Name : salwa khan

Teacher name : maam shamal habib

Semester : 3RD

DEPARTMENT : CHEMISTRY

Ephedra: Distribution, Features and Reproduction

Distribution of Ephedra:
Ephedra (commonly known as joint pine, joint fir, Mormon tea or Brigham tea) is the only genus in
family Ephedraceae and order Ephedrales. It is represented by 50 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. These species are distributed in dry parts of Punjab, Haryana, Rajasthan and parts of Kashmir to
Sikkim.

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.
Internal Structures of Ephedra:
1. Stem:
The stem is ribbed; so, in tranverse section stem shows ridges and grooves (Fig. 2A).

A T.S. of stem at node shows the following structures (Fig. 2A, B):
a. Epidermis:
It is the outermost layer of thick walled cells, covered with a thick layer of cuticle. Sunken stomata
are present on the slopes of the ridges in the circular pits.

b. Hypodermis:
It is present just below the ridges. (Fig 2B). It is made up of sclerenchymatous cells and provides
mechanical strength to the plant.

c. Cortex:
In is present between the thick walled sclerenchyma and vascular cylinder. It can be differentiated
into outer and inner cortex. The outer cortex contains 2-3 layers of radially elongated palisade tissue
and inner cortex consists of 2-3 layers of spongy parenchyma.

The cells of outer and inner cortex are loosely arranged with large intercellular spaces and are
provided with chlorophyll to perform the function of photosynthesis. A few patches of
scleranchymatous cells may also occur in the cortex to provide mechanical support to the young
axis.

d. Endodermis:
It is the innermost layer of cortex. It is not easily distinguishable from the cortical cells.

e. Pericycle:
It is present below the endodermis. It is single layered and ill defined.

f. Vascular Cylinder:
It is endarch, siphonostele and consists of many vascular bundles arranged in a ring. Vascular
bundles are conjointed, collateral, open and endarch. The number of primary vascular strands is
generally eight, out of which four small represent the foliar traces while the other large four are
stem bundles.

Foliar traces run upto the node. Xylem consists of tracheids, vessels and xylem parenchyma. Due to
the presence of the vessels the Ephedra resembles angiosperms. The phloem consists of sieve cells,
phloem parenchyma and albuminous cells. Phloem and xylem are separated by a narrow strip of
cambium.

g. Medullary rays:
Broad, parenchymatous medullary rays are present in between the vascular bundles. Medullary rays
connect the pith with cortex.

It is present in the centre. It is made up off thin walled parenchymatous cells. Near the node its cells
become strongly lignified forming a peridermal diaphragm which accounts for the rapid separation
of the branches in the region (Fig. 3).

Secondary growth:

The secondary growth takes place by the activity of intrafascicular cambium and interfascicular
cambium. After forming a complete ring of cambium, the cambial cells cut of secondary phloem on
the outer side and secondary xylem towards the inner side. (Fig. 4)
Due to formation of the secondary tissues, primary phloem is crushed and the primary xylem is
pushed towards the inner side at the base of the secondary xylem. In addition to vascular tissue
cambium also forms medullary rays (secondary). These rays are long, broad (multiseriate) and
traverse between secondary xylem and secondary phloem.

Radial Longitudinal Section (R.L.S.):

In R.L.S. xylem tracheids, vessels and medullary rays are clearly visible. Medullary rays are cut
lengthwise and their length and height are revealed (Fig. 5A). Each medullary ray is composed of
irregularly dispersed ray cells and ray tracheids. Tracheids possess bordered pits on their radial and
tangential walls. In vessels, the bordered pits are also arranged in the same way as tracheids (Fig. 5
B, C).

Tangential Longitudinal Section (T.L.S.) of wood:

Like R.L.S. in T.L.S. also, the xylem, tracheids, vessels and medullary rays are clearly visible but they
are cut transversely here. (Fig. 6). Bordered pits and simple pits are seen on the radial and tangential
walls. The medullary rays are elongated and on their tangential walls are present simple pits.
2. Leaf:
The transverse section of scaly leaf is oval in shape and can be differentiated into epidermis,
mesophyll tissue and vascular tissue.

a. Epidermis:
It is outer most single layer of thick walled elongated cells. The cells are covered with thick cuticle.
Sunken stomata are present (Fig. 7).

b. Mesophyll tissue:
Two or three layers of palisade tissue are present inner to epidermis. The cells are filled with
chloroplast and large intercellular spaces are present between them. In the centre of the leaf
parenchymatous tissue is present.

c. Vascular tissue:
Two vascular bundles are embedded in the parenchymatous tissue. The vascular bundles are
collateral and closed. Xylem is present towards the upper side.

3. Root:
The transverse section of root shows single layer epiblema, outer cortex (composed of
collenchymatous cells), inner cortex (composed of parenchymatous cells) endodermis and pericycle.
Vascular bundles are radial and exarch. The root may be diarch or triarch.
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).

Development of microsporangium:
The development of microsporangium is eusporangiate. Microspangia arise at the tip of
microsporangiophore. The microsporangiophore arises as small protuberance in the axil of the fertile
bract of male strobilus. The apex of microsporangiophore becomes lobed after growing for some
time.

Each lobe represents a sporangium. Few hypodermal cells in each lobe enlarge in size. These cells
consist large nuclei, denser cytoplasts and are known as archesporial cells. These cells divide
periclinally into outer primary wall cells or primary parietal cells and primary sporogenous cells (Fig.
9A).

Primary sporogenous cells further divide by two periclinal divisions to differentiate middle wall layer,
inner tapetal layer and sporogenous cells. The primary wall cells function directly as the outer wall of
the sporangium.

However, according to some workers, the primary wall cells divide periclinally to form three layered
thick wall. The sporogenous cells divide further to form large number of microspore mother cells.
Each microspore mother cell divides by meiosis to form four haploid microspores arranged in a
linear tetrad.

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).

Ephedra: Anatomy and Reproductive Structures:

Anatomy of Different Parts of Ephedra:

Cut thin transverse sections of internodes of young and old stems, stain them in
safranin-fast green combination, mount in glycerine and study.

Stem: T.S. Young Internode:

1. The outline shows many ridges and grooves (Figs. 50, 51).
2. Outermost layer in epidermis with a thick layer of cuticle. Continuity of the
epidermis is broken by many sunken stomata present in the grooves.

3. Below the ridges are present the patches of sclerenchyma.

4. Cortex is separated into palisade and spongy parenchyma. Some sclerenchyma

patches are also irregularly distributed in the cortex.

5. Vascular bundles are arranged in a ring and the stele is ectophloic siphonostele.

6. Stele is bounded by a layer of endodermis and unilayered pericycle.

7. Each vascular bundle is conjoint, collateral, open and endarch (Fig. 51).

8. In each vascular bundle phloem is present on the outer side and xylem on the inner
side and both are separated by cambium.

9. Parenchymatous pith is present at the centre of the stem.

Stem: T.S. Old Internode:

1. It is wavy in outline with ridges and grooves and remains surrounded by a single-
layered, heavily cuticularized epidermis. The cortex is also differentiated into palisade
and spongy parenchyma (Figs. 52,53).
2. In older stems, the periderm replaces these primary cortical structures.

3. Secondary tissue is externally bounded by a layer of sclerotic cells or stone cells.

4. Cambium cuts secondary phloem towards outer side and secondary xylem towards
inner side.

5. Secondary tissue crushes and pushes the primary phloem towards outer side and
primary xylem towards inner side.

6. Thin-walled spring wood and thick-walled autumn wood are present in the secondary
xylem in the form of regular alternate rings.

7. Sieve tubes and phloem parenchyma are present in phloem.

8. Presence of vessels is the characteristic feature of the wood of Ephedra.

9. Primary medullary rays connect primary phloem and primary xylem while the
secondary medullary rays connect secondary phloem and secondary xylem. Medullary
rays are uniseriate in young stem while in the old stem these are multiseriate.

10. Resin canals are absent.

11. Parenchymatous pith is present in the centre.

R.L.S. Wood:
1. Secondary xylem and medullary rays are present (Fig. 54).

2. Tracheids form the major part of secondary xylem.

3. Bordered pits are present on the radial walls of tracheids. They are usually circular
and never polygonal in shape.

4. Bordered pits are either arranged in 2 or 3 rows or they are scattered irregularly.
Below the bordered pits are also present Bars of Sanio.

5. A few vessels with perforated end walls are also present.

6. Medullary rays are cut lengthwise showing their length and height. They are
uniseriate or multiseriate. The height of medullary rays reaches upto 50 cells.

7. In the region of secondary xylem, the medullary rays are made of ray cells and ray
tracheids. Bordered pits are present in the tangential walls of ray cells. Ray tracheids
are thick-walled and contain bordered pits on their radial and tangential walls.

8. Starch cells surrounded by albuminous cells on both sides constitute the medullary
ray in the region of phloem.

T.L.S. Wood:
1. Tracheids, vessels and medullary rays (Fig. 55) are cut transversely in T.L.S.
2. Bordered pits are observed in the surface view.

3. A small disc torus is present in each bordered pit.

4. The height and breadth of medullary rays can be observed in this plane because they
are cut transversely.

5. Simple pits are present on the tangential walls of medullary rays.

Reproductive Parts Ephedra:

1. Ephedra is heterosporous, i.e., two types of spores (microspores and megaspores) are
present.

2. Microspores are present in male flowers while the megaspores in female flowers.

3. These flowers are present in the form of cone-like compound strobili.

4. Male flowers are present in the form of male strobilus while many female flowers
form the female strobilus.

5. Each strobilus arises on the node in the axil of a scaly leaf.

6. Plant is generally dioecious.

Male Strobilus:
1. Male strobili develop in the axil of scaly leaves on the nodes (Fig. 56).
2. Each strobilus is round or ovoid in shape.

3. In the centre is present the strobilus axis or cone axis.

4. On the strobilus axis are arranged 2 to 12 pairs of bracts in opposite decussate

manner (Figs. 57, 58).

5. All the bracts are fertile except a few on the lower side.
6. A single staminate flower or male flower arises in the axil of each bract.

7. Each male flower consists of two bracteoles and a stamen (Fig. 59).

8. Each stamen is a stalked structure with 2 to 4 anthers or microsporangia at the top.

9. Each anther or microsporangium is bilocular or trilocular, and each locule is

surrounded by a double-layered wall and an innermost layer of tapetum (Fig. 60).

10. May pollen grains or microspores are present in each locule. Each pollen grain is a
uninucleate structure surrounded by a thin intine and thick outer layer of exine.

Female Strobilus:
1. Similar to the male strobilus, the female strobilus also develops in the axil of the leaf
on the node.

2. It is sessile and smaller than male strobilus.

3. Two to four pairs of bracts are arranged in opposite decussate manner on the
strobilus axis (Figs. 61, 62).
4. Except the uppermost pair of bracts, all are sterile.

5. Two ovules are present in the axil of uppermost pair of bracts (Fig. 62B), out of which
generally only one ovule survives.

Ovule:
1. It is covered by a cup-shaped outer integument and an inner integument.

2. Outer integument is attached at the basal part of the ovule (Fig.62D).

3. Inner integument protrudes out of the bracts and outer integument in the form of a
long tubular structure.

4. Outer integument is also known as perianth or involucre.

5. A long micropyle is formed by inner integument (Fig. 62D).

6. Integuments enclose the nucellus.

7. Near the micropyle develops a small pollen chamber in the nucellus.

8. Pollen chamber in Ephedra is deepest known among Gymnosperms.

9. Female gametophyte is present in the centre.

10. Archegonia are present in the female gametophyte near the micropylar end.

11. Haustorial region bearing some haustorial processes is present at the end opposite to
the micropylar end.

L.S. Seed:
1. Bracts and outer integument surround the entire seed (Fig. 63).

2. Bracts are thick, fleshy and form additional envelope.

3. The outer integument is a very thick structure.

4. The inner integument persists only at the micropylar end.

5. Nucellus is present in the form of a disorganised and shrivelled layer.

6. Embryo remains surrounded by endosperm.

7. Two large cotyledons are present in the embryo.

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.

The end