GYMNOSPERMS

Introduction
• All seed bearing plants are included under the division Spermatophyta.
• It consists of two sub-divisions, Gymnosperms & Angiosperms.
• Gymnosperms - simple and primitive seed-bearing plants without
flowers.
• Naked seed plants (ovule without ovary)
• Gymnos = naked; sperma = seed
• Possess unprotected naked ovules, borne on megasporophylls.
• Also known as “phanerogams without ovary”
• Originated in Palaeozoic era (541- 252 million years ago)
• Dominant in Jurassic and Cretaceous periods of Mesozoic era
• Many primitive gymnosperms are extinct (Cycadofilicales, Cordaitales,
Bennettitales)
• Examples of some Gymnosperms – Cycas, Pinus, Gnetum, Ephedra, Taxus,
Ginkgo, Cedrus, Welwitschia, Podocarpus, Abies, Araucaria, etc.
▪ A small group of plants
▪ 70 genera and 900 species
▪ Distribution: Temperate & Tropical regions
▪ The present day gymnosperms are Cycadales, Coniferales, Gnetales & Ginkgoales
▪ Cycadales & Ginkgoales have long geological history and hence they are often
described as living fossils.
▪ Cycadales: cycads whose leaves are like palm leaves
▪ Coniferales (cone bearing evergreen gymnosperms): largest group with 500 sp. The
largest & longest living vascular plants belong to this group are Sequoia (tallest tree in
the world), Pine, Spruce, Cedar etc.
▪ Gnetales: Gnetum, Ephedra etc. Ephedra is economically important and it yields
ephedrine used for the treatment of asthma & some respiratory ailments.
▪ Ginkgoales: Ginkgo tree (maiden hair tree) is the only living representative
 General Characters
Morphological Characters
• Living gymnosperms are evergreen trees or shrubs with
xerophytic adaptations.
• Plant body is sporophytic, differentiated into-root, stem and leaves
• Roots show symbiotic association with algae (coralloid roots of Cycas with
Nostoc & Anabaena) and fungi (Mycorrhizal association of Pinus roots).
Algae helps in nitrogen fixation and fungi helps in absorption of minerals.
Morphological Characters
• The stem is generally erect, branched and woody. However, it is unbranched
in case of Cycas and underground in Zamia.
✓ Presence of leaf scar is a characteristic feature of gymnosperm

✓ Leaves are generally dimorphic in nature.

• Cycas shows circinate vernation (young leaves curved inside).
✓ Presence of circinate vernation in Cycas is a strong evidence for the
pteridophytic origin of Gymnosperm. Therefore, Cycas acts as a connecting
link between Pteridophyta & Gymnosperm.
Anatomical Characters
• Leaves have thick cuticle & sunken stomata.
• Mesophyll in leaves usually differentiated into palisade and spongy tissues
in Cycas. But is undifferentiated in Pinus and Cedrus.
• Leaves do not have lateral vein. Lateral translocation of nutrients takes place
through transfusion tissue.
• Gymnosperm possess well developed vascular system. Vascular bundle is
open and collateral.
• Xylem consists of tracheids and parenchyma.
• Vessel is present in Gnetum only.
• Phloem consists of tubes and phloem parenchyma.
• Stem shows secondary growth. The wood may be manoxylic (Cycas) and
pycnoxylic (Pinus).

• Tanniferous cells are present in cortex region.

• Roots are diarch (two vascular bundles) to polyrach (many vascular
bundles).
Reproductive Characters
• Gymnosperms are heterosporous (i.e. produce two different spores).
• Megasporangia are produced on megasporophyll
• Microsporangia are produced on microsporophyll
• These sporophylls aggregated to form compact cones or strobili.
• Cones/ strobili are monosporangiate. Ephedra - bisporangiate
• Male cones in gymnosperms are short-lived; however female cones
persist for many years.
• Microsporangia are found on the abaxial (lower surface) of
microsporophyll. Microsporangial development is eusporangiate
type.
Male and female cones in gymnosperms
• Female cone is formed by the aggregation of megasporophylls.
• The megasporophylls may be foliar as in Cycas or cauline (woody) as in
Pinus.
• The megasporangium is better known as ovule.
• Ovules are orthotropous and unitegmic (ex. Cycas), bitegmic in Gnetales
• Ovular integument in Gymnosperm is differentiated into three layers
• The outer and inner layers are fleshy, while the middle layer is stony.
• Microspores are liberated at the 3-celled stage in Cycas, 4-celled stage in Pinus,
and 5-celled stage in Ephedra.
• Male gametes are non-motile, with the exception of Ginkgo and Cycas
• The number of archegonia in a female gametophyte may vary. There are several
archegonia in Cycas, and only one in Pinus. The archegonium of Gnetum is
represented by ovum only. Archegonium has a single egg and a venter canal cell
There are no neck canal cells.
• All gymnosperms are usually wind-pollinated.
• Pollen grains deposited in wet pollen chamber.
• Fertilization is siphonogamous (through pollen tube).
• The pollen tube function as sperm carrier.
• Embryo development is meroblastic (i.e. embryo develops from some part of
zygote). Angiosperms also exhibit meroblastic development.
✓ Endosperm development takes place before fertilization. Hence, endosperm is
haploid in nature.
✓ Double fertilization or triple fusion is absent in gymnosperm.
• Polyembryony (development of many embryo) is very common in Gymnosperms.
• Polyembryony results from
a) fertilization of more than one egg or
b) division of zygote (Cleavage polyembryony).
• The naked ovule develops into seed & ovular integuments into seed coat.
• The number of cotyledons in a seed is 1 or 2 (Cycas), or many (Pinus).
• Seed winged in Pinus
• There is distinct alternation between sporophytic and gametophytic generations.
• The diploid sporophytic phase is dominant, whereas the haploid gametophytic phase
is reduced
• Gametophytic phase is dependent on the sporophytic phase
 Classification of Gymnosperms

• The most recent system of classification for gymnosperms is proposed by

Christenhusz et al. (2011).
(i) Order Pteridospermales (Cycadofilicales)
• Commonly known as seed ferns.
• The members of this order are extinct.
• They form a connecting link between cycads and ferns.
• Originated in the Devonian period of the Palaeozoic era and formed the
dominant part of the vegetation of the carboniferous period.
• Pteridospermales are trees or small plants with fern-like leaves.
• Seeds are borne on frond-like leaves.
• Cones are altogether absent.
• Members: Lyginopteris, Medullosa, Calamopitys.
ii) Order Bennettitales (Cycadeoidales)
• Members of this order are also extinct.
• They originated during the Mesozoic era, became dominant in the
Jurassic period, and declined in the upper Cretaceous period.
• Bennettiales are trees whose stem is covered with an armour of
persistent leaf bases.
• Male sporophylls are frond-like and they form a loose crown.
• Female sporophylls are found in a cone-like structure.
• Members: Williamsonia, Cycdeoidea, Bennettites, Ptilophyllum
(iii) Order Pentoxylales
• A group of low shrubs.
• Female inflorescence consists of numerous cones, spirally arranged on a
peduncle.
• Each inflorescence bears numerous ovules borne on a central axis.
• Megasporophylls, ovuliferous scales, or interseminal scales are absent.
• Male flowers are borne terminally on dwarf shoots.
• Unilocular sporangia terminate in short branchlets of the appendages or
in microsporophylls.
• Members: Pentoxylon, Sahnia.
(iv) Order Cycadales
• Members of this order are known as cycads.
• They are believed to have originated from seed ferns.
• This order includes 9 living genera.
• The members have palm- like habit.
• Male and female sporophylls are grouped into cones except in the genus
Cycas where the female sporophylls form a loose crown.
• Members: Cycas, Zamia, Ceratozamia, etc.
(v) Order Cordaitales
• This order consists of extinct plants which originated in the Devonian
period.
• In the Carboniferous period, they flourished well and formed the thick
forests of the world.
• Finally, in the Triassic period, they vanished from the scene, Cordaitales
are large trees with flat and strap-shaped leaves.
• Fructifications are formed as cones.
• Members: Cordaites, Callixylon, Mesoxylon.
(vi) Order Coniferales
• This is the largest order of gymnosperms, having 52 genera.
• They originated in the Carboniferous period.
• The members of this order constitute nearly three-fourth of the living
gymnosperms.
• Coniferales include large trees and shrubs.
• Leaves are needle-like, scale-like, or sometimes flattened.
• Male flowers are arranged in more or less compact clusters or catkins.
• Female flowers are usually clustered to form a cone.
• Members: Pinus, Araucaria, Podocarpus, Thuja, etc.
vii) Order Taxales
• This order includes a single family, Taxaceae, with only 5 genera.
• Plants are characterized by the absence of resin canals in the aerial
parts.
• Male cones are borne on the axils of leaves, either singly or in groups.
• Ovules are borne terminally on axillary branches.
• Members: Taxus, Austrotaxus, Amentotaxus, Nothotaxus
(viii) Order Ginkgoales
• Members were wide spread during the Triassic and Jurassic periods.
• They began to decline gradually in the Jurassic period.
• Now, there is only one living species, Ginkgo biloba - "living fossil".
• Ginkgoales are medium-sized trees.
• Leaves are flattened and lobed in various ways with dichotomous
venation. Male and female flowers are collected into strobili.
• Members: Ginkgo biloba, the only living member. Baiera, Ginkgodium,
Sphaenobaiera, etc are extinct forms.
(ix) Order Gnetales
• Most highly evolved group of gymnosperms.
• It is regarded as a connecting link between gymnosperms and
angiosperms.
• The members are small trees or climbing shrubs.
• Opposite leaves, dioecious, and dicotyledonous embryos are the major
characteristics of the group.
• Members: Gnetum, Ephedra, Welwitschia.
 CYCAS
Division: Gymnospermae
Class: Cycadopsida
Order: Cycadales
Family: Cycadaceae
• Widely distributed genus
• It includes 25 species worldwide and six species are common in India
• Occurs wild or cultivated in tropical and sub-tropical regions.
• Mostly found in South of Eastern Hemisphere
• It is evergreen plant, in India represented with 6 species- Cycas
revoluta, C. pectinata, C. siamensis, C. beddomei, C. rumphii and C.
circinalis.
• C. circinalis and C. beddomei are the common South Indian
species.
• The plants grow in xerophytic conditions.
• It is cultivated as ornamental plant in the garden.
• Cycas is called living fossil.
External morphology
• Evergreen, slow-growing and long-living palm like plant (1.5-3m)
• Adult cycas plant is a sporophyte
• The tallest species is C. pectinata (12m)
• Sporophyte is dioecious
• Plant body is differentiated into roots, stem and leaves.
Root
• Two types of roots:
✓ Normal positively geotropic tap roots
✓ Negatively geotropic coralloid roots
i. Normal tap roots
• Primary root - short living and forms the tap root. Further it
maybe replaced by adventitious roots.
• Primary root is thick and short and its lateral branches are thin
and long. They are called normal roots.
• These roots are positively geotropic & their main functions are
anchorage and absorption of water and mineral nutrients.
• Root hairs and rot caps are absent
ii. Coralloid roots
• Specialized apogeotropic roots grow on the surface of soil
• They are repeatedly dichotomously branched and appear
as coral-like masses (hence the name coralloid roots).
• They are more frequent on young
plants.
• A specific algal zone with colonies of
Anabaena or other blue-green algae is
present in the cortex of these roots.
• The exact function of endophytic
algae in coralloid roots is not known.
• But, they may perhaps help in
nitrogen-fixation.
• Coralloid roots possess lenticels
which help in respiration.
Stem
• Young stem is tuberous and subterranean and its
apical part is covered with brown scale leaves.
• Older stem - thick, columnar and woody, with a
crown of pinnately compound and fern-like leaves
at its top.
• Stem is covered with persistent and woody leaf
scars.
• Stem is usually unbranched.
• But sometimes, after a certain age, or in response to
injury, it may give out branches.
• In this case, the shoot meristem gets divided into two parts
and the stem becomes dichotomously branched.
Leaves
(a) Foliage leaves or assimilatory leaves
• Large, green, pinnately compound and spirally arranged photosynthetic
leaves with circinate vernation.
• They form a crown at the top of the stem.
• Each leaf has 80-100 pairs of leaflets arranged on either sides of the rachis in
opposite or alternate manner.
• The leaflets are sessile, elongated and ovate or lanceolate.
• Each leaflet has only a single midvein, and it is devoid of lateral veins.
• The rachis of a very young leaf is circinate with coiled leaflets, similar to
those of ferns.
• Young leaves are densely covered with ramenta (one cell-thick, thin and
brown scales).
(b) Scale leaves
• These are small, dry, rough, triangular and non-photosynthetic leaves, thickly
covered with ramenta.
• Their function is to protect the apical meristem and other aerial parts.
• These leaves possess persistent leaf bases which form a part of the armour of
the old stem.
• Often, bulbils develop in the crevices of scale leaves.
• When detached from the plant, they help in vegetative propagation.
• Bulbils may be provided with scale leaves and foliage leaves.
• Both foliage leaves and scale leaves are arranged in close alternate whorls at
the apex of the stem.
 Internal structure of rachis
• Rachis is circular, biconvex, or flat in outline.
• Woody & thick
• Differentiated into epidermis, hypodermis, ground tissue & vascular tissue.
• EPIDERMIS
✓ Outermost layer
✓ Uniseriate
✓ Covered with thick cuticle
✓ Stomata sunken & irregularly scattered
✓ Stoma consists of two guard cells & two subsidiary cells
• Just below epidermis, is the hypodermis.
• Composed of two types of cells.
• Outer 2 or 3 layers are composed of thin-walled chlorenchymatous cells
& remaining layers consist of sclerenchymatous cells.
• Below hypodermis, parenchymatous ground tissue, with many mucilage
canals.
• VBs are arranged in a typical inverted omega like structure (Ώ) or horse
shoe shaped manner.
• VB are diploxylic, conjoint, collateral, & open.
• Surrounded by uniseriate or multiseriate pericycle.
• Bundle surrounded by a single layered bundle sheath composed of
thin-walled sclerenchymatous cells.
• They are diploxylic (both centrifugal [endarch] & centripetal [exarch]
xylem.
• Xylem is located towards the centre .
• Phloem towards the periphery
• Between them thin strip of cambium is present
• There is difference between the arrangement of xylem & phloem in VB in
the upper, middle & basal regions of the rachis.
• Basal region – VB have only centrifugal xylem
• Middle region – Groups of thick-walled cells developed just behind the
protoxylem elements & finally differentiate into centripetal xylem.
• Upper region – More centripetal xylem than the centrifugal xylem.
• At extreme tip, centrifugal xylem is completely absent.
• Base – Endarch
• Middle – Psuedoendarch
• Apex - Exarch
 INTERNAL STRUCTURE OF LEAFLET
• Dorsiventral
• Xerophytic adaptations
• Single, unbranched midrib without lateral veins.
• Leaflet is swollen in the midrib region.
• Margins are curved in some species Eg: C. revoluta & C. beddomei
• Straight or flat in others (Eg: C. circinalis, C. pectinata & C. numphii)
• 3 main parts
Epidermis , hypodermis & mesophyll
• EPIDERMIS
✓ Outermost layer
✓ Uniseriate
✓ Cuticle present
✓ Upper epidermis is continuous
✓ Lower epidermis is interrupted by minute pits.
✓ Sunken haplochelic stomata present
• Sclerenchymatous hypodermis.
• Multi-layered in the midrib region & single layered in other parts.
• Hypodermis is followed by the mesophyll which is differentiated into
palisade & spongy parenchyma.
• Palisade parenchyma – consists of single layer of vertically elongated &
compactly arranged columnar cells. Act as photosynthetic or assimilatory
zone.
• Spongy parenchyma – Consists of several layers of loosely arranged cells
that are confined to the wings. Act as aerating zone.
• Both are rich in chloroplasts.
• In the wing region, long, colourless &
transversely elongated parenchyma cells
present between palisade & spongy
parenchyma which represent secondary or
accessory transfusion tissue.
• Single VB in the midrib region.
• Surrounded by parenchymatous bundle
sheath.
• VB is conjoint, collateral , open &
diploxylic.
• Centripetal xylem & two groups of
centrifugal xylem, one on either side of the
centripetal protoxylem.
• Phloem is composed of sieve tubes & parenchyma.
• Thin layer of cambium present
• Sphaeraphides present
XEROPHYTIC ADAPTATIONS OF LEAFLETS
• Presence of thick cuticle
• Epidermis with thick-walled parenchyma cells.
• Sclerenchymatous hypodermis
• Stomata are sunken & distributed only on the lower epidermis.
• Unbranched midrib.
• Presence of transfusion tissue.
Internal structure of normal root
• Similar to dicot root
• It has three parts,
❖ Epiblema
❖ Cortex
❖ Stele or central vascular cylinder
• Epiblema, or piliferous layer - single layer of thin-walled cells.
• Some cells give rise to unicellular root hairs.
• Cortex - multilayered & composed of thin-walled parenchyma cells filled
with starch.
• Some cells contain tannin and mucilage.
• Mucilage cells are secretory and usually
they remain bordering mucilage canals.
• Innermost layer of cortex forms
endodermis.
• Casparian bands are present in the
endodermal cells.
• Stelar region is bounded by a
multilayered and parenchymatous
pericycle.
• Its cells are thin-walled and starch-filled.
• Vascular tissues form a central diarch to
tetrarch stele, or very rarely a polyarch
stele.
• Diarch stele - presence of two patches of
protoxylem points.
• Xylem consists of xylem tracheids.
• Tracheid - one celled, non-living,
elongated xylem element with thick
lignified and pitted cell walls
• Vascular bundles show radial
arrangement of xylem and phloem.
• Xylem bundles alternate with phloem bundles.
• Xylem is exarch.
• The tracheids of the protoxylem have spiral thickenings, while those of the
metaxylem have scalariform thickenings.
• The centre of the root is occupied by metaxylem, or metaxylem and
parenchyma.
• Pith is reduced or absent.
• In young root, pith is present. But, it disappears, or becomes reduced, once
secondary xylem develops during secondary growth.
• Root shows secondary growth which starts by the formation of cambium
strips inner to the primary phloem strands.
• These strips cut off secondary phloem
towards the outer side and secondary
xylem towards the inner side.

• As a result, the primary phloem gets

crushed, but the primary xylem still
occupies the centre of the stele.

• Secondary xylem maybe traversed by

parenchymatous medullary rays.

• A distinct phellogen, or cork cambium,

develops from the outer layers of the
cortex.
• It forms phellem or cork on its outer
side and phelloderm or secondary
cortex towards the inner side.

• Phellem, phellogen and phelloderm

collectively constitute the periderm.

• Periderm formation is same as in dicots

Internal structure of coralloid root
• Anatomy of coralloid root is similar to that
of normal root.
• But, in coralloid root, cortex is
differentiated into three distinct regions,
outer, middle and inner.
• Outer cortex is composed of compactly
arranged polygonal cells
• Inner cortex is formed of thin-walled
parenchyma cells
• Middle cortex forms the algal zone
• Algal zone is made up of a single layer of
loosely arranged thin-walled and radially
elongated cells, with intercellular spaces.
• These cells are occupied by blue-green
algae, such as Anabaena cycadae, Nostoc
punctiforme, etc., in a symbiotic
association with the plant.
• Some fungi and bacteria (e.g.,
Pseudomonas, Azotobacter) are also found
in this zone.
• Algal zone formation takes place after the
entrance of endophytic algae.
• Coralloid roots show little or no
secondary growth.
• Cortex, in general, contains tannin cells
and sphaeraphides.
• Sphaeraphides are globular clusters of
minute crystals.
• In structure, coralloid roots differ from
normal roots in the following respects.
• (i) Their cortex is differentiated into three
regions.
• (ii) Secondary tissues are poorly developed
or absent in them.
Internal structure of stem
• Anatomy of Cycas stem is similar to the stem of dicotyledonous
angiosperms.
• Distinguishing features of Cycas stem - large pith, broad cortex and a narrow
zone of conducting tissue.
• Also, the stem consists of leaf bases and woody scales.
• Young stem is irregular in outline.
• It is differentiated into epidermis, cortex and vascular cylinder.
• Epidermis - outermost layer, covered with a thick cuticle. It is discontinuous
due to the presence of persistent leaf bases.
• Cortex - major part, composed of parenchymatous cells, rich in starch grains.

• Cortex is traversed by many mucilaginous canals and leaf traces.

• Leaf traces are concentric and hence are called girdles.

• Innermost layer of cortex is endodermis and is followed by pericycle.

• Both these layers are not distinct in young stem.

• Vascular cylinder is very small. It consists of several vascular bundles,

arranged in a ring, forming an ectophloic siphonostele.

• Vascular bundles are conjoint, collateral and open and the xylem is endarch.
• Individual bundles are separated from each other by parenchymatous
medullary rays.
• Xylem is composed of only tracheids and xylem parenchyma; vessels are
altogether absent.
• The tracheids of protoxylem have spiral thickenings, whereas those of
metaxylem have scalariform thickenings.
• Phloem is composed of sieve tubes and phloem parenchyma. Companion
cells are absent.
• An ill-defined cambium may be present between primary xylem and primary
phloem.
• A well developed parenchymatous pith is present in the centre.
• Its cells are rich in starch. Some cells contain tannins and mucilaginous
substances also.
• Pith may be traversed by scattered mucilage canals.
• A striking anatomical feature of Cycas stem is the presence of leaf traces, or
girdle traces, in the cortical region.
• They are the vascular strands, supplied to leaves.
• Each leaf receives four traces. Two of them arise on the same side where the
leaf is located.
• They enter the leaf directly and hence are called direct traces.
• The other two are opposite the leaf. They enter the leaf after turning around
the vascular cylinder. They form a girdle around the vascular cylinder, they
are called girdle traces.
• Before entering the rachis, these traces divide to form several strands and
hence there are several vascular bundles in the rachis.
Secondary growth of the stem
• Secondary thickening takes place in young stem essentially in the same way
as in a dicot stem.
• It is accomplished by the activity of a continuous cambial ring, formed from
fascicular and interfascicular cambia.
• Interfascicular cambium, formed between the vascular bundles, unite with
fascicular cambium to form a complete cambial ring.
• This ring cuts off secondary xylem on the inner side and secondary phloem
on the outer side.
• Multiseriate bordered pits are present on the walls of the secondary xylem.

• The cambium also forms well developed parenchymatous medullary rays.

• After a short period, cambial ring stops functioning. Then, a new cambium
develops in the pericycle or in the inner cortical layers.

• This cambium, like old ones, forms secondary xylem towards the inner side
and secondary phloem towards the outer side.

• This cambium also becomes inactive after some time and then a new cambium
develops once again.

• The whole process is repeated several times at irregular intervals, resulting in

the formation of a few alternating concentric rings of xylem and phloem.
• In C. pectinata, about 20 rings of cambium may be formed. Thus, the
monoxylic young stem becomes polyxylic.
• Secondary growth in Cycas is somewhat anomalous and it takes place not
annually, but at irregular intervals. So, the growth rings are not annual rings.
• Cambium is not functional throughout, it is alternately active and inactive.
• Secondary growth results in a polyxylic condition by the development of
several concentric rings of vascular bundles outside the primary one.
• Presence of leaf gaps and concentric vascular bundles in Cycas are certainly
prominent pteridophyte features and they suggest a strong affinity between
Cycas and ferns.
• Extrastelar secondary growth takes place by the formation of phellogen (cork
cambium) which forms phelloderm (secondary cortex) on the inner side and
phellem (cork) on the outer side. Lenticels are also formed in the usual way.
Cycas - Reproduction
• Cycas reproduces by vegetative and sexual methods.
Vegetative reproduction
• Vegetative reproduction takes place by means of bulbils.
• Develop in crevices of scale leaves and leaf bases at the basal part of
an old stem.
• Produces new plant on detachment.
• The bulbils, formed in male plants, develop to male plants, and those
formed in female plants develop to female plants.
• From roots, suckers may often develop. They grow horizontally on the
ground for some distance and then develop to new plants (e.g., C. circinalis)
Sexual reproduction
• Cycas - heterosporous, heterothallic and dioecious plant.
• Heterosporous - two different kinds of spores (microspores & megaspores).
• Heterothallic - male and female gametophytes develop from different kinds
of spores, namely from microspores and megaspores respectively
• Dioecious - male and female sex organs develop in separate plants. So, there
are separate male plants and female plants.
• Male reproductive organs are called cones or strobili, and female organs are
called megasporophylls.
• Microspores – microsporangia - microsporophylls
• Megaspores – megasporangia - megasporophylls
✓ Microsporophylls get clustered and compactly arranged on male plants,
forming male cones.
✓ But, megasporophylls are only loosely arranged on female plants, without
forming female cones.
• Plants do not normally produce male cones or female megasporophylls until
they are more than ten years old.

Microsporophyll
Microsporophyll
• Microsporophylls are soft and fleshy when they are young and at maturity,
they become hard & woody.
• They are large in the middle region and small at the base and tip of the cone
axis.
• A mature microsporophyll is a more or less triangular and flattened structure.
• It is differentiated into a proximal wedge-shaped fertile part and a distal disc-
like sterile part.
• The sterile part tapers to an upcurved projection called apophysis.
✓ The upper or adaxial surface of the microsporophyll is sterile. But, its lower
or abaxial surface is fertile.
• Numerous microsporangia, arranged in definite groups called sori. Each
sorus consists of 3-6 microsporangia, arising from a central indusial papilla.
• In b/w the sporangia, there are many delicate unicellular or two-celled
epidermal hairs, called indusial hairs - protect young sporangia and also help
in the dissemination of microspores.
Microsporangia
• Microsporangia or pollen sacs are the microspore-producing organs.
• attached to the lower surface of microsporophylls by a short stalk.
• A microsporangium consists of a central core of sporogenous tissue, enclosed
by sporangial wall.
• Microsporangial wall is multi-layered and is differentiated into 3 regions -
outer exothecium, middle endothecium, and inner tapetum.
• Exothecium - thick-walled, cutinized cells.
• Endothecium - thin-walled cells
• Tapetum - nutritive cells
• Sporogenous tissue consists of a large number of microspore mother cells
(microsporocytes) and microspores.
Development of microsporangia
• Development of microsporangia is eusporangiate type (each microsporangium
develops from a group of hypoderemal initials).
• Each hypodermal cell of the fertile lower surface of the microsporophyll can
function as a sporangial initial.
• During sporangial development, it divides periclinally and forms an outer
primary wall cell and inner archesporial cell.
• The wall cell undergoes repeated anticlinal and periclinal divisions and forms
the multilayered sporangial wall.
• The archesporial cell undergoes repeated irregular divisions in all possible
planes and gives rise to the sporogenous cells.
• The cells of the last generation grow in size, round off and function as
microspore mother cells (pollen mother cells or microsporocytes).

• These cells represent the last stage of sporophytic generation.

• Tapetal and endothecial layers of the sporangial wall disintegrate and form a
nutritive medium for the development of microspores

• Exothecial cells thicken along their radial and tangential walls.

• Microspore mother cells float in this medium for some time. Then, they
undergo meiosis and form numerous tetrads of haploid microspores or pollen
grains.
Dehiscence of microsporangium
• With the maturity of microspores, microsporangium dehisces and releases the
microspores.
• During this, the cone axis elongates. This causes the separation of individual
microsporophylls from each other and the exposure of microsporangia.
• Soon, the wall of the exposed sporangia dries up and breaks open along the
line of dehiscence. This results in the shedding of microspores.
Microspores
• Microspores, or pollen grains, are globular uninucleate and haploid
structures, covered by a thick spore wall.
• Spore wall has two layers - outer exine and inner intine.
Megasporophylls

• Cycas is unique in that in female plants megasporophylls are only loosely

arranged and are not compactly organised to form female cones. So,
megasporophylls directly serve as female organs.

• In the absence of cone formation, apical meristem remains unaffected.

• Megasporophylls arise in close spirals around the stem apex. They are
produced in large numbers, more numerous than foliage leaves.

• Megasporophylls are considered as modified foliage leaves.

Cycas-Megasporophyll
The megasporophylls
for Cycas is leaf-like
with ovules at the
base.

The enlarged one has

been fertilized and is
developing into a seed
• Each of them has two parts, namely a basal stalk or rachis and a terminal
pinnate lamina.

• Ovules are formed on the lateral sides of the stalk.

• The number of ovules varies from 2 to 10, depending upon the species.

• The size and shape of megasporophylls are highly variable among the
different species of Cycas.

• The margin of the lamina is serrate or dentate in C. circinalis, very much

dissected and tapering in C. revoluta, and pectinate in C.pectinata.
Ovule or Megasporangium
• The ovule, or megasporangium of Cycas is sessile, or sometimes short-
stalked, (upright with chalaza, funicle and micropyle in straight line), oval or
spherical, and large-sized.
✓ The largest ovule in the plant kingdom.
• When young, it is green and hairy. But, at maturity, it loses its hairs and
becomes red or orange-coloured.
• Ovule consists of a large central mass of parenchymatous cells, called
nucellus.
• It is surrounded by a single integument that protects the ovule.
• The integument fuses with the body of the ovule, except at the apex of the
nucellus, where it forms long micropyle.
• At this part, the apical portion of the nucellus forms a nucellar beak which
forces its way into the micropyle.

• The integument is very thick and is differentiated into three distinct layers,
namely the outer and inner fleshy layers, called outer sarcotesta and inner
sarcotesta, and the middle hard and stony layer, called sclerotesta.

• Mucilage canals and tannin cells are usually present in outer fleshy layer.

• After fertilization, this layer becomes the seed coat or testa.

• Inner fleshy layer is composed of parenchymatous cells. Most of it will be

used for the development of ovule. So, in a mature ovule, it will be a thin
layer.
• A mature ovule is supplied with three vascular strands from the
megasporophyll one median and two lateral.

• The median strand penetrates into the inner sarcotesta extends to the chalazal
end of the nucellus and breaks up into small traces.

• Each lateral strand divides into two branches, one to inner sarcotesta and
other to outer sarcotesta.

Development of ovule

• The ovule of Cycas develops from a few hypodermal cells of the fertile basal
part of the megasporophyll.

• These ovule initials undergo repeated division and form small outgrowth.
• Cell division continues and a mass of parenchymatous cells is formed. It
form the nucellus.
• Then, the basal cells of the nucellus undergo repeated division and the
resulting daughter cells re-organize themselves to form the integument.
• The integument surrounds the nucellus all around, leaving a small opening at
the top, called micropyle.
• Initially, nucellus and integument are free from each other. But later,
integument fuses with the chalazal part of the nucellus due to intercalary
growth.
• At a later stage, a deeply situated cell of the nucellus differentiates into a
megaspore mother cell.
• This cell is larger in size, with dense cytoplasm and prominent nucleus.
• It undergoes meiosis and forms a linear tetrad of four haploid megaspores.

• The lowermost megaspore enlarges in size and becomes the functional

megaspore.

• The others disintegrate to provide nourishment for the functional megaspore.

Gametophytes
• Cycas is heterosporous. It produces microspores and megaspores.
• Microspores germinate to form male gametophytes, and megaspores
germinate to form female gametophytes.
• Thus, spores represent the last stage of sporophytic generation and spore
mother cells represent the first stage of gametophytic generation.
(i) Microspore germination and the formation of male gametophyte
• Microspore is a uninucleate cell and it represents the 1st cell of the male
gametophyte.
• Its germination begins within the microsporangium before liberation and is
completed within the ovule after pollination.
• Thus, the development of male gametophyte is completed partially inside the
microsporangium before pollination, and partially within the pollen chamber
of the ovule after pollination.
Development of male gametophyte before pollination
• Microspore germination begins in situ within the microsporangium before the
release of spores.
• During this, each microspore divides transversely to form two unequal cells,
an upper small prothallial cell and a lower large antheridial cell.
✓ Prothallial cell remains undivided. But, antheridial cell divides to form a
small generative cell and a large tube cell.
✓ The dispersal of microspores takes place at this 3-celled stage.
• Microspores are very light and are carried away by air currents.
• Further development of male gametophyte takes place within the pollen
chamber of the ovule.
Pollination
• Anemophily
• Mature microsporangium dehisces, liberating large numbers of 3-celled
microspores.
• Microspores are blown away by wind to the ovules of female plants.
• Here, they get entangled in an adhesive fluid or mucilage that oozes out from
the micropyle of the ovule.
• With the drying up of this fluid, the microspores are sucked into the pollen
chamber where they remain dormant for nearly four months.
Development of male gametophyte after pollination
• After a period of nearly four months, the generative cell of the microspore
within the pollen chamber divides to form a stalk cell or sterile cell and a
body cell or spermatogenous cell.
• Exine of microspore breaks open and intine grows out as a pollen tube.
• The pollen tube penetrates the nucellar tissue and grows towards the female
gametophyte.
• It acts as a haustorium and also as a sperm carrier.
• The stalk cell, which is in contact with the prothallial cell does not divide
further.
• The body cell increases in size and divides into two sperm mother cells
which transform to male gametes (antherozoids).
• The division of body cell and the formation of sperm takes place just before
fertilization.
✓ The antherozoids of Cycas are uninucleate and multiflagellate, and the largest
in the plant kingdom.
• They are visible to naked eye.
(ii) Germination of megaspore and formation of female gametophyte
• Female gametophyte develops from the functional megaspore.
• This development is in situ, taking place within the nucellus.
• Megaspore has a thin and fibrillar inner wall called endospore, and a thick and
papillate outer wall called exospore.
• The functional megaspore increases in size and its nucleus divides repeatedly
and produces a large number of nuclei.
• Now, in the centre of the megaspore, a vacuole develops, pushing the
cytoplasm and the nuclei towards the periphery.

• Soon, cytoplasm collects around each nucleus and wall formation starts
centripetally from periphery to centre.

✓ Thus, the entire gametophytic tissue becomes cellular, each cell having a
single nucleus. The haploid tissue thus formed is called megagametocyte or
female prothallus, or endosperm.

✓ It develops without fertilization and that is why it is haploid.

• It differs from the triploid endosperm of angiosperms which is formed by

triple fusion.
Development of archegonia

• Archegonia develop from some cells at the micropylar end of the

gametophyte.

• These cells enlarge in size and become the archegonial initials.

• During this, their nucleus moves towards the periphery and the cells become
distinctly different from other endosperm cells.

• Now, the nucellar tissue above the archegonial initials disintegrates and
forms an archegonial chamber.

• Then, each archegonial initial divides periclinally and forms a small primary
neck cell on the outer side and a large central cell on the inner side.
• The neck cell divides anticlinally and forms a two celled archegonial neck.

• The central cell enlarges considerably and forms the venter. Its nucleus
divides and forms a small venter canal nucleus and a large egg nucleus.

• Neck canal cells are not formed.

• The egg of Cycas is the largest among the living plants.

• Soon, the venter gets surrounded by a nutritive jacket, formed by the

gametophytic cells. It is called archegonial jacket.

• The number of archegonia in an ovule is different among different species of

Cycas.
Fertilization
• Fertilization in Cycas is siphonogamous because of the formation of pollen
tube.
• Since the sperms move with the help of their flagella, it is also called
zoodiogamous.
• The pollen tube acts both as a haustorium and a sperm carrier.
• It is highly turgid since it is filled with a fluid of high osmotic potential. In
this fluid, sperms can actively swim about.
• During fertilization, the pollen tube grows towards the embryo sac through
the nucellus and reaches the archegonial chamber.
• Due to high turgor pressure of its fluid contents, it bursts open violently and
discharges its contents, including sperms to archegonial chamber.
• As soon as a swimming sperm comes in contact with the cells of the
archegonium, it is sucked in forcibly.

• Normally, only one sperm can get into the archegonium. It moves to the egg
nucleus.

• Nuclear fusion follows between the small sperm nucleus and the large egg
nucleus. This is called syngamy and it results in a diploid zygote or oospore.

• If more than one sperm happen to enter the archegonium accidently, the
condition is called polyspermy.

• In such cases, only one survives and the others get disorganized. This is
polyembryony (an abnormal condition).

• Zygote represents first cell or beginning of sporophytic generation.

Embryogeny (development of young sporophyte)
• Zygote is the mother cell of the sporophytic phase.
• It increases in size and undergoes repeated free nuclear division, forming
numerous daughter nuclei.
• Then, a central vacuole appears and the nuclei are pushed towards the
periphery.
• Centripetal wall formation follows from the periphery to the centre. This
results in a multicellular proembryo.
• Soon, it gets differentiated into three regions, namely the upper haustorial
region near the micropyle, the middle-suspensor zone, and the lower
embryonal region.
• In an ovule, several zygotes and proembryos may be formed.

• But, only one embryo attains maturity.

• The cells of the haustorial region absorb nutrients for the developing embryo.

• The cells of the suspensor region grow much faster and become elongated
and coiled.

• They push the embryo deep into female gametophyte.

• The embryonal region forms the main part of the embryo. The embryo takes
about one year for its complete development.

• In an embryo, two cotyledons are formed. The radicle is enclosed in a pad-

like hard covering, called coleorhiza, provides protection.
Structure of seed
• After fertilization, the ovule transforms to a seed.
• During this, nucellus is used up by the developing embryo.
• Mature seed is fleshy and red, orange, or dark-brown in color.
• It is surrounded by fleshy seed-coat or testa, formed from the integument of
the ovule.
• The outer fleshy layer of the integument forms the sarcotesta, and the middle
stony layer forms sclerotesta.
• The inner layer remains thin and papery.
• Seed-coat encloses the shrivelled nucellus, endosperm and a straight embryo
with two unequal cotyledons, plumule and radicle.
• Mature seed of Cycas represents three generations. Seed-coat (formed from
the integument of the ovule) and nucellus represent first sporophytic
generation, endosperm represents gametophytic generation, and embryo
represents second (new) sporophytic generation.
• The seeds of Cycas have attractive colouration, sweet taste and a pleasant
smell. These features are mainly responsible for zoochorous (mostly
ornithochorous) dispersal.