious (both these
types of spores are produced on two
different plants of the same species. E.
fuliata is monoecious. Microspores are
formed in male flowers while megaspores
are formed in female flowers.

These flowers are present in the form of

cone like compound strobili. Male flowers
are present in the form of male strobilus
while female flowers are present in the form
of female strobilus. Both male and female
strobili are compound i. e.,the cone axis
bears pairs of bracts which subtend either
microsproangiate or ovulate shoots.

Male Strobilus (Staminate Strobilus):

Male strobili arise in clusters from the nodes of the branches. Each strobilus is rounded, ovoid or spherical in
shape and arises in the axis of a scale leaf. Their number at the node depends upon the number of scale
leaves.

Each strobilus has a central axis which bears 2-12

pairs decussately arranged simple, broad and
cupped bracts. Lower most 1-2 pairs of bracts are
sterile. In the axil of each fertile bract arises a male
flower or staminate flower (Fig. 8 A-C). A male
strobilus with several male flowers can be
compared with an inflorescence.

Male flowers:

Each male flower has two lipped thin bractioles

(perianth) which encloses a stamen. Bracteoles are
united at the base. The flower has a short stalk
known as microsporangiophore and two, eight to
twelve microsporangia at its tip (Fig. 8 D).

Microsporangia are sessile and dehisce terminally.

Male flower is also called simple strobilus. A
compound male strobilus, therefore, consists of
many such strobili.

Structure of microsporangium:

Each microsporangium has 2-3 loculi and is often

called as synangium. Its wall is two layered
followed by a prominent tapetal layer enclosing a
sporangial sac having many pollen grains or
microspores (Fig. 8E).
Structure of pollen grain:

Pollen grain is the first cell of the male gametophyte. Each pollen grain is elliptical, uninucleate and has two
wall layers. The outer wall layer is thick and is called exine while the inner male layer is then and is called
intine (Fig. 10A, B).

Female Strobilus (Ovulate Strobilus) or Female Cone:

They usually arise in pairs at each node in the axil of

scale leaves. A female strobilus appears to be an elliptical
structure with a pointed apex (Fig. 11 A, B). It retains the
same compound structure as the male strobilus. It
consists of a short axis to which are attached three or four
pairs of decussate bracts.

In E. Americana these bracts are swollen and juicy (Fig.

11E). All the pairs of bracts are sterile except the
uppermost one which bears a pair of ovules in its axil
(Fig. 11C, D) and may be variously coloured. Out of the
pairs of the ovules only one survives and it takes up a false terminal position.

Female flower:

The female flower has short stalk and an ovule

(megasporangium)

Structure of ovule (megasporangium):

Longitudinal section of an ovule shows that it

consists of a mass of parenchymatous cells in the
centre. It is called nucellus. The nucellus is
surrounded by a two-layered envelope. These are
usually designated as outer and inner integuments.
The outer envelope is formed by four segments and
receives four bundles while the inner one is formed
of two segments and receives two bundles.

The lower half of the inner envelope is fused to the

nucellus but upper half is free and prolongs into a
long micropyle tube. By the time of pollination just
below the micropyle pollen chamber develops.
Pollen chamber in Ephedra is the deepest known
among the Gymnosperms. The floor of the pollen
chamber is formed by female gametophytic tissue
and not by the nucellus as in other gymnosperms.
(Fig. 12).

Development of Ovule:

Development of the ovule takes place in the form

of a small cellular protuberance. This protuberance
increases in size and becomes the nucellus. Soon
neighbouring cells of the base forms inner and
 outer integuments. Inner integument surrounds the nucellus except
the top where it form a small opening called micropyle.

A hypodermal archesporial cell differentiates in the nucellus. It

divides periclinally into outer parietal cell and inner megaspore
mother cell. The latter is pushed quite deep into the nucellar
tissue.

The megaspore mother cell divides meiotically to form four

hapliod megaspores. Generally the lowermost megaspore (towards
the chalazal end) remains functional. It enlarges and gives rise to
female gametophyte (first cell of the female gametophyte) and the
remaining upper three megaspores degenerate.

Gametophytic Phase:

The sporogenesis results in the formation of micro- and

megaspores representing the gametophytic stage. They undergo
gametogenesis to form the male and female gametophytes
respectively.

Development of male gametophyte before pollination:

It takes place in microsporangium. After the reduction division

spores tetrads are formed. The four cells of the tetrad separate and
develop into microspores. The microspore divides by a transverse
wall to form a small prothallial cell and a large outer cell is (Fig.
13 A). The outer cell again divides by a transverse wall and forms
a second prothallial cell and an antheridial cell. (Fig 13 B).

The antheridial cell divides to form a small generative cell and a

large tube cell (Fig. 13 C, D). The generative cell soon divides
into the nuclei of stalk cell and body cell. The nuclei of stalk cell
and body cell remain surrounded by a common mass of cytoplasm (Fig. 13 E, F). Pollens are shed at this
five celled stage.

Development of female gametophyte:

As mentioned earlier, the functional megaspore is the first cell of the female gametophyte. It enlarges and its
nucleus divides into two. These nuclei move towards the opposite pole and are separated by a large central
vacuole.

Later these two nuclei divide by free nuclear division to form as many as 256 nuclei. These nuclei are
arranged in a peripheral layer around the central vacuole. Later the central vacuole disappears and free
nuclei are evenly distributed throughout.

Centripetal wall formation (from periphery towards the centre) starts and thus a mass of cellular tissue is
formed. It is called female gametophyte or endosperm. Gradually the female gametophyte is differentiated
into two regions.

Micropylar region and antipodal region. Micropylar region consists of loosely arranged thin walled cells,
which later on give rise to archegonia. Antipodal region is further differentiated into lower storage zone and
basal haustorial zone. Storage zone comprises of bulk of endosperm. This zone consists of compactly
arranged cells which are full of starch and other food. The cells of the haustorial zone absorb the food
material from the nucellus.
Structure and development of archegonium:

Archegonia arise in the micropylar region. The

number of archegonia in Ephedra varies from 1-
3 but they are generally two in number. Any
superficial cell of female gametophyte towards
micropylar region acts as archegonial initial (Fig.
14A). It divides by a transverse division to form
outer primary neck cell or neck initial and inner
central cell (Fig. 14B). The neck cell undergoes
a number of divisions to form a long neck of 8 or
more tiers (minimum of 32 cells). It encloses no
neck canal.

The neck of archegonium of Ephedra is the

longest in the gymnosperms. The central cell
enlarges in size. Its nucleus divides into a ventral
canal nucleus and an egg nucleus but no wall is
laid down between the two.

As the archegonium reaches towards maturity,

the cells of neck usually merge with
surrounding gametophytic cells and become
undistinguishable from the surrounding cells of
female gametophyte. The cells adjacent to the
central cell divide transversely to form a
distinct jacket layer, which may be two or three
layer thick.

A mature archegonium consists of a long neck

and a central cell having a ventral canal
nucleus and egg nucleus (Fig. 13, 14).

Pollination:

The pollination is anemophilous i.e. it takes

place by wind. Pollen grains are carried by the
wind on the female strobilus. The cells of the
nucellus secrete pollination drop which comes
out through the micropylar canal. Pollen grains
to adhere to the pollination drop. Pollen grains
are sucked inside and come to lie in a deep
pollen chamber.

Development of male gametophyte after

pollination:

Pollen grains germinate in the pollen chamber.

The exine ruptures and intine comes out in the
form of pollen tube. The nucleus of the body
cell divides to form two male gametes (Fig. 16)
 Fertilization:

It occurs 10 hours after pollination. The pollen tube along with its four
nuclei (2 male nuclei, 1 stalk nucleus and 1 tube nucleus) gradually
penetrates the neck cell of the archegonium and discharges all the four
nuclei into the egg.

One male nucleus fuses with the egg nucleus forming the zygote (2x) or
oopsore. Khan (1941) observed in E.foliata that second male gamete
fuses with the ventral canal nucleus (double fertilization) but this diploid
nucleus does not develop into embryo Oospore is the first cell of the
sporophytic phase (Fig. 17).

Embryogeny:

More than one archegonium may be fertilized in an ovule, but only one oospore develops into embryo. The
zygote nucleus divides by three free nuclear divisions to form eight nuclei. These nuclei are irregularly
distributed in the cytoplasm of the archegonium.

Later wall-formation takes place and this structure is known

as proembryo. Each cell of inproembryo is capable to
develop into an independent embryo. Three to five of these
nuclei individually enclose in somewhat irregular walls and
become globular.

These are known as pro-embryonal cells, each of which

produces an independent embryo. In Ephedra, this type of
polyembrony without any cleavage, it unique among
gymnosperms. Because the polyembryony occurs without
any cleavage, it is known as embryo sac polyembryony.
Each proembryo grows into tubular structure called the
suspensor (Fig. 18A-C).

Tube nucleus of the proembryo divides into two. Both these

nuclei move into the tube. A wall separates these two
daughter nuclei and forms two cell(Fig. 18D). The cell
towards the micropylar and disintegrates while the cell
formed towards the chalazal end of the tube survives and is
called embryonal initial.

The tube grows

more and carries
the embryonal
initial deep into theprothallus tissue. This embryonal initial divides
into a proximal suspensor cell and a distalembryo cell. The embryo
cell divides and develops into the embryo proper which contains
two cotyledons (Fig. 18E-G).Although several embryos may
develop in a single ovule but only one survives and reaches at
maturity as seed.

Structure of Seed:

Longitudinal section of the seed shows that it consists of a

dicotyledonous embryo in the centre. This embryo is situated at the
tip of the elongated suspensor and remains embedded in the
endosperm (Fig. 19). The nucellus is consumed during the development of embryo and persists as a nucellar
cap at the micropylar end of the seed.

The seed is enclosed by the seed coat which consists of two

separate layers derived from the two envelopes. At the time of
maturity, the subtending bracts of the megasporangiate strobilus
become thick and fleshy and form an additional covering around
the seed e.g.,E. foliata.

Germination of the seed:

Seeds germinate without undergoing a period of rest if the

atmospheric conditions are favourable. The seed germination is
epigeal (Fig. 20A-G).

Economic Importance of Ephedra:

1. An alkaloid ephedrine is obtained from E. gerardiana, E.

intermedia, E. nebrodensis etc. It is used in preparation of
medicines that cure cough, bronchitis, asthma and hay fever.

2. Tincture of E. gerardiana
is also used as a cardiac and
circulatory stimulant.

3. Decoction of the stem and

roots is used to cure
rheumatism and syphilis
e.g.,E. antisyphilitica.

4. The juice of berry is used

to cure respiratory disorders.

5. Mormon tea is brewed

from the species of Ephedra
in south western United
States.

6. Some species are grown

as ornamentals.