Morphological Features of Cycas:

Cycas is perennial, slow growing evergreen plant and is referred as

living fossil because it occurs as a fossil e.g., C. fusiana. It looks like a
palm tree. Its main plant body is sporophytic, diploid, dominant and
can be differentiated into three parts – roots, stem and leaves. Tallest
species of Cycas is C. media with 20 feet height.

1. Roots:
They are of two types – normal and coralloid roots. Normal roots grow
deep into the soil and form tap root system. Later it is replaced by
adventitious roots. The function of these roots is to fix the plant in the soil
and to absorb water and other minerals.

From the normal roots develop some small lateral apogeotropic

branches near the ground surface. These lateral roots get infected with
bacteria, fungi as well as algae. The entry of these organisms is said to
be responsible for the characteristic, swollen, knob like or coral like
appearance and hence, these roots are called as coralloid roots or
corallorhiza. These roots have minute pores (lenticels like) which are
respiratory in function (aeration). Root cap and root hairs are absent
in coralloid roots (Fig. 2).

2. Stem:
ADVERTISEMENTS:

It is thick, erect, woody, aerial and usually unbranched (caudex).

Branching is rare and it is due to injury or development of
adventitious buds. Surface of the stem is rough due to the presence of
persistent woody leaf bases (Fig. 4). These leaf bases form thick
armour around the stem.

In the armour are distinctly visible the alternating bands of large and
small rhomboidal leaf bases. Larger ones are of foliage leaves and
smaller ones are of scaly leaves and megasporophylls in the female
plant. The leaf bases are spirally and compactly arranged with each
other (Fig. 4).
At t he top is present a crown of leaves
(Fig. 1).

3. Leaves:

Leaves are dimorphic i.e., of two types – scale leaves and foliage
leaves. Both these types of leaves form a crown at the top of the stem.

(a) Scale leaves:

These are small, dry, brown, triangular structures with a thick
covering of brown hairs or rameta. These leaves alternate with green
foliage leaves. These leaves protect the shoot apex and reproductive
structures (Fig. 3).
(b) Foliage leaves:
These leaves are also produced in a crown at the apex of the stem.
According to Coulter and Chamberlain (1910) one crown of foliage
leaves is formed yearly while D.D. Pant (1953) observed the formation
of two crowns per year in C. circinalis. In case of C. revoluta the leaves
are 30 to 150 cm long but in case of C. circinalis they are up to 270 cm
long.

A single foliage leaf is pinnately compound. It is unipinnate and

paripinnate. Each leaf has 80-100 pairs of leaflets which are arranged
on both the sides of adaxial groove of the rachis in opposite or
alternate manner. The rachis is spiny below with the sheathing leaf
base (Fig. 6A). these spines are modified leaflets. Each leaflet is
leathery in texture, sessile elongated, ovate or lanceolate in shape and
has entire margin with acute apex. Each pinna or leaflet contains a
midrib without lateral veins.

In C. micholitzii the leaflet is repeatedly and deeply dichotomised (Fig.

5).
Margins of the pinnae are flat (Fig. 6B) but sometimes they are curved
downwards and inwards (revolute) (Fig. 6C) which give the plant a
specific name C. revoluta. Young leaves have circinately coiled leaflets
which are also covered by hairs or ramenta like those of ferns (Fig. 6
D, E).
Internal Structure of Cycas:
1. Root:
(i) Normal root:
Its internal structure is exactly similar to that of dicot root. It is
circular in outline and can be differentiated into epiblema, cortex and
vascular tissue.

a. Epiblema:
It is the outermost limiting layer and consists of single layer of thin
walled cells. Some of its cells give rise to root hairs.

b. Cortex:
Epiblema surrounds the multilayered zone of thin walled
parenchymatous cortex with numerous intercellular spaces. The cells
of the cortex are filled with starch. Some tannin cells, mucilage cells
and sometimes sphaeraphides (calcium oxalate crystals) are also
present in the cortex. The innermost layer of the cortex forms the
endodermis which is characterised by the presence of casparian strips.

c. Vascular tissue:
Endordermis is followed by multilayered parenchymatous pericycle.
Vascular bundles are radial. Xylem is diarch and exarch i. e.,
protoxylem is towards the periphery). The protoxylem consists of
spiral tracheids whereas the metaxylem consists of scalariform
thickenings. Vessels are absent. Alternating with the protoxylem
groups are present phloem cells consisting of sieve tubes and phloem
parenchyma. The companion cells are completely absent (Fig. 7A, B).

Secondary Growth:
The mature normal root shows secondary growth on both the lateral
sides of primary xylem. Along with the inner side of primary phloem
develops the cambium. It cuts off secondary phloem on outer side and
secondary xylem on the inner side. After sometime the cells of the
pericycle opposite to the protoxylem strands also become
meristematic and behave as cambium, cutting phloem on the outer
side and xylem on the inner side.

Thus, a complete ring of cambium is formed which forms a complete

ring of secondary xylem on the inner side and complete ring of
secondary phloem on the outer side. The primary phloem is crushed in
the due course of development and appears in the form of crushed
layer above the secondary phloem.

Simultaneously the formation of periderm also starts. The cells of the

outermost layer of the cortex become meristematic (also called cork
cambium) and start cutting cork cells on the outer side and secondary
cortex on the inner side. In the course of the formation of cork, the
cells of the epiblema are crushed (Fig. 8 A, B)

(ii) Coralloid Root:

The transverse section of the coralloid root is similar to that of normal
root and it can be differentiated into epidermis, cortex and vascular
tissue.

a. Epidermis:
In young root, it is similar to normal root. However, in old root the
outermost tissue is periderm. It consists of 2 to 5 layers of dead cells.
b. Cortex:
The cortex is wider in comparison with the normal root. A greenish
algal zone is present almost in the middle of the cortex and divides it
into outer cortex and inner cortex (Fig. 9A, B).

The algal zone consists of loosely connected, radially elongated thin

walled cells occupied by blue green algae (Anabaena cycadae, Nostoc
punctiforme, Oscillatoria), bacteria (Azotobacter, Pseudomonas
radicicola) and some fungi. The main function of these roots is
nitrogen fixation due to the presence of cyanophycean members.
Endodermis is similar to normal root.

c. Vascular tissue:
Endodermis is followed by multilayered parenchymatous pericycle.
Vascular bundles are radial. Xylem is triarch and exarch.
Secondary growth is very rare or absent. No secondary xylem or
secondary phloem are developed although cork and cork cambium are
present.

Comparison between Normal Root and Coralloid Root:

Normal root:
1. Develops from the radicle, tap root system

2. Geotropic

3. Such characters are absent

4. Such infection is absent

5. Root hairs are present

6. Cortex is smaller

7. Such division is absent

8. Diarch

9. Secondary growth present

10. Main functions are: fixation of plant, absorption of water and

mineral nutrients

Coralloid root:
1. Develops from the normal roots

2. Apogeotropic

3. Develops from the normal roots

4. Dichotomously branched and appears like coral

5. Gets infected with algae, bacteria and fungi

6. Absent
7. Cortex is wider in comparison

8. Due to presence of the algal zone in the cortex, it is differentiated

into outer cortex and inner cortex

9. Very little or absent

10. Main function is nitrogen fixation

2. Stem:
A transverse section of young stem is similar to dicot stem. It is
irregular in outline due to persistent leaf bases. Internally, it can be
differentiated into epidermis, cortex and vascular cylinder.

a. Epidermis:
It is the outermost layer of the stem. It is made up of compactly
arranged thick walled cells. Epidermis is ruptured due to the armour
of persistent leaf bases (Fig. 11A).

b. Cortex:
Epidermis encloses the cortex. It forms the major portion of the stem.
It is composed of parenchymatous cells which are filled with large
number of starch grains. These starch grains are the source of sago
starch. Therefore, C. revoluta is popularly known as sago palm.
Scattered in the cortex are various mucilage canals. Each mucilage
canal is lined by many radially elongated epithelial or secretory cells
(Fig. 10). which secrete mucilage. These canals are connected with
those of the pith with the help of the medullary rays. The innermost
layer of cortex is endodermis. It is not distinct.

c. Vascular Cylinder:
The vascular cylinder is surrounded by not very conspicuous pericycle.
Like dicot stems vascular cylinder consists of many conjoint,
collateral, open, endarch vascular bundles arranged in a ring
(ectophloic slphonostele). The xylem consists of tracheids and Xylem
parenchyma (Fig. 11B).

Vessels are absent. Outside the xylem is the phloem which consists of
sieve tubes and phloem paraenchyma. Companion cells are absent.
The Xylem is separated from the phloem with the help of primary
combium. The cells of the primary cambium are brick shaped.

The cells lying in between the vascular bundles form the medullary
rays. These are parenchymatous and connect the pith with the cortex.
Each medullary ray is one celled wide and 1 to 20 cells long.

3. Pith:
In the centre of the stem is present large canals leaf traces massive
pith consisting of parenchymatous cells which are rich in starch (sago
starch). A large number of mucilage canals are also present, which are
exactly similar in structure with the mucilage canals present in the
cortex.

4. Leaf Traces and Girdle Traces:

The leaf traces are scattered in the cortex of the stem and constitute
the vascular tissue of the leaves from the main vascular cylinder. Each
leaf receives four traces, two of which are direct traces and the other
two are given out from the opposite side of the direct traces.

These two traces take a round around the main vascular cylinder in
opposite direction through cortex and then enter the leaf base of
opposite side from the point of their origin from the stele. These leaf
traces are known as girdle traces or indirect trances and are peculiar
structures in the stem of Cycas.

Secondary growth:
It is a slow process. At first a complete ring of cambium is formed by
the development of interfascicular combium in between the adjacent
vascular bundles. The cambium cuts off secondary xylem on the inner
side and secondary phloem on the outer side.
Tracheids consist of multiseriate bordered pits. This cambial ring is
short-lived and new cambial ring is formed every year in the pericycle
of the cortex. Wood formed by this method (more than one) cambium
ring is polyxylic and manoxylic (large amount of parenchyma is cut off
in the xylem. (Fig. 13).

5. Rachis:
A transverse section of the rachis is somewhat rhomboidal in outline,
but a little higher up it is shield shaped. Its internal structure can be
differentiated into epidermis, cortex and Vascular bundles.

a. Epidermis:
It is the outermost covering. It is made up of compactly arranged thick
walled cells. It is single layered, covered with thick cuticle and has
stomata.

Hypodermis:
Epidermis is followed by hypodermis. It is differentiated into outer 2-
3- layers of chlorenchyma (Chlorophyll containing thin walled cells)
and inner 4-6 layers of sclerenchyma (thick walled, lignified cells; Fig.
14A, B).

Ground tissue:
Below the sclerenchyma is present a large tissue made up of thin
walled parenchymatous cells. It is called ground tissue. In this region
are present many mucilaginous canals and vascular bundles.
b. Vascular bundles:
Vascular bundles are arranged in the shape of inverted Greek letter
‘omega’ [Ω; Fig. 14 A], Each vascular bundle is conjoint, collateral,
endarch, open and diploxylic i. e., consists of centripetal and
centrifugal Xylem and is surrounded by bundle sheath.

Xylem is present towards the inner side and consists of tracheids and
xylem parenchyma. Vessels are absent. Phloem is present towards the
outer side of the vascular bundle. It consists of sieve tubes and phloem
parenchyma. Companion cells are absent, Cambium is present in
between the xylem and phloem.

In rachis the vascular bundles are endarch at the base (centrifugal

xylem is well developed, protoxylem faces towards the centre showing
endarch condition, centripetal xylem is not developed), mesarch in the
middle (centripetal and centrifugal xylem are present showing
diploxylic condition) and exarch at the apex (centripetal xylem is well
developed, triangular and exarch, centrifugal xylem is much reduced
and in the form of two patches lying one on each side of the
protoxylem elements of centripetal xylem) due to twisting of the rachis
(Fig. 15 A-C).

6. Leaflet:
The leaflet of Cycas is dorsiventral and hypostomatic (the stomata are
present at the lower surface only). In a transverse section the leaflet
can be differentiated into a swollen midrib portion and two lateral
wings (Fig. 16A, B).

Its internal structure is as follows:

a. Epidermis:
It is the outer most single layer made up of squarish cells. The upper
epidermis is complete whereas the lower epidermis is interrupted by
several sunken stomata present in the region of the wings. The upper
and lower epidermis is covered by a thick layer of culicle.

b. Hypodermis:
Below the epidermis occurs the thick walled sclerenchymatous
hypodermis. It is single layered in the region of blade but in the region
of mid rib it becomes 2-3 layered thick. Two to five layers of
sclerenchymatous cells are also present above the lower epidermis
only in the region of the mid rib. It helps in checking the rate of
transpiration and protects the tissue from excessive heat.

c. Mesophyll:
A well-developed mesophyll tissue is present in the leaflet. It is
differentiated into palisade tissue and spongy parenchyma. Palisade
tissue is present in the form of continuous layer below the
sclerenchymatous hypodermis. Spongy parenchyma present only in
the wings directly above the lower epidermis. It is made up of loosely
arranged oval cells filled with chloroplast. These cells have many
intercellular spaces filled with air.

d. Vascular bundle:
A single large vascular bundle is present in the mid rib region of the
leaflet. It is surrounded by a single layer of sclerenchymatous cells,
known as bundle sheath. The vascular bundle is conjoint, collateral,
open and diploxylic. Xylem is present towards the dorsal surface and
phloem is present towards the ventral surface.

Xylem and phloem are separated by a non-functional strip of

cambium. Centrifugal xylem is represented by two small groups on
either side of the protoxylem. The remaining space of the vascular
bundle is filled with thin walled parenchymatous cells.

e. Transfusion tissue:
Groups of tracheidal cells, separated by some parenchymatous cells, or
directly in contact with the centripetal xylem, the bundle sheath are
present in the leaflet. It is called primary transfusion tissue. The cells
of this tissue are short and wide with are reticulate or bordered pitted
walls.

A zone is present on either side of the midrib between the palisade and
spongy layers. It is three layered and is composed of elongated
colourless cells. These cells run paralled to the leaf surface from the
midrib to the margin. This zone is called accessory transfusion tissue
or secondary transfusion tissue or hydrostereom or radial
parenchyma.
On either side of the leaflet it is connected with the primary
transfusion tissue present around centripetal xylem of the vascular
bundle. Primary and secondary transfusion tissue help in the lateral
conduction of water. The presence of transfusion tissue is to
compensate for the unbranched condition of the midrib and it
probably serves as a later conducting channel of water.

Reproduction in Cycas:
Cycas reproduction by two method – Vegetative and Sexual

1. Vegetative reproduction:
It is the simplest method of reproduction. It takes place by the
formation of bulbils or adventitious buds. These buds develop on the
stem in the axil of the scale leaves. A bulbil is an oval structure, broad
at the base and pointed at the apex. It consists of dormant stem in the
centre covered by numerous brown scaly leaves.

On detachment from the stem, a bulbil starts to germinate by

producing many roots from the lower side and a leaf towards the
upper side. A bulbil from male plant will develop only into male plant
while the bulbil from the female plant will form only female plant
because cyas is strictly dioecious (Fig. 1A, B).

2. Sexual Reproduction:
Sexual reproduction in cycas is oogamous (the female gamete i.e., egg
cell is significantly larger than the male gamete and is non-motile).
Cycas is sporophytic and strictly dioecious i.e., male and female sex
organs are borne on separate plants.
Male Reproductive Organs:
Male plant of cycas produces every year a single male cone (Fig. 1B) at
its apex. In the formation of the male cone the apical meristem is used
up, and therefore, the growth of the steam checked for some time
some time but later an apical meristem is formed at the base of the
cone, which pushes that on one side so that the growth of the stem is
resumed again.

Such growth of the stem is called sympodial (Fig. 18 A, B). The male
cone is largest in the plant kingdom (approximately 500 cm or more in
length).
Longitudinal Section of Male Cone:
Each cone is an ovoid or conical structure (Fig. 19A). A longitudinal
section of male cone shows that each one consists of a central axis
around which, a large number of leaf like structures called as
microsporophylls are attached at right angle in a compact, spiral,
acropetal succession (Fig. 19.B). The maturation of the sporophylls
takes place in a spiral manner i.e., from apex to base. However, a few
sporophlills at the apex and base remain sterile. Fig. 19 B.
Structure of microsporophyll:
Each microsporophyll represents a stamen. It is a flattened, woody
and triangular structure. It is differentiated into upper or distal, sterile
region called apophysis (Fig. 20 A) and proximal wedge shaped fertile
part. Each microsporophyll bears several hundred microsporangia
(pollen sacs) on its abaxial surface (more than 1000, Fig. 20B-D).

Microsporangia are arranged in clusters of 3 to 6. Each cluster or

group of microsporangia is called sorus. In between the
microsporangia indusial hairs are present which help in the dispersal
of the microspores and protect young sporangia.
Development of Microsporangium:
The development of microsporangium is of eusporangiate type i. e.,
develops from a group of cells called microsporangial initials (Fig.
21A). These cells are hypodermal in origin and divide by periclinal
walls into upper primary wall cells and lower primary sporogenous
cells.

Primary wall cells divide and redivide to form three to six layered wall
below the epidermis. Simultaneously primary sporogenous cells also
divide and redivide irregularly to form a mass of cells known as
sporogenous tissue.

At this time from the peripheral cells of the sporogenous tissue or the
inner most wall layer differentiates into a single celled, nourishing
layer known as tapetum (Fig. 21C, D). The outermost layer of the
sporangial wall forms the epidermis or exothecium.

The rest of the sporogenous tissue increases in size and functions as

spore mother cells. These are the ultimate cells of sporophytic phase.
Each spore mother divides cell by reduction division to form four
haploid microspores (Fig. 21D-F).
A mature microsporangium is sessile or shortly stalked consists 5-6
wall layers (outer single layer exothecium, innermost single layer
tapetum and rest endothecial layers). The wall of sporangium encloses
large number of haploid microspores. The tapetum is used up during
the development of microspores. Only the signs of disintegrated
tapetum can be seen at maturity.

Female Reproductive Organs:

Female reproductive organs are megasporophylls. Each female plant
every year produces numerous megasporophylls in acropetal
succession above each crown of foliage and scaly leaves. There is no
female cone formation. The number of the megasporophylls is much
more than the number of the foliage leaves on the stem.

During the formation of the megasporophylls the apical meristem is

not used up like that of male cone and therefore, the growth of the
stem continues, and thus in female plant growth is monopodial.

Structure of Megasporophyll:
Each megasporophyll (carpel) is regarded as a modified leaf. It is
about 12.7 cm to 25.4 cm long and can be divided into 3 parts: upper
leafy portion, middle ovule bearing portion and lower stalk. Ovules are
formed on the lateral side of the middle portion. The upper portion is
pinnate and each pinna is tapering to a point.

Two lateral rows of ovules are present on the lateral side of the middle
portion. In Cycas there is a great variation regarding the pinnate
character of megasporophyll and the number of ovules per sporophyll
as a result of which in various species of Cycas gradual reduction in
megasporophylls can be traced.

The megasporophylls of C. revolula (Fig. 22A) are pinnate whereas

those of C. circinalis C. rumphii and C. beddomei (fig 22 B-D) are
ovate lanceolate structures. In C. pectinata and C. siamensis they are
orbicular or rhomboidal structures (Fig. 22B, F).

The laminar portion is well developed in C. revoluta, C.pectinata and

C. siamensis but reduced in C. circinalis, C. beddomei and C. rumphii
(fig. 22). The margin of lamina is serrate or dentate in C.circinalis,
C.beddomei and C. rumphii. The number of ovules differ in different
species of Cycas. It is 1-6. pairs in C.revoluta, C. Circinalis and only
one pair in C. norambyana. Megasporophylls are covered by yellow or
brown hairs.
Structure of ovule (megasporangium):
The ovules are sessile and are borne laterally on the stalk. The ovules
of Cycas are largest in plant kingdom (7 cm long in C. thoursaii, 6 cm
long x 4 cm diameter in C. circinalis) and can be seen by naked eye.
The ovule is green when young and is covered by hairs. At maturity its
colour changes to orange and hair also fall off.

The ovules are orthotropus (short and straight) and unitegmic (with
one integument).

The integument is very thick and consists of three distinct

layers:
(i) Outer, green or orange fleshy layer called outer sarcotesta

(ii) Middle, yellow stony layer called sclerotesta and

(iii) Inner fleshy layer or inner sarcotesta.

The parenchymatous tissue inside the integument is called nucellus.

The integument encloses all the nuclellus except at one point. This
point or opening is called micropyle. Just below the micropyle, the
cells of the nucellus form the nucellar beak.

Some of the cells of the nucellar beak dissolves and forms a cavity like
structure called pollen chamber. Just below the pollen chamber is
present an archegonial chamber. Micropyle leads into the pollen
chamber. Just below the floor of the archegonial chamber 3-6
archegonia are present towards the micropylar end.

The ovule is supplied by three vasular traces (Fig. 23). The central
vascular trace enters the chalazal end of the nucellus. The inner and
outer vascular traces divide into two each, one branch supplies the
outer fleshy layer and the inner fleshy layer. Thus, the outer and inner
fleshy layers receive the vascular supply but the middle stony layers
get no vascular supply (Fig. 23).
Development of Ovule:
When megasporophyll is young; in its middle portion 4-6 ovules arise
as a hypodermal mass of meristematic cells on the lateral side. These
meristematic cells divide and redivide to form a mass of
parenchymatous cells known as nucellus.

Soon the neighbouring cells at the base are also activated and they
grew upwards forming the integument which surrounds the nucellus
on all sides except at the top where a small opening is left which is
known as micropyle (Fig. 24A, B). In the beginning the nucellus and
integument are free but afterwards due to intercalary growth both of
them fuse except in the region of micropyle.
Deeply situated in the nucellus, any one cell enlarges and functions as
megaspore mother cell or embryo sac cell. It divides by meiosis to
form a linear tetrad of four megaspores. Upper three degenerate (Fig.
24 C-F) and lower most is functional. It is the first cell of female
gametophyte.

Gametophytic Phase:
Microspores are formed in microsporangia and megaspores are
formed in ovules after meiosis. Cycas is heterosporous because it
produces two types of spores.

Development of male gametophyte:

Each microspore is unicellular, unicelled structure with two layered
wall. Outer wall is known as exine while inner is known as intine.

The development of male gametophyte takes place in two

stages:
Stage I:
Development of male gametophyte before pollination: Development of
male gametopheyte or germination of pollen grains starts in situ i. e.,
they are still inside the microsporangium.

First its nucleus divides into two, one of them goes towards the lower
side and is separated from the other by a cresent shaped wall resulting
in the formation of two unequal cells. The lower smaller cell is called
as prothallial cell and the upper bigger one as antheridial cell (Fig. 25
A, B).

The prothallial cell does not divide further but the antheridial cell
divides to form a generative cell (Fig. 25C) which is in close contact
with the prothallial cell and distal tube cell with a large nucleus. This
stage is called 3-celled stage consisting of a prothallial cell, generative
cell and tube cell.

At this stage microsporangia dehisce and the shedding of the pollen

grains takes place, (tapetum disintegrates, sporangium becomes dry,
cells shrink, sporangial wall ruptures radially at the line of
dehiscence). Further development of the pollen grains (II stage) takes
place after pollination.
Development of Female Gametophyte:
The functional megospore (also called embryo sac cell or first cell of
female gametophyte) is haploid and it starts its development in situ i.
e., within the nucellus (Fig 26 A). It absorbs the surrounding cells of
the nucellus and enlarges considerably. Its nucleus divides by free
nuclear divisions and as a result a large number of nuclei are formed.
A vacuole develops in the centre.

It pushes the free nuclei and cytoplasm of the megaspores towards its
periphery. (Fig. 26 B) Now the wall formation starts from periphery to
the centre (centripetal wall formation). It results in the formation of a
cellular tissue called endosperm or female prothallus or
megagametophyte (Fig. 26 C). The endosperm in Cycas is a haploid
tissue formed before the fertilization.
Endosperm is nutritive in function. Simultaneously, a tiny space
develops on the upper side of the ovule between nucellus and the
female gametophyte due to degeneration of certain nucellar cells. This
is called archegonial chamber.

Development of archegonium:
The archegonia develop from the gemetophytic cells lining the
archegonial chamber towards the micropylar end. Any cell enlarges in
size and functions as archegonial initial (Fig. 27A). It divides
transversely into an upper primary neck cell and a lower central cell
(Fig. 27 B). The primary neck cell divides by a longitudinal division to
form two neck cells. These cells form the neck of the archegonium.

The central cell enlarges its size and its nucleus divides to form a
ventral canal nucleus and an egg nucleus (Fig.27 C). No wall is formed
between the venter canal nucleus and egg nucleus. Therefore, there is
no neck canal cell. Later on venter canal nucleus disorganises. The egg
of the Cycas is largest in all living plants measuring 5 mm in diameter.
Structure of Archegonium:
The mature archegonium consists of a neck of two neck cells. The
archegonial neck opens in the archegonial chamber. There is no neck
canal cell. There is no venter either. The egg and the venter canal
nucleus remain surrounded by the cells of the endosperm. These cells
act as archegonium jacket.

Pollination:
The pollination is anemophilous. The cells of nucellar beak present in
the pollen chamber disintegrate and form a viscous fluid. This fluid is
cohesive in nature. This fluid oozes out of the micropyle and collects in
the form of a pollination drop.

The pollen grains present in the air current at their 3-celled stage, are
entangled in the pollination drop. Gradually the pollen drop dries up
and the pollen grains are sucked into the pollen chamber through
micropyle. Further drying of this drop seals up the micropyle (Fig. 28)
Pollen drop helps in collecting the pollen grains at the micropyle in all
gymnosperms.
II Stage. Development of male gametophyte (after
pollination):
After a definite period of rest (pollen grains may lie inside the pollen
chamber for quite some time say for four months, the interval between
pollination and fertilization is about 20 days in C. revoluta). Further
development of the male gametophyte takes place in the pollen
chamber. The exine breaks up and intine comes out in the form of
pollen tube.

It may be branched or unbranched and acts as an absorbing organ

(Pollen tube acts only as an absorbing organ or hustorium because the
sperms are formed only after the complete development of the pollen
tube) Pollen tube, penetrates the nucellar tissue and comes to lie in the
archegonial chamber. The generative cell divides into a lower stalk cell
and upper body cell (Fig. 25D). The stalk cell does not divide further.
The body cell divides at the time – micropyle of fertilization.

Now two blepharoplasts appear, one at each pole of the nucleus of the
body cell in a transverse position. Body cell divides longitudinally into
two sperm cells, each of these having a single nucleus, blepharoplast
and a small amount of cytoplasm. Each sperm cell later on develops
into a sperm and blepharoplast gives rise to cilia. The sperms are
liberated in the pollen tube by the breaking of the sperm mother cell
(Fig. 25E, F).

The sperms of the Cycas are largest in the plant kingdom (180-210 µ)
and visible to the naked eye. Each sperm is more or less triangular, top
shaped mobile structure having five to six spiral bands with thousands
of cilia with a single large nucleus. By means of their cilia, they move
freely in the pollen tube (Fig. 25F).

Fertilization:
At the time of fertilization, the nucellar tissue between the pollen
chamber and the archegonial chamber disorganise and simultaneously
the venter canal nucleus also disintigrates. The pollen tube reaches the
archegonial chamber (Fig. 29). The tip of the pollen tube ruptures
releasing two male gametes and fluid contents.

Due to this archegonial chamber becomes moist and the sperms move
freely in it with the help of cilia. Only a single sperm enters violently in
each archegonium through neck. Only the male nucleus of the sperm
fuses with the egg nucleus to form a zygote or oospore (2x). The
fertilization in Cycas takes place with the help of motile sperms.

This process is known as zooidogamy. It is accompanied by pollen

tube formation, a phenomenon known as siphonogamy. Sometimes
more than one sperm enter the archegonium but the male nucleus
which first reaches near the nucleus fertilizes the egg. Rest male nuclei
degenerate. It is called polyspermy.
Embryogeny:
The fertilized egg, zygote or oospore is the first cell of the sporophyte.
The zygote contains dense cytoplasm and a large nucleus. It enlarges
in size and finally forms the embryo. In this whole process one year
time is utilised. The nucleus moves at the base and starts dividing by
free nuclear divisions to form about 256 free nuclei (Fig. 30A, B).

The nuclei and cytoplasm of the central region disorganise forming a

central vacuole (Fig. 30C). Subsequently wall formation takes place
from the base and advances towards the upper side forming a small
mass of cells. This embryonal mass of cells is called proembryo (Fig.
30D). It is meristematic in nature. Some of the nuclei in the upper
region remain without cell walls.

The proembryo soon gets differentiated into three zones

(Fig. 30E):
(1) Upper haustorial zone at the micropylar end

(2) The middle suspensor zone

(3) Tha basal embryonal zone.

The haustorial zone absorbs the nutrition for the developing embryo.
Suspensor zone elongates considerably forming a long, spirally coiled
suspensor, which pushes the embryo in the food containing cells of the
endosperm. The embryonal zone is differentiated into embryo. It is
differentiated into two cotyledons (three in C. circinalis).

A plumule is then differentiated in between the cotyledonary

depression. The embryonal part lying below the cotyledonary
attachment is the hycotyl. A radicle is now differentiated at the base of
the hypocotyl. It is protected by a hard pad like protective covering
called coleorhiza. It provides protection to the radicle (Fig. 30 F, G).

Several archegonia may be fertilized in an ovule of Cycas. It results in

the formation of several zygotes. These zygotes may undergo initial
development but usually one embryo reaches maturity.
Structure of seed:
After fertilization, the ovule is transformed into seed. The nucellus and
the inner layer of integument are used up as nourishment by
developing embryo. The mature seed appears as orange-red or reddish
brown structure.

It comprises the following structures:

a. Testa or seed coat:
It is formed by the outer brightly coloured fleshy layer and the middle
layer of the integuments.

b. Micropyle:
It is present in the form of small opening at the top of the seed.

c. Endosperm:
Inner to the seed coat lie the wall tissues called endosperm. The cells
store a large amount of food material.

d. Embryo:
Embedded in the endosperm lies the embryo. It consists of two
cotyledons, plumule and radicle. The embryo remains suspended in
the endosperm by a long spirally coiled suspensor (Fig. 32).
Thus, a mature seed of Cycas represents three generations:
Seed Coat:
It is formed by the integument and represents parent-sporophytic
generation.

Endosperm:
Represents the gametophytic generation.

Embryo:
It represents the new sporophytic generation.

Testa is sweet in taste and emits pleasant odour. The two

characteristics i. e., red colour and pleasant odour are responsible for
their zoochorus (orinthochorous) dispersal.
Germination of seed:
Seeds remain viable for not more than a few months. Under suitable
conditions the seed starts germination. It absorbs water and embryo
expands. The expansion of the embryo breaks open the hard seed coat.

The coleorhiza protrudes out and is pierced by the growing radicle

which grows down and forms the tap root or primary root (Fig. 32A).
The cotyledons do not come out of the seed coat but they absorb food
from the endosperm for the growing embryo.

The stalk of the cotyledons elongates to carry the plumule out of the
seed coat. The growing plumule at first, forms a few scaly leaves and
then foliage leaves. The young foliage leaves show circinate vernation
(Fig. 32 B-D).

The seed germination is hypogeal because the cotyledons remain

underground enclosed in the female prothallus or endosperm.
However, according to some workers cotyledons do not come out of
the seed during germination as they are haustorial in nature, but their
bases remain exposed, hence the seed germination may be said to
epigeal.

Economic Importance of Cycas:

1. Several species of Cycas of e.g., C. revoluta, C. circinalis C. bedomei
are grown in the gardens as ornamental plants.

2. From the stem ‘sago’ a kind of starch, is obtained. Hence, it is also

called sago palm (C. revoluta)

3. Leaves of C. circinalis are used to prepare hats, baskets and mats.

The leaves of Cycas are extensively used for decorative purposes and
floral decoration.

4. The resin obtained from C. rumphii is applied to malignant ulcers.

5. Juice of tender leaves of Cycas is useful for vomiting and flatulence

(the presence of excessive gas in the digestive tract).
6. Pollens of C. circivalis are narcotic. Its seeds ground to paste with
coconut oil are useful for sores and swellings.

7. Seeds of Cycas are roasted and are used as food in Assam and
certain islands.

8. The seeds and stem of C. revoluta are used in making wire in Japan.

9. The flour of seeds of Cycas is called Indum Podi and is used in the
preparation of cakes and porridges.

10. The young succulent leaves of Cycas are also cooked as vegetable.
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