Essay on the Life Cycle of Cycas | Class

Cycadopsida | Gymnosperms | Botany

https://www.biologydiscussion.com/essay/gymnosperms/essay-on-the-life-cycle-of-cycas-class-
cycadopsida-gymnosperms-botany/76688

Here is an essay on the ‘Life Cycle of Cycas’ for class 11 and 12. Find paragraphs, long
and short essays on the ‘Life Cycle of Cycas’ especially written for school and college
students.
Essay # 1. Sporophyte of Cycas:
The Cycas plant looks like a palm tree with a columnar aerial trunk and a crown of
pinnately compound leaves (Fig. 1.12). Generally the stem is unbranched, but older
trees sometimes exhibit branching. The branch is developed from the adventitious
bud called ‘bulbil’ which arises from the leaf base. Thus the bulbil helps in vegetative
propagation of the plant. The stem is covered by an armour of alternating bands of
large and small rhomboidal leaf bases.
These leaf bases are persistent of which the large bases belong to foliage leaves, while
the small ones belong either to scale leaves (in male plants) or to sporophyll (in
female plants). Cycas exhibits leaf dimorphism in possessing large green foliage
leaves and small scale leaves or cataphylls. The Primary tap root system is replaced by
a number of” strong, branched adventitious roots arising from the stem.
Essay # 2. Reproduction of Cycas:
Cycas reproduces both by vegetative and sexual methods:
I. Vegetative Reproduction:
Cycas reproduces vegetatively by means of bulbils or adventitious buds. These bulbils
develop from the parenchymatous cells of the cortex in the base of the stem at the
crevices between persistent leaf bases. Initially bulbils are covered only with scale
leaves, but a few foliage leaves develop with further growth.
Bulbil arising on male plant gives rise to male plant, if it forms on female plant it
produces a female plant. This is a very common method of vegetative propagation in
C. revoluta. The vegetative propagation in C. circinalis takes place by suckers that
develop on roots. With further growth they start producing new plants.
II. Sexual Reproduction:
Cycas is strickly a dioecious plant, but the male and female plants are
indistinguishable at the vegetative stage. The compact male cone develops at the apex
of the stem in the male plant. However, Cycas is the only genus of Cycadaceae which
does not produce any female cone. Instead, several megasporophylls arise spirally in
acropetal succession around the stem apex of the female plant.
Male Cone:
The surface of male cone is covered with brown scales at young stage. At maturity, the
male cone becomes very large (40-80 cm in length), oval or conical in shape (Fig.
1.17A) which emits odour that can be smelt from quite a moderate distance. The male
cone of Cycas is the largest among the plant kingdom. The male cone has a central
cone axis and numerous microsporophylls are arranged spirally and acropetally
around the axis.
A single micro- sporophyll is a nearly triangular flattened woody structure. It is
differentiated into a proximal wedge-shaped fertile part and a distal sterile part,
tapering into an upcurved apex called apophysis (Fig. 1.17C). Numerous
microsporangia (700 in C. circinalis, 1100 in C. media) are borne on the abaxial
(lower) surface of the microsporophyll except at the apex and the base (Fig. 1.17B).
Microsporangia are borne in groups of 3-5 forming sori that are surrounded by many
single- called delicate hairs (Fig. 1.17D). Each sporangium is oval or circular in shape
having a very short massive stalk.
The dehiscence of sporangia takes place by longitudinal slit (Fig. 1.17D). The
development of sporangia is of eusporangiate type. The sporangial wall is
multilayered with a thickened epidermis and a ill- defined tapetum enclosing
numerous microspore (pollen) mother cells.
Further, microspore mother cells through meiotic division produce numerous
microspores or pollen grains. The pollen grain is oval-shaped having a large rounded
monosulcate aperture. The pollen is bounded by two concentric wall layers; the outer
thick exine and the inner thin intine (Fig. 1.20A).

Female Fructification:
The megasporophylls of Cycas are not organised into a definite cone. Instead, they
arise at the stem apex spirally and acropetaily forming a loose crown. The
megasporophylls are pinnate in nature and are covered with brown hairs called
rementa.
Hence the megasporophylls of Cycas are considered to be the modified foliage leaves.
Megasporophylls are produced every year like the foliage leaves, though they are
produced more in numbers than the foliage leaves.
A single megasporophyll is a flat dorsiventral structure, measuring up to 30 cm in
length. It is differentiated into an upper pinnate lamina and a basal stalk that bears
two rows of opposite or sub-opposite, one to six pairs of ovules (Fig. 1.18A-F). There
is a great variation in structure of megasporophylls and in number of ovules per
megasporophyll in Cycas and these criteria can be applied to identify the species of
Cycas.
There is a great reduction in the structure of megasporophylls and in number of
ovules among the various species of Cycas. C. revoluta is the most primitive species
where the megasporophyll lamina is much dissected and tapers into a point, bearing
3-4 pairs of ovules (Fig. 1.18A). In C. pectinata, the mere serrated lamina is orbiculate
with pectinate margins, bearing 2-3 pairs of ovules (Fig. 1.18 B).
In C. circinalis, the lamina is lanceolate with dentate margins and acuminate apex,
bearing 4-6 pairs of ovules (Fig. 1.18C). The further reduction in lamina has been
observed in C. rumphii (Fig. 1.18D) and C. beddomei (Fig. 1.18E) where lamina is
ovate-lanceolate with acuminate apex, bearing 2-3 pairs of ovules. The maximum
reduction in megasporophyll has been noted in C. normanbyana. Its lamina shows a
mere serrated margins bearing only one pair of ovule (Fig. 1.18F).

Ovule:
The ovules of Cycas are orthotropous and shortly stalked. The ovules are large,
attaining a length of about 6-7 cm with a diameter of 4 cm and is perhaps the largest
amongst the plant kingdom. The mature seeds are elliptical, bilobed and slightly
flattened — they become fleshy and bright orange or red in colour.
There is a single integument which is fused with the nucellus except the top (Fig.
1.19). The integument is differentiated into three layers. The outer layer is fleshy and
pulpy and becomes variously coloured at maturity. The middle layer is very hard and
stony, while the inner layer is fleshy and becomes papery before maturity. There are
frequent mucilage canals and tannin cells in the integument.
The ovule is supplied with three vascular traces (Fig. 1.19). The median strand
supplies the base of the integument, which extends up to the chalazal end of the
nucellus and further ramifies abruptly. The two side strands pass to the integument
which again divide — one branch supplies to the outer pulpy layer and the other to
the inner soft layer.

Megasporogenesis:
A deeply situated cell of the nucellus is differentiated into a large megaspore mother
cell which undergoes meiotic division to form a linear tetrad of four megaspores. Out
of the four megaspores the outer three degenerate, while the lowermost megaspore
becomes functional (Fig. 1.21 A).
The upper free opening of the integument forms the micropyle and a concavity in the
top of the nucellar tissue forms the pollen chamber. After pollination, the pollen
grains are collected in the pollen chamber and the development of pollen grains takes
place in the nucellar tissue.
Essay # 3. Gametophyte of Cycas:
The spore is the first phase of gametophyte generation. The microspore or pollen
grain is the male gametophyte, while the megaspore represents the first stage of
female gametophyte which develops to form a female gametophyte.
Development of Male Gametophyte before Pollination:
The pollen nucleus undergoes mitosis to form endosporically a small lens-shaped
prothallial cell and a large antheridial initial (Fig. 1.20B). The prothalial cell does not
divide further, while the antheridial initial divides into an antheridial cell and a tube
cell (Fig. 1.20C). The pollen grains are released from the sporangium at the 3-celled
stage. After pollination, the further development takes place within the nucellus
tissue of the ovule.

Development of Male Gametophyte after Pollination:

The further development of male gametophyte takes place within a week after
pollination. The tube cell of 3-celled male gametophyte comes out through the pollen
aperture in the form of a pollen tube which penetrates into the nucellar tissue. The
pollen tube destroys all the tissues between pollen chamber and the female
gametophyte.
Thus, the fertilisation is siphonogamous, though it is haustorial in nature. Now, the
antheridial cell within the pollen tube divides into a stalk cell and a spermatogenous
(body) cell (Fig. 1.20D). The spermatogenous cell enlarges considerably and two
blepharoplasts develop at the opposite end of the spermatogenous cell.
Then the spermatogenous cell divides to form two large, top-shaped, motile sperms
with spiral band bearing numerous flagella around the distal end of the sperm (Fig.
1.20E). The sperms of Cycas are very large (180-210 pm) and can be visible to the
naked eye (Fig. 1.20F).
Development of Female Gametophyte:
The female gametaphyte of Cycas develops from the functional megaspore
surrounded by a tough membrane that persists in the gametophyte (Fig. 1.21B). The
nucleus of the megaspore divides by free nuclear divisions forming a large number of
nuclei unaccompanied by wall-formation. In the meantime, the megaspore increases
in size and develops a vacuole in the center which forces the cytoplasm along with
nuclei towards the periphery.
Thus the nuclei lie in a thin film of cytoplasm around the vacuole (Fig. 1.21B). The cell
wall formation starts in a centripetal fashion, from periphery inwards. The wall
formation proceeds very rapidly and as a result the central vacuole in obliterated (Fig.
1.21C).
The entire gametophyte becomes cellular and the tissue thus formed is called
endosperm. The peripheral cells of the gametophyte are small, isodiametric devoid of
any food reserves, while the inner cells are large, irregular and filled up with starch
grains.

Development of Archegonia:
Two to eight cells of the female gametophyte at the micropylar end enlarge in size and
have dense cytoplasm and prominent nuclei. These cells function as archegonial
initials (Fig. 1.21 D). Each archegonial initial divides periclinally to form an outer
small neck initial and a large central cell (Fig. 1.21 E). The neck initial divides by a
vertical (anticlinal) wall to form two neck cells (Fig. 1.21F).
A second division takes place prior to fertilisation, thus the neck actually comprises of
four cells. The central cell enlarges considerably in size and its nucleus divides into a
small ephemeral ventral canal nucleus and a large egg cell (Fig. 1.20G).
The egg of Cycas is reported to be the largest among the living plants, thus it can be
seen with naked eyes. The nucellar tissue above the archegonia disorganises to form
an archegonial chamber (Fig. 1.21G).

Pollination:
Cycas is anemophilous i.e., wind pollinated. The airborne pollen grains remain
suspended in the air and at the same time some cells of the nucellar beak in the ovule
are disorganised to form a viscous fluid. This fluid comes out through the microphyle
in the form of a ‘pollination drop’.
Thus, some of the airborne pollen grains come in contact with the fluid and are
sucked into the pollen chamber through the micropylar canal. The pollen grains are
then deposited and concentrated within the pollen chamber as a result of drying off
the fluid (Fig. 1.20G). At this stage, the ovule increases in size.
Fertilisation:
The sperms and the cytoplasm of pollen tube are released in the archegonial chamber
by the rupture of the basal end of pollen tube. The osmotically rich pollen tube
cytoplasm causes the rupture of neck cells. The motile sperms enter into the
archegonium with a forward and circular motion, ciliary band forming the anterior
end.
The archegonial chamber is flooded with the fertilisation fluid produced by nucellar
cells. The ciliary band of the sperm is left behind on the top of the egg cell. The sperm
nucleus fuses with the egg nucleus and thus a zygote in formed.
Embryogeny:
The zygote enlarges considerably and undergoes numerous free nuclear divisions. As
much as 256 (C. revoluta) to 512 (C. circinalis) free bryo. The cell formation in the
proembryo begins nuclei have been reported. A large central from the basal part and
extends up the periphery, vacuole is formed and the nuclei are arranged.
The entire embryo never becomes cellular. The around the central vacuole (Fig.
1.22A). Most of basal cells are smaller with dense cytoplasm the free nuclei move to
the base of the proem- forming the embryo (Fig. 1.22B).
A dicotyledeous embryo is developed at the tip. The upper cells elongate greatly to
form the suspensor which may be coiled and twisted (Fig. 1.22C). Several egg cells
may be fertilised to form many zygotes, but ultimately only one embryo in an ovule
attains maturity. The layer of cells covering the outer curved face of the embryonal
mass is called the cap.
Seeds:
Cycas seeds are fleshy and remain covered with an orange red-coloured thick seed
coat. The embryo in the seed enlarges much. The nucellus is used up and the seed is
covered with the three-layered seed coat (Fig. 1.22D). The coleorrhiza is the first
region to differentiate in the embryo proper which is quite hard. The plumule
develops later inside the base of the cotyledons. Subsequently the suspensor dries up.
The growth of the embryo in seeds is very slow, taking over a year to mature. Cycas
shows three years reproductive cycle where pollination in first year winter,
fertilisation in second year summer and seed shedding in third year summer. The
development of embryo continues even after the seeds are shed.
The seeds are dispersed by birds due to their attractive fleshy outer pulp. The
germination of the seed is epigeal. The first leaves come out at right angle to the
cotyledons in a decussated fashion, although the remaining leaves develop spirally
(Fig. 1.23).
 Related Articles:
1. Male Gametophytes of Cycas and Pinus | Gymnosperms
2. Essay on Cycas: Top 5 Essays | Class Cycadopsida | Gymnosperms | Botany
Essay on the Life Cycle of Pinus | Class
Coniferopsida | Gymnosperms | Botany

https://www.biologydiscussion.com/essay/gymnosperms/essay-on-the-life-cycle-of-pinus-class-
coniferopsida-gymnosperms-
botany/76971#:~:text=Pinus%20reproduces%20sexually.,formed%20on%20the%20upper%20branch
es.&text=The%20female%20cones%20grow%20very,may%20persist%20for%20several%20years.

In this essay we will discuss about the life cycle of Pinus, explained with the help of
suitable diagrams.
Sporophyte of Pinus:
Pinus is a tall evergreen tree giving rise to a series of widespread horizontal branches
(Fig. 1.57A). In each year, a whorl of branches is produced in the axil of scale leaves.
The branching is restricted to the upper part of the stem, thus giving the tree a
pyramid-like appearance. The main stem is cylindrical and covered with scaly bark.
The branches are dimorphic, bearing two types of shoots: long shoots and dwarf
shoots, or spurs or brachyblasts (Fig. 1.57B). Pinus exhibits two types of leaves, the
scale leaves and the green acicular foliage leaves called needles (Fig. 1.57B). The plant
has a tap root system which becomes elongated at maturity and possesses strong
lateral roots.
The plants are monoecious where the male and female cones are borne on separate
branches in the same plant.
Reproduction:
Pinus reproduces sexually. Pinus is monoecious, but the male and the female cones
are produced on separate branches of the same plant. The male cones develop on the
lower branches, while the female cones are formed on the upper branches. The male
cones, which replace the dwarf shoots, develop in clusters on the base of the current
year’s long shoot at early spring (Fig. 1.61 A).
The number of male cones in a cluster varies considerably from 15 (P. wallichiana) to
140 (P. roxburghii). At the onset of spring, the male cones fall off and simultaneously
the young female cones are borne in pairs or in clusters round the tip of the long
shoot (Fig. 1.61 B).
The female cones grow very slowly and the growth may persist for several years.
Thus, the female cones of different ages may be seen in acropetal succession in the
long shoot (Fig. 1.61B).

i. Male Cone:
The male cone is small (2-4 cm in length) and oval in shape that develops in the axil
of scale leaves. The male cone has a central axis on which 60-150 microsporophylls
are spirally arranged around the axis (Fig. 1.62A).
A single microsporophyll is a membranous stalked structure with a distal expanded
roughly triangular sterile part called apophysis (Fig. 1.62B). Each microsporophyll
bears two sac-like microsporangia on the abaxial surface.
The development of microsporangia is of eusporangiate type, i.e., it develops from a
group of hypodermal cells of the microsporophyll. A mature microsporangium
consists of a multilayered wall, tapetum and microspore mother cells (Fig. 1.62D).
Each microspore mother cell — by meiotic division — produces four microspores or
pollen grains.
Thus, at maturity, a single microsporangium contains numerous pale yellow pollen
grains. The pollen grains are boat-shaped with monosulcate apertures and are
bounded by two concentric wall layers: the outer thick exine and the inner thin intine
(Fig. 1.62C).
The exine on the lateral sides of the pollen is expanded to form two wings (sacci).
Thus the pollen grains of Pinus are bisaccate indicating their anemophilous mode of
pollination. The dehiscence of sporangia takes place by longitudinal slit in dry and
warm environment.
Pinus is wind-pollinated (anemophilous). The pale-yellow pollen gains are released
into the air in a large quantity, so that a pine forest appears yellow at the time of
pollination. This is popularly called ‘sulphur showers’ which occurs specially in the
spring when pine trees are shaken by strong winds.

ii. Female Cone:

Female cones are produced in pairs or in clusters in the axil of the scale leaves. The
female cones mature very slowly. The first year young cone (Fig. 1.61 A, B) is small (1-
2 cm in length), soft, compact and red-green in colour. The second year cone (Fig.
1.61 A, B) is comparatively large (5-8 cm in length), woody, compact and green in
colour.
The fully matured third year cone is much larger (15-60 cm in length), woody, loose
and brown in colour. Here megasporophylls are separated from each other due to the
elongation of the cone axis.
The female cone of Pinus represents a compound shoot; it is a complicated structure.
The female cone is composed of a central axis on which 80-90 megasporophylls,
axillary to bract scale/scale leaves, are arrange spirally (Fig. 1.63A).
The bract scale and ovuliferous scale thus form a seed-scale complex.

A single megasporaphyll consists of two types of scales:

(a) a large woody ovuliferous scale or seminiferous scale bearing two ovules on the
adaxial surface, and
(b) a bract scale or cone scale on the abaxial surface (Fig. 1.63B, C).
Initially, the ovuliferous scale is much smaller than that of bract scale, but after
pollination it becomes larger than the bract scale. The ovuliferous scale is a thick,
large, woody, roughly triangular and brownish structure. Its upper thick exposed part
is known as apophysis.
In the mature cone, the tip of the apophysis becomes the ‘umbo’ (Fig. 1.63C). There
are two separate vascular traces, one supplies to the ovuliferous scale and the other to
the bract scale (Fig. 1.63B). There is no separate vascular trace for ovule.
The development of megasporangium (ovule) is of eusporangiate type i.e., an ovule
develops from a group of superficial cells of the ovuliferous scale.

Ovule:
The ovules of Pinus are anatropous, unitegmic and crassinucellate (Fig. 1.64). The
single integument is free from the nucellus except at the chalazal end. There is a fairly
broad micropylar tube which becomes inwardly curved during pre-pollination stages
and becomes outwardly curved at the time of pollination.
The integument is three-layered, the outer fleshy, the middle stony and the inner
fleshy.

Megasporogenesis:
A hypodermed cell in the nucellar tissue at the micropylar end is differentiated into
an archesporial cell. It divides periclinally to form an upper parietal cell and a lower
megaspore mother cell. The parietal cell further divides to form tapetal layer. The
megaspore mother cell undergoes meiotic division to form a linear tetrad of four
megaspores.
The outer three megaspores degenerate, while the lowermost megaspore becomes
functional (Fig. 1.66A). The upper free opening of the integument forms the
micropyle and a concavity in between the integument and nucellus in the upper part
of the ovule forms the pollen chamber. After pollination the pollen grains are stored
in the pollen chamber and further development of pollen grains takes place in the
nucellar tissue.
Morphological Nature of Ovuliferous Scale:
There is a great controversy regarding the morphological nature of ovuliferous scale
and several hypotheses have been put forward.
Some of the earlier theories which have only a historial values are briefly
discussed:
1. According to Robert Brown (1827), the ovuliferous scale represents an open foliar
carpel bearing naked ovules, present in the axil of bract scale.
2. According to Schleiden (1839), the ovuliferous scale represents an axillary placenta
which is situated in the axil of an axillary leaf (bract scale).
3. According to Alexander Brown (1842), the ovuliferous scale is equivalent to the
first two leaves of an axillary shoot which had fused.
4. According to van Tieghem (1869), the ovuliferous scale represents a single leaf
branch, present in the axil of a leaf (bract).
5. According to the foliar theory of Delpino (1889), both the ovuliferous scale and the
bract scale are the parts of a tripartite bract, where the two lateral fertile lobes of the
bract were fused to form ovuliferous scale and the median sterile part formed the
bract scale.
6. According to ligular or excrescence theory of Sachs (1882) and Eichler (1889), the
female cone represents a simple flower where the cone axis is equivalent to receptacle
or thalamus and bract scale to free carpels. The ovuliferous scale represents an
outgrowth of the carpel (bract scale) as in ligule of Selaginella or the placenta of
angiosperms.
7. According to brachyblast theory of Braun, the female cone is equivalent to an
inflorescence, where ovuliferous scale represents a determinate axillary shoot bearing
two fertile leaves with a single ovule on the dorsal surface of each leaf.
8. Modern hypothesis: On the basis of the comparative studies of the fossil
Cordaitales and Voltziales members, Florin (1951) introduced a terminology ‘seed-
cone complex’ for ovuliferous scale and bract scale, According to Florin, the female
cone of Pinus represents an inflorescence where cone axis is the peduncle, bract scale
is a true bract and the ovuliferous scale is a rudimentary female flower i.e., a modified
reproductive shoot.
This hypothesis has been substantiated by fossil evidences. The occurrence of several
Upper Carboniferous to Triassic seed-cone complex in the members of Cordaitales
and Voltziales supports the above hypothesis (Fig. 1.56A-D). In Cordaianthus,
Emporia and Ernestiodendron of Upper Carboniferous, the cone axis bears several
secondary female reproductive shoots in the axils of bracts.
Each secondary shoot consists of many spirally-arranged sterile and fertile (with
terminal ovule) scales (showing radial symmetry).
The further evolution took place in the Triassic Voltziales like Glyptolepis,
Wallchiostrobus, etc. where the scales were planated (showing bilateral symmetry)
and tangentially fused to form a rudimentary ovuliferous scale following Telome
hypothesis. Thus, it confirms that the ovuliferous scale is a modified reduced
secondary female reproductive shoot and bract scale is actually a reduced bract.
Gametophyte of Pinus:
The spore is the first phase of gametophyte generation. The microspore or pollen
grain represents the male gametophyte, while the megaspore represents the first
stage of female gametophyte which develops into a female gametophyte.
i. Development of Male Gametophyte before Pollination:
The basic pattern of the development of male gametophyte of Pinus is similar to that
of Ginkgo. The pollen grains undergo endosporic development. The pollen nucleus
divides mitotically to produce a small lens-shaped first prothallial cell towards the
proximal end and a large central cell on the distal end (Fig. 1.65A).
The central cell again cuts off a second prothallial cell and an antheridial initial (Fig.
1.65B). Both the prothallial cells are ephemeral and the second prothallial cell
remains attached to the first prothallial cell.
The antheridial initial divides to form a small antheridial cell and a large tube cell
(Fig. 1.65C). The pollen grains are released from the microsporangium at the 4- celled
stage (2 prothallial cells, an antheridial cell and a tube cell).
ii. Development of Male Gametophyte after Pollination:
After pollination, the 4-celled pollen stores in the pollen chamber and remains
ungerminated for about 11 months. The pollen develops the next spring. The tube cell
of the pollen comes out through the pollen aperture in the form of a pollen tube. The
pollen tube proceeds towards the archegonium, penetrating the nucellar tissue of the
ovule. The antheridial cell within the pollen tube divides to form a stalk cell and a
spermatogenous (body) cell (Fig. 1.65D).
The spermatogenous cell divides to form two male nuclei just prior to fertilisation
(Fig. 1.65E). The male nuclei are actually the male gametes which are non-motile and
ephemeral.
iii. Development of Female Gametophyte:
The female gametophyte of Pinus develops from the functional megaspore which
enlarges considerably (Fig. 1.66A). The nucleus of the megaspore divides mitotically
forming a large number of nuclei unaccompanied by wall formation.
The number of free nuclei is constant for a particular species, say for example, it is
2,000 for P. gerardiana and 2,500 for P. roxburghii and P. wallichiana. With the
increase in size, the megaspore develops a vacuole at the centre which forces by
cytoplasm along with nuclei towards the periphery.
Thus, the nuclei lie in a thin film of cytoplasm around the vacuole (Fig. 1.66B). There-
after, the cell wall formation starts in a centripetal fashion, from periphery inwards.
At this stage, numerous radially elongated multinucleate tube-like cells called alveoli
are formed and the wall formation takes place through alveoli. Each alveolus
containing a nucleus at its mouth directs its growth.
Then, cross-walls are laid down on each alveolus to form uninucleate cells. In this
way, the entire gametophyte becomes cellular and the tissue thus formed represents
endosperm or female prothallus (Fig. 1.64).
iv. Development of Archegonia:
The development of archegonia in Pinus is similar to that of Ginkgo. Two to four cells
of the female gametophyte at the micropylar end enlarge in size and have dense
cytoplasm and prominent nuclei. These cells function as archegonial initials. Each
archegonial initial divides periclinally to form an outer small primary neck initial and
a large central cell.
The primary neck initial divides by two vertical walls at right angles to each other
forming a neck of four cells. Thus, the four neck cells are arranged in a single tier as
in P. roxburghii and P wallichiana. However, in P. rigida, P. austriaca the four neck
cells again divide transversely to form eight cells which are arranged in two tiers. The
central cell enlarges very rapidly and its cytoplasm becomes vacuolated (Fig. 1.66C).
The nucleus of the central cell divides into an upper ephemeral ventral canal cell and
a large egg cell. A nutritive layer called archegonial jacket is differentiated around the
archegonium. The nucellar tissue above the archegonia disorganises to form an
archegonial chamber.
v. Pollination:
Pinus is anemophilous i.e., wind-pollinated. The pollen grains are dispersed and
remain suspended in the air for some time. At the same time, the nucellar beak in the
ovule disorganises forming a viscous sugary liquid containing glucose, fructose and
sucrose. This fluid comes out in a cyclic phenomenon (24 hr. cycle) through the
micropyle in the form of a pollination drop either at night or in the early hours of
morning. The pollen grains are caught in the pollination drop and are collected in the
pollen chamber as a result of drying off the fluid. The mouth of the micropyle is then
sealed from the outer environment.
vi. Fertilisation:
The fertilisation takes place after one year of pollination. The pollen tube enters the
tip of the archegonium by forcing itself between the cells of the nucellus. The pollen
tube wall is disintegrated by the enzymes secreted from the egg and eventually two
male nuclei are released. One of the male nuclei fuses with the egg cell and thus a
zygote is formed.
vii. Development of Proembryo:
The zygote nucleus divides by two mitotic divisions forming four nuclei which move
to the base of zygote (Fig. 1.67A). All the four nuclei are arranged in one plane and
only two nuclei are thus visible in lateral view. A synchronous division gives rise to
eight nuclei arranged in two tiers of four each (Fig. 1.67B).
Thus, the upper group of four cells forms the primary upper tier (these cells have no
wall towards upper side) and lower group of four cells forms the primary embryonic
tier (these cells are bounded by wall all around).
The internal division in both the tiers forms four tiers of four cells each (Fig. 1.67C),
and the proembryo thus consists of 16 cells. The lowest tier is known as embryonal
tier, (Fig. 1.67D) which further divides to form embryo. The next tier, called
suspensor tier, elongates considerably to form the embryonal suspensor.
The third tier is known as dysfunctional suspensor (earlier known as rosette tier)
which shows abortive meristematic activity. The uppermost or the fourth tier is called
upper tier or nutritive tier which provides nutrition.

viii. Embryogeny:
The developing embryonal cells are deeply embedded into the gametophyte by the
sevenfold elongation of embryonal suspensor (Fig. 1.67E). Thus the several
embryonal suspensors (designated as Es1, Es2, Es3 and so on) are formed. Proximal
cells of the embryonal mass elongate unequally to form embryonal tubes.
The cells of the embryonal tier are separated from each other at the time of
embryonal suspensor elongation, thus four independent embryos are formed (Fig.
1.67E). This phenomenon is known as polyembryony, because more than one embryo
is formed from a zygote (Fig. 1.67F).
As the polyembryony occurs due to the splitting of a zygote, it is called cleavage
polyembryony. Only a single deep-seated proembryo develops into an embryo and
the growth of other embryos is arrested at different stages of development.
The proembryo divides transversely to form two cells which by further repeated
divisions form an embryo. The embryo is comprised of 3- 18 cotyledons, a distinct
epicotyl root axis and a hypocotyl shoot axis with remnants of suspensor (Fig. 1.68A).
Both the two-year and three- year reproductive cycles are exemplified by Pinus
species.
In two-year type, the pollination and fertilisation take place in late spring of first and
second year, respectively. In three-year type the pollination takes place in spring of
first year, and fertilisation in the spring of third year, a lapse of two years. The seeds
shed in autumn.
Seeds of Pinus:
The seeds are endowed with a well-developed wing which is thin and papery and is
easily detachable at maturity (Fig. 1.68B). The outer fleshy layer of integument and
part of ovuliferous scale contribute to the wing formation (Fig. 1.68C). The seeds are
usually dispersed by wind. The embryo remains embedded within the endosperm.

The seeds of Pinus remain viable for a long time. The germination of seed is epigeal
(Fig. 1.69A, B).
Origin and Relationship of Pinus:
The conifers are large and diversified group of extant gymnosperms. The Pinaceae is
the largest family of the modern conifers. The conifers had evolved from the members
of Permo-Carboniferous Voltziales, commonly known as “transition conifers”. They
reached their climax in mid-Mesozoic forming extensive forests in Northern Europe.
Among the living families, Pinaceae and Araucariaceae are more primitive, probably
evolved during the Triassic age. On the other hand, Cupressaceae and
Cephalotaxaceae are comparatively younger which have evolved probably during
Upper Jurassic to Lower Cretaceous period.
Pinus is the large and well-representative genus of the family Pinaceae. It indicates
relationship with Cycadales, Ginkgoales and Cordaitales.
Relation to Cycadales:
The similarities between Pinus and Cycadales are:
i. The stem anatomy shows a broad pith, large cortex and centripetal wood.
ii. The presence of haplocheilic, sunken stomata in leaves.
iii. The presence of leaf sclerenchyma.
iv. The seeds are with three-layered integument.
v. The presence of free-nuclear divisions in the development of female gametophyte.
Relation to Ginkgoales:
The resemblance between Pinus and Ginkgoales are:
i. The plants are profusely branched showing excurrent habit.
ii. The shoots are dimorphic, bearing long shoots and dwarf shoots
iii. The wood is pycnoxylic.
iv. The leaves are with haplocheilic sunken stomata.
v. The mature wood shows pitting and Bars of Sanio.
Relation to Cordaitales:
Conifers have derived from the members of Voltziales which are considered as
‘transition conifers’ between Cordaitales and Conifers. Thus, Pinus shows striking
similarities with Cordaitales.
These include:
i. The plants are tall-branched trees.
ii. The leaves are sample with paralled veins.
iii. The presence of sclerenchymatous hypodermis in the leaves.
iv. The presence of pycnoxylic wood.
v. The pollen grains are winged.
vi. The ovules are bilaterally symmetrical.
Figure 1.70 shows the life cycle of Pinus:
 Related Articles:
1. Male Gametophytes of Cycas and Pinus | Gymnosperms
2. Essay on Cordaitales | Class Coniferopsida | Gymnosperms | Botany
Essay on the Life Cycle of Gnetum | Class
Gnetopsida | Gymnosperms | Botany

https://www.biologydiscussion.com/essay/gymnosperms/essay-on-the-life-cycle-of-
gnetum-class-gnetopsida-gymnosperms-botany/76855

Here is an essay on the ‘Life Cycle of Gnetum’. Find paragraphs, long and short
essays on the ‘Life Cycle of Gnetum’ especially written for school and college
students.
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.

Essay # 1. Sporophyte of Gnetum:

Gnetum resembles very much in its characteristics to an angiosperm than a
gymnosperm. Based on the studies of 58 characteristics, Gnetum shares more than
60% of the characteristics with angiosperm and about 30% characteristics with
gymnosperm. Gnetum is easily mistaken for a dicot plant unless it is in flowering or
fruiting stage. The plant body is differentiated into root, stem and leaves (Fig 13.2).
1. Root of Gnetum:
External Morphology:
Gnetum shows a typical tap root system which is profusely branched. The mature
roots show normal secondary growth.
Internal Structure:
Internally, the root of Gnetum resembles the root of angiosperm. The root is
differentiated into the outermost layer, epidermis, multilayered cortex and diarch
vascular cylinder. The cortex consists of polygonal or oval-shaped parenchymatous
cells containing starch grains. The thick-walled fibre cells are often present in the
cortex. A single-layered endodermis encircles a multilayered pericycle.
The primary vascular cylinder is diarch, radial and exarch. The secondary growth in
roots is of normal type. The arcs of cambium form below the phloem groups and
above the xylem groups which join together to form a cambium ring. The secondary
xylem consists of tracheids possessing uniseriate bordered pits with conspicuous
Bars of Sanio. The vessels are also present.
The pits on the vessels are bordered or simple, small and multiseriate with or without
inconspicuous Bars of Sanio. The xylem ray is composed of thin-walled
parenchymatous cells containing starch grains. The phloem consists of sieve cells
and parenchyma. The periderm is formed due to the extrastelar secondary growth.
2. Stem of Gnetum:
External Morphology:
Almost all the species of Gnetum, except the tree type like G. gnemon, exhibits two
types of branches viz. dwarf shoots or branches of limited growth and long shoots or
branches of unlimited growth. In climbing and scandent species of Gnetum, the stem
is articulated with prominent joints.
The joint consists of two parts: one just above the node and the other below the node
and these two are separated by an annular groove. In arboreal species such as G.
gnemon, the stem exhibits uniform type of branching.
Internal Structure:
Internally, a Gnetum stem is similar to that of a dicot. In T.S. the stem exhibits more
or less circular outline (Fig. 1.71 A). The following regions are discernible from out-
side inward: the heavily cutinised single-layered epidermis with sunken stomata, a
several- layered (12-16 layers) thick cortex which is differentiated into an outer
chlorenchymatous, a middle parenchymatous and an inner sclerenchymatous region.
The cells of the inner regions are referred to as spicular cells (Fig. 1.71C). Many
fibrous cells are often present which have branched or unbranched pit canals. The
cortex is followed by endodermis and pericycle layers which are not distinct. Next to
pericycle, there is an endarch siphonostele.
The vascular cylinder is comprised of 20-24 collateral, open and endarch vascular
bundles, arranged in a ring (Fig. 1.71 A). The vascular bundles are separated from
one other by wide primary medullary rays. The xylem consists of tracheids and few
vessels (Fig. 1.71 B).
The protoxylem of tracheids have annular or spiral thickening and the metaxylem
have reticulate thickening with uniseriate bordered pits. The phloem is composed of
sieve cells and phloem parechyma. Laticiferous elements are often found in pith as
well as in cortex.

Secondary Growth in Thickness:

In tree species like G. gnemon, the secondary growth is of normal type. The fascicular
cambium joins with interfascicular cambium to form a complete cambium ring that
cuts off a continuous cylinder of secondary xylem towards the inside and secondary
phloem towards the outside. The secondary xylem is produced much more than the
secondary phloem.
In climbing species (G. ula, G. africanum), the anomalous secondary growth is noted.
Initially, the stem shows normal secondary growth. Later, several successive
cambium rings are formed in the cortex.
Thus, several extra- stellar rings of xylem and phloem are formed which are
separated into wedge-shaped bundles because of medullary rays (Fig. 1.71D). Some
of these accessory rings remain incomplete and, as a result, the vascular bundles
become eccentric with regard to pith (Fig. 1.71 E).
The periderm is also formed by the activity of phellogen during the third or fourth
year of growth of the stem.
The wood of Gnetum is composed of vessels, tracheids and xylem parenchyma. The
vessels are of different sizes and have a single pore on their end walls (Fig. 1.72A).
The tracheids are much longer than the vessels and have bordered pits on both of
their radial and tangential walls. The xylem parenchyma cells have simple pits. There
are many uni- to multiseriate vascular rays (Fig. 1.72B, C).
The secondary phloem consists of sieve cells and phloem parenchyma. The companion
cells are totally absent in Gnetum, although some small cells are present near the sieve
tube.
The vascular rays are very large and multi- seriate which appear boat-shaped in T.L.S.
Occasionally, some short biseriate or uniseriate rays are present. The ray cells are
thick-walled and pitted.
3. Leaf of Gnetum:
External Morphology:
Gnetum exhibits leaf dimorphism bearing both the foliage leaves and scale leaves.

A dwarf shoot bears 9 to 10 foliage leaves, arranged in an opposite decussate manner.

The leaves are large, simple having an oval-shaped broad coriaceous lamina with
unicostate reticulate venation. The leaves are exstipulate, shortly petiolate with
entire margin. The leaf of Gnetum resembles a dicot leaf.

Internal Structure:
Anatomically, the leaves of Gnetum resemble those of dorsiventral dicot leaves. The
epidermis (both upper and lower) has undulating walls, covered with thick cuticle.
The mesophyll is differentiated into palisade and spongy parenchyma.
The palisade consists of single-layered compactly arranged elongated cells. The
spongy parenchyma is composed of loosely arranged lobed cells. In addition, some
spicular cells, sclerotic cells, latex tubes and fibres are also present.
The vascular bundles are arranged in a curved in the midrib region. Each bundle is of
conjoint, collateral and endarch type with an adaxial xylem and abaxial phloem.
There are distinct patches of stone cells outside the phloem.
The xylem is composed of vessels, tracheids and xylem parenchyma, while the
phloem consists of sieve cells and phloem parenchyma. Transfusion tissue is present
in the petiole in association with vascular bundles.
According to Takeda. (1913) and Florin (1931), the stomata of Gnetum is of
syndetocheilic type (i.e., both the guard cells and subsidiary cells develop from a
common stomatal initial). But, Maheswari and Vasil (1961) reported that the
development of stomata in G. ula is haplocheilic where the stomatal initial forms
only guard cells, while irregularly arranged epidermal cells around them do not
function as subsidiary cell.

Essay # 2. Reproduction:
Gnetum reproduces sexually. Gnetum is dioecious and both the male and female
strobili (= inflorescence) are compound. The inflorescence is either axillary or
terminal in position which occurs singly or in groups. The inflorescence is composed
of a stout long axis with two opposite decussate, connate bracts at the base and a
series of cup-like bracts called cupules or collars that are superposed one above the
other (Fig. 1.73, 1.74A).
The strobilus becomes compact at young stage because of the suppression of
internodes. Thus, the collars of a young strobilus appear to be continuous (Fig. 1.73).
At maturity, the axis becomes elongated and the collars get separated. There are
many rings of flowers in the axil of collars. The collars are developed in acropetal
succession and the flowers are initiated as mounds of meristematic cells from the
lower surface of a collar.

Male Strobilus:
The male strobilus is compound and has a long slender axis bearing 10-25 whorl of
bracts (collars) (Fig. 1.73). About 12-25 male flowers are arranged in three to six
rings above each collar (Fig. 1.74A). A single ring of 7-15 imperfect female flowers or
abortive ovules is present just above the male flowers.
Male “Flower”:
A male flower consists of two unilocular anthers on a stalk (antherophore) enclosed
in a small sheathing perianth (Fig. 1.74B). The stalk of the anther elongates rapidly at
maturity pushing the anther beyond the collars through a slit in the perianth.
Female Strobilus:
The female strobilus is very much similar to that of the male strobilus in the young
stage. Like male strobilus, the female strobilus consists of an axis bearing several
whorl of collars arranged one above the other (Fig. 1.75A). A ring of 4-10 ovules
(female flowers) is present above each collar (Fig. 1.75B). The male flowers are not
found in the female strobilus. The upper few collars are devoid of ovules and are thus
sterile (Fig. 1.75B).
Ovule:
A single ovule represents a female flower. The ovule is stalked in G. ula, but may be
sub- sessile or even sessile. The ovules are orthotropous, crassinucellate (with
massive nucellar tissue) and are protected by three envelopes (Fig. 1.75C). The outer
envelop which becomes thickened and succulent at maturity is considered to be the
perianth corresponding to the perianth of male flower.
The middle and the inner envelopes are actually the integuments. Numerous
laticiferous ducts and sclerides are present in the perianth with some epidermal
stomata. The middle envelop is called the outer integument which is anatomically
similar to the outer envelop. The inner envelop, i.e., the inner integument, elongates
far beyond the apical cleft of the perianth and forms a long micropylar tube (Fig.
1.75C).
Anatomically, the inner integument is different from the other two envelopes,
because neither sclerides nor stomata develop in the inner integument. The inner
integument is free from the nucellus except at the chalazal end.
Two sets of vascular bundles are formed (Fig. 1.75C), of which the outer set passes to
the perianth and the inner set again divides and one of its branches passes to the
outer integument and the other to the inner integument. All the three envelopes of
ovules develop in acropetal manner. A shallow pollen chamber is present at the tip of
the nucellus.
Megasporogenesis:
Two to four hypodermal cells in the nucellar tissue at the micropylar end is
differentiated into primary parietal cells towards outside and the primary
sporogenous cells towards inside. The primary parietal cells together with nucellar
epidermal cells divide repeatedly to produce a massive nucellus.
The primary sporogenous cells divide to form 8-20 sporogenous cells which are
linearly arranged. The sporogenous cells function as megaspore mother cells which
undergo meiotic division. Since no walls are laid down after meiotic division of
megaspore mother cells, all the four nuclei remain within the mother cell to form a
tetranucleate coenomegaspore. Thus, the female gametophyte of Gnetum is
tetrasporic.
Essay # 3. Gametophyte of Gnetum:
The spore is the first phase of gametophyte generation. The microspore or pollen
grain represents the male gametophyte, while the tetranucleate coenomegaspore
represents the first phase of female gametophyte which develops into a female
gametophyte.
Development of Male Gametophyte before Pollination:
The pollen grain is inapeturate and spherical, bounded by the two concentric wall
layers: the outer thick, spiny exine and the inner thin intine (Fig. 1.74C). The pollen
nucleus divides mitotically to produce a small lens-shaped prothallial cell and a large
antheridiai initial (Fig. 1.76B). The prothallial cell does not divide further and even-
tually degenerates.
The antheridiai initial divides to form an antheridiai cell and a tube nucleus (Fig.
1.76C). The antheridiai cell directly functions as spermatogenous cell. The pollen
grains are released from the microsporangium at this 3- celled stage (one prothallial
cell, an antheridiai or spermatogenous cell and a tube nucleus).
According to Thomson (1961), the prothallial cell is not formed in Gnetum. He
proposed that the pollen nucleus cuts off a tube nucleus and a generative cell (Fig.
1.76B’). The generative cell again divides forming a stalk cell and a body cell (Fig.
1.76C’). Thus, the pollen grains are released at this 3-celled stage (tube nucleus, stalk
cell and body cell).
Development of Male Gametophyte after Pollination:
The exine is cast off during pollen germination. The tube cell of the pollen comes out
in the form of a pollen tube which traverses the nucellar tissue through intercellular
spaces. The prothalial cell remains within the pollen grain and eventually
disorganises.
The spermatogenous cell moves into the pollen tube and subsequently it divides to
form two equal (e.g., G. ula, G. gnemon) or unequal (e.g., C. africanum) male cells
just prior to fertilisation (Fig. 1.76D). The male cells are actually the male gametes
which are non-motile. However, according to Thomson (1961) the stalk cell remains
within the microspore and eventually degenerates. The body cell moves into the tube
where it divides to form two male cells (Fig. 1.76D’).
Development of Female Gametophyte:
At the initial stage, before the gametophyte formation, the nucellar cells immediately
below the megaspore mother cell divide to form a tissue. The cells of this tissue are
arranged in radiating rows. This tissue is termed as ‘pavement tissue’ which
eventually gets absorbed and seems to be nutritive in function.
There is a free nuclear division in the coenomegaspore, as a result a large number of
free nuclei are formed (Fig. 1.77A). The number of nuclei thus formed varies in
different species, viz. 256 in G. gnemon, 512 in G. africanum and 1500 in G. ula. At
this stage, a large central vacuole appears and the free nuclei lie in a thin film of
cytoplasm around the vacuole towards the periphery (Fig. 1.77B). Later, the nuclei in
the peripheral cytoplasm divide repeatedly.
At this stage, the upper part of the gametophyte surrounding a vacuole widens, while
the lower part of the gametophyte shows accumulation of cytoplasm. Thus, the
gametophyte becomes an inverted flask-shaped structure (Fig. 1.77C). The wall
formation starts very slowly from the chalazal end towards the micropylar end. Thus,
the nuclei remain free at the microphylar end even at the time of fertilisation.
The important characteristic in the female gametophyte of Gnetum is the absence of
archegonia. One to three nuclei of the gametophyte in the micropylar end enlarge
several times and accumulate dense cytoplasm around them. These large and densely
cytoplasmic cells are the eggs (Fig. 1.77B). It is important to note that all the eggs do
not mature simultaneously.

Pollination:
Gnetum is wind-pollinated. The pollen grains are dispersed from the anther and remain
suspended in the air for some time. At the free nuclear stage of the female gametophyte, the
nucellar beak in the ovule disorganises forming a viscous sugary liquid which comes out
through the microphyle in the form of a pollination drop. The pollen grains are caught in the
pollination drop.
Due to the drying off of the fluid, the pollen grains are sucked into the micropylar canal and
are finally collected in the pollen chamber. The mouth of the micropyle is then sealed from
the outer environment due to the development of flage (a circular rim or an umbrella-shaped
structure develops from the inner integument) and micropylar closing tissue (a tissue
develops by the proliferation of the inner epidermis of integument at the level of flage).
Fertilisation:
The pollen tube enters the female gametophyte and the male gametes move ahead of tube
nucleus (Fig. 1.77B). The pollen tube ruptures to discharge the male gametes into the egg
cell.
The cell sheath of male cell is left outside the egg cell. Usually one of the male nuclei fuses
with the egg nucleus and thus a zygote is formed. Sometimes, two male gametes may fuse
two different eggs if those eggs are in the vicinity of the pollen tube.
Endosperm:
In gymnosperms, endosperms are cellular and haploid and are formed before fertilisation.
However, in Gnetum the development of endosperms starts before fertilisation very slowly
from lower part of the gametophyte which eventually proceeds upward. After fertilisation,
the wall formation starts in such a way that the cytoplasm divides into many multinucleate
compartments (Fig. 1.78A). Later, the nuclei in each cell fuse to form a single polyploid
nucleus (Fig. 1.78B).
In this stage, the lower part of the gametophyte becomes cellular, while the upper part
remains free nuclear even after fertilisation (Fig. 1.77C). Thus, the development of endo-
sperm takes place even after fertilisation. There is a great variation in the development of
endosperm in Gnetum.
In some cases, the wall formation starts either from the upper part or from the middle part of
the gametophyte instead of the lower part and the whole gametophyte may become cellular.
Though some portions of the endosperms are formed after fertilisation, the characteristic
triploid endosperm through double fertilisation is, however, absent in Gnetum.

Embryogeny:
In all angiosperms (except, Paeonia), the division of zygote is accompanied by wall-
formation; while in all gymnosperms (except Sequoia, Welwitschia), there is a free
nuclear phase in the zygote during the development of embryo. However, Gnetum
occupies an intermediate stage between gymnosperms and angiosperms with regard
to embryo development by having both the free-nuclear divisions as well as cell
divisions.
There is a great variation in the early development of the embryo in different species
of Gnetum.
In C. gnemon, the zygote develops 1-3 small tubular outgrowths. Only one of the pro-
tuberances receives the nucleus and survives, while the remaining protuberances die
out (Fig. 1.79A).
The surviving tubular outgrowth becomes much elongated and branched and
develops in various directions invading the intercellular spaces of the endosperm.
These tubes are called primary suspensor tubes or proembryonal tubes (Fig. 1.79B).
All the primary suspensor tubes remain coiled around each other. At the tip of the
primary suspensor tube, a small cell is cut off which eventually divides by a
transverse wall, followed by a longitudinal wall resulting into four cells.
This is further followed by irregular divisions to form a group of cells. Now, further
divisions take place in some of these cells which eventually elongate to form
secondary suspensor. The rest of the cells at the tip form an embryonal mass (Fig.
1.79C, D).

In G. ula, the early development of the embryo up to the primary suspensor cells is
almost similar to that of G. gnemon. The nucleus of the primary suspensor cell (Fig.
1.80A) divides to form two unequal nuclei, of which the smaller nucleus is cut-off by
a thin wall. This cell is called peculiar cell which forms the embryo (Fig. 1.80B).
The peculiar cell divides twice forming a four-celled stage (Fig. 1.80C, D) which
further divides transversely resulting into a 8-celled embryo (Fig. 1.80E). The
embryonal mass increases in size by the further irregular divisions (Fig. 1.80F). Some
cells of the embryonal mass adjacent to the primary suspensor elongate to form the
secondary suspensor.
Irrespective of the pattern of formation of embryonal mass and secondary suspensor,
the cell of the embryonal mass in Gnetum are small and compact with dense
cytoplasm forming the embryo-proper. The cells of the secondary suspensor are thin-
walled, uninucleate and highly vacuolated. Both the primary as well as secondary
suspensors push the embryo deep inside the endosperm for the nourishment of the
embryo.
At the tip of the embryonal cells, a stem tip with two lateral cotyledons is
differentiated (Fig. 1.81 B). A root tip with a root cap also develops at the opposite
end of the stem tip. Simultaneously, a hump-like structure called feeder is developed
in-between the stem and root tips (Fig. 1.81B). Thus, a mature embryo is composed
of a stem tip, two cotyledons, a large feeder and a root tip covered with root cap.
In Gnetum, polyembryony takes place in various ways. Each of the primary
suspensor tube may develop an embryo, thus a large number of embryos are formed
from a single zygote (Fig. 1.79C).
Sometimes additional embryos may develop due to the proliferation of the
proembryonal mass present at the tip of the secondary suspensor. Sometimes, the
primary suspensor tube branches giving rise to several primary suspensor tubes,
each of which may develop an embryo at its tip.
 Seeds:
Gnetum seeds are oval in shape (Fig. 1.81 A) and green to brown-red in colour. The
seeds remain covered with a three-layered envelop, of which outer is fleshy, middle is
stony and inner is pepary. The nucellus is used up and the embryo is embedded
within the endosperm. Gnetum shows one-year reproductive cycle where pollination,
fertilisation and development of embryo take place in one year.
The germination of seed is epigeal. The seeds of G. ula germinate after one year’s of
res-ting phase.

Figure 1.82 shows the life cycle of Gnetum.

Related Articles:
1. Essay on the Life Cycle of Cycas | Class Cycadopsida | Gymnosperms | Botany