Ephedra

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 pit
on their radial and tangential walls. In vessels, the bordered pits are also arranged in the same

way as tracheids (Fig. 5 B, C).

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).
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.

Biota
Biota, a common ornamental plant of Indian gardens, is a monotypic genus of Cupressaceae
and represented by only B. orientalis. It is often confused with and was once included under
Thuja as one of its species. On the basis of some morphological peculiarities, Endlicher
(1847) gave Biota a generic rank, and the same was supported by Pears (1937) on the basis of
his studies on wood anatomy, Bucholz (1948) on the basis of his studies on polyembryony
and Martin (1950) on the basis of his studies of embryology. Biota is an evergreen tree with a
deep penetrating tap root system and numerous green branches. Mature plants reach up to 10-
12 metres in height. The older branches remain covered with smooth brownish bark. Young
plants bear vertical lateral branches which become horizontal in the later conditions.
Persistent foliage leaves are present on the adult plants. The leaves are closely appressed (Fig.
11.51 A), acute and decussate. Each leaf remains fused with the stem along one-third of its
length.
Anatomically, young stem remains covered by a thickly circularized epidermis, a well-
developed cortex and a ring of vascular bundles which are conjoint, collateral, open and
endarch. Resin canals are present in the cortex. Medullary rays are broad and the xylem
consists of only tracheids. Companion cells are absent in the phloem Pith is small. Anatomy
of old stem exhibits secondary growth, due to which cortex becomes highly reduced and pith
gets completely obliterated. Only tracheids are present in the wood. Medullary rays are
uniseriate.

Internally; leaf shows xerophytic characters. Single-layered epidermis is thickly-circularized

and contains sunken stomata. Hypodermis is single- layered and made of lignified cells.
Mesophyll is differentiated into palisade and spongy parenchyma. Resin canals are present in
spongy parenchyma region. A single collateral vascular bundle is present in the centre of the
lamina. Phloem faces dorsal side while protoxylem of vascular bundle faces ventral side of
the lamina. Biota is monoecious, i.e. both the types of cones develop on the same plant. Male
and female cones (Fig. 11.51 B-C), however, develop usually on separate branchlets.

Female cone-bearing branches are usually curved downwards (Fig. 11.51 C). Each female
cone (measuring about 20mm in length and 15mm in diameter) contains 3-4 pairs of
decussate, fleshy ovuliferous scales, the tip of each of which is spiny and curved.

On the basal part of the ventral surface of each ovuliferous scale are present 1 to 3 ovules
(Fig. 11.51 D). Usually the ovule-bearing fertile scales in each cone are only three.
Uppermost and a few lower scales are stenle. As many as 10-12 ovules are present in the
female cone of Biota. Each ovule (Fig. 11.51 E) has a massive nucellus covered by an
integument.

In the upper part, the integument is free from the nucellus and forms a micropyle. Nucellus
contains one or two sporogenous cells which divide and form the sporogenous tissue. In the
centre of the sporogenous tissue develops a megaspore mother cell which divides
reductionally and form a row of three cells.

Lower two cells represent megaspores, of which only the chalazal megaspore remains
functional. The uppermost cell of the 3-celled stage is an undivided dyad cell.

Each male cone (measuring about 2.5 to 4mm in diameter) bears 4 to 8 microsporophyll’s
arranged in an opposite decussate manner Each microsporophyll (Fig. 11.5IF) is a shortly-
stalked body with a slightly broader base and an inwardly curved margin, and contains 3-5
microsporangia on its dorsal surface. Microsporophyll’s get separated exposing the
microsporangia in the mature male cones (Fig 11.51 B).
Each microsporangium, when young, is a round or elongated, shortly- stalked body
consisting of sprogenous cells surrounded by a three-layered wall (Fig. 11.51 G). The
innermost wall layer represents tapetum. The microsporangium development is of
eusporangiate type.

The cells of the sporogenous tissue represent microspore mother cells. They undergo
reduction division and form tetrads of haploid microspores (Fig. 11 51H). Several starch
grains and a large nucleus are present in each microspore. A thin exine and a thick intine
surround each microspore (Fig. 11.51I).

Female gametophyte develops from the functional megaspore (Fig. 11.52 A). The nucleus of
the megaspore divides repeatedly without any wall formation, and thus results in the
formation of several free nuclei (Fig. 11 52B,C) in the young and now enlarged gametophyte.
After the formation of about four thousand free nuclei, the wall formation starts in the
gametophyte.

Wall formation is centripetal and the entire gametophyte becomes cellular (Fig. 11 52D,E).
Mature gametophyte is long and slightly tapered at the chalazal end.

A large number of archegonia (15 to 30 but usually 22) develop near the micropylar end of
the female gametophyte in the form of an archegonial complex (Fig. 11.52F). A common
jacket layer surrounds the archegonial complex. Several of the cells of this jacket are bi-
nucleate.
Male gametophyte develops from the microspore or pollen grain (Fig. 11.51 I). In the initial
stages the number of starch grains increases, nucleus moves to one side and divides to form a
small antheridial cell and a large tube cell (Fig. 11.52G). Shedding of pollen takes place at
this two-celled stage. Its exine bursts as it happens to lie on the nucellus surface.
The intine comes out in the form of a small pollen tube which moves through the nucellar
tissue. The antheridial cell becomes free, moves down in the pollen tube and divides to form
a stalk nucleus and a body cell (Fig. 11 52H). Sometimes the pollen tube becomes branched.
The body cell divides and form two male cells. Stalk nucleus and tube nucleus ultimately dis-
organise.

Fertilization process starts after the entry of the pollen tube in the archegonial chamber. Neck
cells degenerate and a passage is formed for the entry of the male cells in the archegonium.
Fusion between the nucleus of one male cell and the egg nucleus results in the formation of a
diploid zygote.

Embryo development starts after the migration of the diploid zygotic nucleus towards the
archegonial base (Fig. 11.52 I). This diploid nucleus divides by free-nuclear divisions to form
4 and then 8 nuclei. Soon the wall formation takes place and 8 cells arranged in two tiers are
formed.

Lower tier of embryonal cells usually consists of 4 cells but sometimes of 6 or 5 cells also.
Upper tier usually consists of 4 cells but sometimes of only 2 or 3 cells.

The cells of upper tier divide to form an open tier and a middle pro-suspensor tier (Fig.
11.52K). Three distinct tiers of cells are thus formed: open tier, pro-suspensor tier and
embryonal tier (Fig. 11.52K). The cells of the pro-suspensor tier elongate, thus pushing the
embryonal tier into the gametophyte. The cells of the embryonal tier divide and form primary
suspensor cells and embryonal cells (Fig. 11 52L).

Primary suspensor cells elongate, and at the tip of each of these cells is present a small
embryonal cell. After a period of rest, during which primary suspensor cells elongate too
much, each embryonal cell divides by two vertical divisions at right angles to one another.
Several irregular divisions in this dividing embryo result in the formation of a globular mass.

Sometimes the secondary suspensors are also formed. All embryos abort gradually except the
one which is situated deepest in the gametophytic tissue. A root tip, stem tip and two
cotyledons are present in a mature embryo (Fig. 11.52M).

Hypocotyl is formed by the elongation of the region between root tip and stem tip. A mature
seed remains surrounded by a 20-25 cells thick seed coat. Seeds show almost no dormancy,
and the germination is epigeal.
 Gnetum
1. Distribution of Gnetum:
Gnetum, represented by about 40 species is confined to the tropical and humid regions of the
world. Nearly all species, except G. microcarpum, occur below an altitude of 1500 metres.
Five species (Gnetum contractum, G. gnemon, G. montanum, G. ula and G. latifolium) have
been reported from India. Gnetum ula is the most commonly occurring species of India.

2. Habit of Gnetum:
Majority of the Gnetum species are climbers except a few shrubs and trees. G. trinerve is
apparently parasitic. Two types of branches are present on the main stem of the plant, i.e.
branches of limited growth and branches of unlimited growth. Each branch contains nodes
and intemodes Stem of several species of Gnetum is articulated

In climbing species the branches of limited growth or short shoots are generally un-branched
and bear the foliage leaves. The leaves (9-10) are arranged in decussate pairs (Fig. 13.2).
They often lie in one plane giving the appearance of a pinnate leaf to the branch. The leaves
are large and oval with entire margin and reticulate venation as also seen in dicotyledons.
Some scaly leaves are also present.

3. Anatomy of Gnetum:
(i) Root:
Young root (Fig. 13.3) has several layers of starch-filled parenchymatous cortex, the cells of
which are large and polygonal in outline. An endodermal layer is distinguishable. Casparian
strips are seen in the cells of the endodermis. The endodermis follows 4-6 layered pericycle.
Roots are diarch and exarch. Small amount of primary xylem, visible in young roots,
becomes indistinguishable after secondary growth.

The secondary growth is of normal type. A continuous zone of wood is present in the old
roots (Fig. 13.4). It consists of tracheids, vessels and xylem parenchyma. The tracheids have
uniseriate bordered pits along with bars of Sanio.

Vessels have simple or small multiseriate bordered pits. Some of the xylem elements have
starch grains. Bars of Sanio are generally absent in the vessels. Phloem consists of sieve cells
and phloem parenchyma.

(ii) Young Stem:

The young stem in transverse section is roughly circular in outline, and resembles with a
typical dicotyledonous stem. It remains surrounded by a single-layered epidermis, which is
thickly circularized and consists of rectangular cells. Some of the epidermal cells show
papillate outgrowths. Sunken stomata are present.

The cortex consists of outer 5-7 cells thick chlorenchymatous region, middle few-cells thick
parenchymatous region and inner 2-4 cells thick sclerenchymatous region. Endodermis and
pericycle regions are not very clearly distinguishable. Several conjoint, collateral, open and
endarch vascular bundles are arranged in a ring (Fig. 13.5) in the young stem . Xylem consists
of tracheitis and vessels. Presence of vessels is an angiospermic character. Protoxylem
elements are spiral or annular while the metaxylem shows bordered pits which are circular in
outline. The phloem consists of sieve cells and phloem parenchyma. An extensive pith,
consisting of polygonal, parenchymatous cells, is present in the centre of the young stem.

(iii) Old Stem:

Old stems in Gnetum show secondary growth. In G. gnemon the secondary growth is normal,
as seen also in the dicotyledons. But in majority of the species (e.g., G. ula, G. africanum,
etc.) the anamolous secondary growth is present.

The primary cambium is ephemeral, i.e., short-lived. The secondary cambium in different
parts of cortex develops in the form of successive rings, one after the other (Fig. 13.6). The
first cambium cuts off secondary xylem towards inside and secondary phloem towards
outside. This cambium ceases to function after some time.

Another cambium gets differentiated along the outermost secondary phloem region, and the
same process is repeated. In the later stages, more secondary xylem is produced on one side
and less on the other side, and thus the eccentric rings of xylem and phloem are formed in the
wood. This type of eccentric wood is the characteristic feature of angiospermic lianes. The
periderm is thin and develops from the outer cortex. It also possesses lenticels. The cortex
also contains chlorenchvmatous and parenchymatous tissues along with many sclereids.
In old stems the secondary wood consists of tracheids and vessels. Tracheids contain
bordered pits on their radial walls while vessels contain simple pits. Transitional stages (Fig.
13.7), containing one to many perforations in the terminal part of the vessels, are also seen
commonly.
In tangential longitudinal section (T.L.S) of the stem (Fig. 13.8), the wood xylem and
medullary rays are visible. Bordered pits on both the radial and tangential walls are present.
Medullary rays are either uniseriate or multiseriate and consist of polygonal parenchymatous
cells. They are boat-shaped (Fig. 13.8) and their breadth varies from 2 to many cells. Sieve
cells of the phloem contain oblique and perforated sieve plates.

(iv) Leaf:
Internally, Gnetum leaves also resemble with a dicot leaf. It is bounded by a layer of thickly
circularized epidermis on both the surfaces. Stomata are distributed all over the lower surface
except on the veins. The mesophyll is differentiated generally into a single-layered palisade
and a well-developed spongy parenchyma.

The latter consists of many loosely-packed cells. Many stellately branched sclereids are
present near the lower epidermis in the spongy parenchyma. Many stone cells and latex tubes
are present in the midrib region of the leaf.

Several vascular bundles in the form of an arch or curve are present in the prominent midrib
region (Fig. 13.9). A ring of thick-walled stone cells is present just outside the phloem. Each
vascular bundle is conjoint and collateral.
The xylem of each vascular bundle faces towards the upper surface while the phloem faces
towards the lower surface. The xylem consists of tracheids, vessels and xylem parenchyma
while the phloem consists of sieve cells and phloem parenchyma.

4. Reproduction of Gnetum:
Gnetum is dioecious. The reproductive organs are organised into well-developed cones or
strobili. These cones are organised into inflorescences, generally of panicle type. Sometimes
the cones are terminal in position. A cone consists of a cone axis, at the base of which are
present two opposite and connate bracts. Nodes and internodes are present in the cone axis.
Whorls of circular bracts are present on the nodes. These are arranged one above the other to
form cupulas or collars (Fig. 13.10). Flowers are present in these collars. Upper few collars
may be reduced and are sterile in nature in G. gnemon.
Male Cone and Male Flower:
The male flowers are arranged in definite rings above each collar on the nodes of the axis of
male cone. The number of rings varies between 3-6. The male flowers in the rings are
arranged alternately. There is a ring of abortive ovules or imperfect female flowers above the
rings of male flowers. Each male flower contains two coherent bracts which form the
perianth (Fig. 13.11). Two unilocular anthers remain attached on a short stalk enclosed within
the perianth. At maturity, when the anthers are ready for dehiscence, the stalk elongates and
the anthers come out of the perianth sheath. In Gnetum gnemon a few (2-3) flowers are
sometimes seen fusing each other (Fig. 13.12).
Development of Male Flower (Figs. 13.13, 13.14):
In very young cones, certain cells below each collar become meristematic. They divide
repeatedly and form a small hump-like outgrowth. Certain cells on the upper side of this
annular outgrowth start to differentiate into the initials of the ovules. They develop into
abortive ovules which form the uppermost ring. The cells of the lower side of this annular
outgrowth form the primordium of male flower.
A central cushion of cells develops by the repeated divisions in the male flower primordium.
This cushion gets surrounded by a circular sheath called perianth. The sheath-like perianth
encloses the central cushion-like mass only partially. With the development of a depression
or notch in the central mass two lobes differentiate and later on develop into two anther lobes.
With the help of many divisions the basal portion of this central mass of cells starts to
differentiate into a stalk. This stalk elongates and pushes the anther lobes towards the outer
side. Each anther lobe remains surrounded by an epidermal layer and a few wall layers which
enclose a microsporangium. The innermost wall layer enclosing the sporogenous tissue is
known as tapetum. The sporogenous cells become loose, contract, round up and change into
the spore mother cells. In the process of microspore formation the tapetum and two wall
layers are used for the developing microspores. The spore mother cells undergo meiosis and
ultimately the spore tetrads are formed. The characteristic radial thickenings develop in the
epidermal cells. They help in the dehiscence of microsporangium. The microspores are
ornamented.

Female Cone:
The female cones resemble with the male cones except in some definite aspects. A single ring
of 4-10 female flowers or ovules is present just above each collar (Fig. 13.15). Only a few of
the ovules develop into mature seeds (Fig. 13.15B).

In the young condition, there is hardly any external difference between female and male
cones. All the ovules are of the same size when young but later on a few of them enlarge and
develop into mature seeds. All the ovules never mature into seeds.

Ovule or Female Flower:

Each ovule (Fig. 13 16) consists of a nucellus surrounded of three envelopes. The nucellus
consists of central mass of cells. The inner envelope elongates beyond the middle envelope to
form the micropylar tube or style. The nucellus contains the female gametophyte. There is no
nucellar beak in the ovule of Gnetum. Stomata, sclereids and laticiferous cells are present in
the two outer envelopes. Madhulata (1960) observed the formation of a circular rim from the
outer epidermis of the inner integument in G. gnemon. Thoday (1921), however, observed the
formation of a second such rim at a higher level. The ovules in G. ula are stalked.

Four to ten ovular primordia differentiate on the annular meristematic ring. This ring
develops below each collar of the female cone in the same manner as that of the male cone.
The ovular primordium divides and re-divides several times to form a mass of cells. All the
three envelopes of the female flower develop around this mass of cells The innermost third
envelope remains fused with the nucellus at the base while its upper portion remains free and
form the long micropylar tube or ‘style’. In the young conditions, an outer epidermal layer is
distinguishable in the nucellus. Two to four archesporial cells develop below the epidermis at
a later stage. The archesporial cells divide periclinally to form outer primary’ parietal cells
and inner sporogenous cells. The primary parietal cells and the epidermal layer divide
periclinally and anticlinally several times resulting into a massive nucellus.
The sporogenous cells divide and re-divide to form megaspore mother cells which remain
arranged in linear rows. All the megaspore mother cells may divide reductionally and form
tetrasporic embryo-sacs but ultimately all, except one, degenerate.

As many as 256 (Gnetum gnemon) to 1500 (G. ula) free-nuclei are formed in the female
gametophyte leaving a vacuole in the centre (Fig. 13.18). The female gametophyte is
tetrasporic in development. It is broader towards the micropylar end and it tapers towards the
chalazal end.

The nuclei near the chalazal end get surrounded by cell walls while those towards micropylar
end remain free. Gametophyte is thus partly cellular and partly-nuclear. The archegonia are
absent in Gnetum. Certain nuclei near the micropylar end start to function as egg nuclei.
According to Swamy (1973) the only nucleus in a uninucleate cell or one of the nuclei in a
multinucleate cell enlarges and functions as the egg in G. ula. The nucellar beak is absent in
Gnetum.

The megaspore mother cell divides reductionally and forms four free haploid nuclei in the
mother cell. Megaspore tetrads are never formed in Gnetum.
Microsporangium and Micro-Sporogenesis:
Development of the microsporangium (Fig. 13.19) can be studied only in young anthers. Two
archesporial cells are distinguished below the epidermal layer (Fig. 13.19A). Archesporial
cells divide and re-divide to form many-celled archesporium (Fig. 13.19B). The outermost
layer of the archesporium divide periclinally to form an outer layer of parietal cells and inner
layers of sporogenous cells (Fig. 13 .19C).

The parietal cells form the wall layers and tapetal layer by periclinal divisions (Fig. 13.19D).
The sporogenous cells develop into microspore mother cells by some irregular divisions.
Tapetal cells later on become bi-nucleate (Fig. 13.19D, E). Microspore mother cells divide
reductionally to form haploid microspores.

The microspores may be arranged in isobilateral, decussate or tetrahedral manner in their

earlier stages. Side by side the wall cells and the tapetal cells degenerate and ultimately dis-
organise. The epidermal cells become thick, cutinized and radially elongated.

Many fibrous thickenings also develop in these cells (Fig. 13.19H). Small globular structures
are present on the inner surface of the epidermis in Gnetum ula and G. gnemon. Anthers
dehisce along a double row of small cells which extends from the tip towards the base.

Male Gametophyte:
Pollen grains or microspores are roughly spherical in outline. They are uninucleate and
remain surrounded by a thick and spiny exine and thin intine. Mature pollen grains are shed
at three-nucleate stage. These include prothallial nucleus, tube nucleus and generative
nucleus (Fig. 13.20, Upper) in Gnetum Africanism and G. gnemon according to Pearson
(1912, 1914). This three-nucleate stage is reached by first dividing the microspore nucleus
mitotically into two and then one of them again gets divided. Further development is affected
only in the pollen chamber. The intine comes out by rupturing the exine and forms a pollen
tube.

The tube nucleus migrates into the pollen tube. The generative nucleus also adopts the same
course and divides into two unequal male gametes in the tube. Prothallial nucleus does not
enter the pollen tube.

Thompson (1916) opined that the prothallial cell does not form at all in the male gametophyte
(Fig. 13.20, Middle). The microspore nucleus divides into a tube nucleus and a generative
cell. The latter divides into a stalk cell and body cell. The tube nucleus and body cell enter in
the pollen tube where the body cell divides into two equal male gametes.

According to Negi and Madhulata (1957) the microspore nucleus in Gnetum gnemon and G.
ula divides into a small lenticular cell and a large cell (Fig. 13.20, Lower). The lenticular cell
does not take part in the further development and ultimately disappears.
The other large nucleus divides into a tube nucleus and a generative cell, both of which pass
into the tube. The generative cell divides into two equal male gametes in the tube. A stalk cell
is never formed in these species.

Pollination:
Wind helps in carrying the pollen grains up to the micropylar tube of the ovule. The
micropylar tube secretes a drop of fluid in which certain pollen grains get entangled and
reach up to the pollen chamber. The nucellus cells below the pollen chamber are full of
starch.

Fertilization:
The fertilization in Gnetum has been studied only by a few workers. Vasil (1959) studied this
phenomenon in G. ula. At the time of fertilization, the pollen tube pierces through the
membrane of the female gametophyte just near to a group of densely cytoplasmic cells. The
tip of pollen tube bursts and the male cells are released. One of the male cells enters the egg
cell.

The male and female nuclei, after lying side by side for some time, fuse with each other and
form the zygote. According to Swamy (1973), the only identifying features of the zygote are
its spherical shape and dense cytoplasm. Both the male cells of a pollen tube may remain
functional if two eggs are present close to the pollen tube.

Endosperm:
In all gymnosperms, except Gnetum, a cellular endosperm (Fig. 13.21) develops before
fertilization. In Gnetum, the cell formation, although starts before fertilization, a part of the
gametophyte remains free-nuclear at the time of fertilization.

After fertilization the wall formation in the female gametophyte starts in such a way that the
cytoplasm gets divided into many compartments. Each of these compartments contains many
nuclei (Fig. 13.21C).

All the nuclei of one compartment fuse and form a single nucleus. The wall formation starts
from the base and proceeds upwards. The wall formation varies greatly in Gnetum. Only the
lower portion of the gametophyte may become cellular leaving the remaining upper portion
free-nuclear. Sometimes the entire gametophyte may become cellular.

In some cases the upper portion may become cellular instead of the lower portion. Sometimes
only the middle portion may become cellular and in still other cases there may not be any
wall formation at all. The characteristic triple fusion of the angiosperms is, however, absent
in Gnetum.
The Embryo:
The embryo development in several species of Gnetum has been studied by many different
workers including Lotsy (1899), Coulter (1908) and Thompson (1916), but the details put
forward by these wokers are highly variable.

Maheshwari and Vasil (1961) have stated that in all the angiosperms the first division of the
zygote is accompanied by a wall formation but in all gymnosperms, except Sequoia
sempervirens, these are free-nuclear divisions in the zygote. Gnetum in this respect forms a
link in between gymnosperms and angiosperms by showing both free-nuclear divisions as
well as cell divisions. Thompson (1916) opined that a two-celled pro-embryo is formed (Fig.
13.22 A). From each of these two cells develops a tube called suspensor (Fig. 13.22B). Now
the nucleus divides and one of the two nuclei undergoes free-nuclear divisions forming four
nuclei. The embryo gets organised by these four nuclei (Fig. 13.22C, D). There is no division
in the other larger nucleus..

Madhulata (1960) has worked on the zygote development in Gnetum gnemon. According to
her 2-4 or sometimes up to 12 zygotes may develop in a gametophyte, of which normally one
remains functional. From the zygote develops generally one or sometimes 2-3 small tubular
outgrowths. Only one of these tubes receives the nucleus and survives while the remaining
tubes disintegrate and soon die. The surviving outgrowth elongates, becomes branched and
grows into different directions through the intercellular spaces of the endosperm. All the
primary suspensor tubes usually remain coiled round each other. A small cell is cut off at the
tip of the primary suspensor tube in Gnetum gnemon. It soon divides first transversely and
then longitudinally resulting into four cells. Now irregular divisions take place forming a
group of cells. Some of these cells divide and elongate to form secondary suspensor (Fig.
13.23). The remaining cells at the tip form the embryonal mass.

In Gnetum ula a small cell is cut off at the tip of the tube called peculiar cell. This peculiar
cell soon divides and forms a group of cells. The secondary suspensor and embryonal mass
are differentiated (Fig. 13.24) from this group of cells. By this time, the wall of the tube starts
to become thick.
What so ever may be the pattern of formation of the embryonal mass and secondary
suspensor, the cells of the former are small, compact, dense in cytoplasm and develop into
embryo-proper while that of the latter (i. e. secondary suspensor) are thin-walled, uninucleate
and highly vacuolated. The primary and secondary suspensors help in pushing the embryo
into the endosperm. Soon a stem tip with two lateral cotyledons form in the tip region of the
embryonal mass. On the opposite side develop the root tip with a root cap. A feeder develops
after the formation of stem and root tips (Fig. 13.25). The feeder is a protuberance-like
structure present in between root and stem tips. Thus, the stem tip, two cotyledons, feeder,
root tip and root cap

are the parts of a mature embryo.

Seed:
Gnetum seeds (Fig. 13.26) are oval to elongated in shape and green to red in colour. It
remains surrounded by a three-layered envelope which encloses the embryo and the
endosperm. Outer envelope is fleshy, and consists of parenchymatous cells. It imparts colour
to the seed.

The middle envelope is hard, protective and made up to three layers, i.e., outer layer of
parenchymatous cells, middle of palisade cells and innermost fibrous region. The inner
envelope is parenchymatous. Branched vascular bundles traverse through all the three
envelopes.
Germination of Seed:
Germination is of epigeal type (Fig. 13.27). The cotyledons are pushed out of the seed. The
hypocotyl elongates, and this brings the cotyledons out of the soil. The first green leaves of
the plant are formed by the cotyledons. The first pair of foliage leaves is produced by the
development of plumule. A persistent feeder is present up to a very late stage in the seed.

5. Relationships of Gnetum:
Gnetum and Other Gymnosperms:
Gnetum shows several resemblances with gymnosperms and has, therefore, been finally
included under this group.

Some of the characteristics common in both Gnetum and

other gymnosperms are under mentioned:
1. Wood having tracheids with bordered pits.

2. No sieve tubes and companion cells are present.

3. Presence of naked ovules.

4. Absence of fruit formation because of the absence of ovary.

5. Anemophilous type of pollination.

6. Development of prothallial cell.

7. Cleavage polyembryony.

8. Resemblance of the vascular supply of the peduncle of the cone of Cycadeoidea wielandii
with that of a single flower of Gnetum.

9. Resemblance of the structure of basal part of the ovule in Gnetum and Bennettites.