GYMNOSPERMS, TAXONOMY OF ANGIOSPERMS

AND ANATOMY

BLOCK-I-GYMNOSPERMS

Unit-1: Distribution, General Characteristics, Classification

and Economic Importance of Gymnosperms

By
Dr. Prabha Dhondiyal
Department of Botany
Uttarakhand Open University Haldwani
E-mail: pdhondiyal@uou.ac.in
CONTENTS

Introduction
General Characters of Gymnosperms
Classification
Distribution
Reproduction
Pollination and Fertilisation
Economic Importance of Gymnosperms
References
Suggested Readings
INTRODUCTION

The word Gymnosperm, “Gymnos” = necked and “Sperma” means seeds

was first used by Theophrastus, a pupil of Aristotle in his famous book
“Enquiry into Plants”. He used this term in all those plants having
unprotected (without covering) seeds.

On the basis of their seeds with or without covering are grouped into
two major categories namely- Angiosperms and Gymnosperms. Thus
the plants or Spermatophyta is divided into two sub groups
Angiosperms and Gymnosperms.

The ovules of gymnosperms have freely exposed before and after

fertilization while in case of angiosperms where (Angios = Vessels and
Sperma = Seeds) the ovules are enclosed within the carpel. Thus due to
this, the angiosperms are considered as the most advanced type of
organisms in plant kingdom. Comparatively to angiosperms, the
gymnosperms are less advanced; they have some specific characteristic
features as-
GENERAL CHARACTERS OF GYMNOSPERMS
❑ They are slow growers and lacks vegetative means of reproduction
such as by cuttings, layering etc.
❑ They are unable to grow under varied habitats means they are able to
grow on some specific habitats and conditions.
❑ They have limited means of dispersal of seeds and can be dispersed
only by wind, animals or by human beings.
❑ Most of the gymnosperms are terrestrial or land loving and unable to
grow in aquatic habitats except a few.
❑They lack vessels in xylem (with few exceptions) and companion cells
in phloem.
❑ Most of the gymnosperms are unisexual, thus due to absence of
bisexuality, chances of self- pollination reduces, and
❑As wind is the main source of pollination hence maximum amount of
pollen grains are wasted.
MORPHOLOGY OF VEGETATIVE ORGANS

The living gymnosperms including, tall trees, shrubs and climbers

includes approximately 70 genera and 725 species.

There are complete absence of herbs and climbers. Vegetative

propagation is not reported in this group but only bulbils are known in
some Cycas as means of vegetative propagation. Generally they possess
tap root but sometimes mycorrhizal and corolloid roots (Pinus and Cycas
respectively) are present in some genera.

Stem: The stem may be aerial, erect, unbranched (e.g. Cycas, Zamia) or
aerial, erect branched (e.g. Pinus, Cedrus etc.). In gymnosperms the
branches may be of two types on the basis of their branching system.

They may be either- two types as in Pinus- i) the long shoots and ii) the
dwarf shoots at their apices and collectively known as spurs.
Leaf: Gymnosperms bears both microphyllous and megaphyllous
leaves. The microphyllous leaves are large and well- developed and their
vascular supply always leaves a leaf- gap in the stem stele is another
characteristic feature of gymnosperms. The leaves may be simple or
compound and vary in shape, size and form, as a minute scale leaf to
several feet long megaphylls (e.g. in Cycas). Gymnosperms show great
diversity in leaf venation, it may be parallel (Welwitschia), reticulate
(Gnetum) or even dichotomous (Ginkgo). The leaves are always evergreen
and mostly possess resin canals as in Pinus, Cedrus and Abies. The leaves
of Gnetales lack resin passages but Gnetum possess latex tubes.

Leaf Arrangements: The arrangement of leaves may be whorled

(Cedrus), opposite or decussate (Gnetum, Ephedra etc.) or spirally arranged
(Taxus, Podocarpus etc.). Conifers usually have sunken stomata. The shape
of leaves may also vary from triangular (Pinus roxburghii), semi-circular
(Pinus sylvestris), bifid or circular (Pinus microphylla), and bifacial leaflet
of Cycas, Zamia, and Gnetum leaves). Likewise leaves the system and
arrangement of vascular bundles also shows great variations in
gymnosperms. In a young stem the ring of discrete vascular bundles. The
leaf base remains permanently merismatic while the tip drying off.
Vegetative organs (leaf) of some gymnosperms
Due to secondary growth gymnosperms possess primary and secondary
wood. The secondary wood is the characteristic feature of cycadophyta.
The wood is porous, soft and more parenchymatous in nature, while
pycnoxylic wood is the characteristic feature of coniferophyta; the wood
is compact and narrow medullary rays, xylem lacks wood vessels except
in Ephedra, Gnetum etc. The xylem is usually endarch or mesarch in stem
while it may be exarch in roots. The vascular bundles are conjoint,
collateral, endarch and open in gymnosperms.
CLASSIFICATION

The group Gymnosperms is a very large class which includes both living
and fossil forms. Due to ample records of fossil forms the classification
has become somewhat complicated. Several workers have classified
Gymnosperms differently from time to time among them the important
ones are as follows.
The pioneer workers in this field are Coulter and Chamerlain (1917)
divided the gymnosperms directly into seven orders viz.
1. Cycadofilicals,
2. Bennettitales,
3. Cycadales,
4. Cordaitales,
5. Ginkoalea,
6. Coniferales and
7. Gnetales.
Jefferey (1917) recognizes two classes among gymnosperms s follows-
A. Class Arachigymnospermae- It includes all those gymnosperms
that resemble from their general appearance and anatomy. They
possess motile spermatozoids. He divided this class into five
orders-

1 Order Pteridospermae- it includes all extinct forms, primitive and that

lived in Paleozoic period.
2. Order Cycadeodea- It includes all extinct forms.
3. Cycadales- Includes extinct and living forms.
4. Order Cordaitales- Includes all extinct forms.
5. Order Ginkgoales- Includes extinct and single living genera.
B. Metagymnospermae -i) plants of this class possess simple leaves and
have no resemblance to the ferns.
ii) Possess non- motile male gametes, the pollen
gains grow into a pollen tube. It includes two orders-
1 Order Coniferales- It includes living genera e.g. Pinus, Cedrus, Taxus,
Podocarpus, Taxodium etc. also includes fossil genera.
2 Order Gnetales- Includes Ephedra, Gnetum and Welwitschia and many
fossil genera.
On the basis of composition of wood the gymnosperms were divided
into two classes-
i) Metaxylic- when possess porous wood and loose texture.
ii) Pycnoxylic- When the wood was compact
The former includes Cycadales, Cycadeiodales and Cycadofilicales
while the latter includes Cordaitales, Ginkgoales, Coniferales and
Gnetales. This classification was given by Soward (1919).
However, Chamberlain in 1934 divided the gymnosperms into
two classes which were further divided into orders with their respective
characters such as-
A. Class Cycadophyta- i) Most of the plants of this group are unbranched,
stems are stumpy.
ii) Male cones large and compact with simple sporophylls bear large ovule.
iii) Anatomically the stems have wide cortex and monoxylic wood.
This class includes three orders-
a) Cycadofilicales- Includes extinct forms and fossils.
b) Cycadeiodales-Includes both living and fossil forms.
c) Cycadales- Includes both living and fossil forms.

B. Class- Coniferophyta- i) profusely branched stem is the characteristic

feature of this group.
ii) Leaves simple and the foliage given leaf like appearance.
iii) Both the strobili- male and female are compact and bear complex
sporophylls.
iv) Wood is pycnoxylic.
It includes four orders-
a. Cordaitales- It Includes extinct order.
b. Ginkgoales- It includes extinct and only one living representative- Ginkgo
biloba.
c. Coniferales- It includes both extinct and living forms.
d. Gnetales- Includes both living and extinct forms.
D. D. Pant (1957) has proposed a classification of Gymnosperms in which
the group is divided into three divisions
a) Cycadophyta
ii) Clamydospermatophyta and
iii) Coniferophyta
An outline of Pant’s (1957) scheme of classification of gymnosperms is as
follows:
After that Andrew (1961) another renowned scientist classified
Gymnosperms and divided it into six divisions as-
i) Pteridospermatophyta
ii) Cycadophyta.
iii) Ginkgophyta
iv) Coniferophyta.
v) Gnetophyta and
vi)Gymnosperms of uncertain affinities.
K. R. Sporne in 1965 classified Gymnosperms in his book namely “The
Morphology of Gymnosperms” based on Pilger and Melchior (1954)
classification-
Gymnosperms- divided into three divisions
1. Cycadopsida 2. Coniferopsida and 3. Gnetopsida
These divisions further divided into orders-
1. Division Cycadopsida-
Order 1. Pteridospermales- Families-(7)-1. Lyginopteridaceae-
(Lyginopteris)
2- Medulosaceae (Medulosa)
3- Calamopteridaceae (Calamopitys)
4- Glossopteridaceae (Glossopteris)
5- Peltospermaceae (Xylopteris)
6- Corystospermaceae (Xylopteris)
7-Caytoniaceae (Caytonia)
.
Order 2. Bennettitales - Families (3) 1- Williamsoniaceae (Williamsonia)
2. Wielandiellaceae (Wielandiella)
3. Cycadeoideaceae (Cycadeidea)

Order 3. Pentoxylaes -Family(1) 1. Pentoxylaceae (Pentoxylon), Sahnia)

Order 4. Cycadales-Family(2) 1. Cycadaceae (Cycas, Zamia etc.)
2. Nilssoniaceae (Nilssonia)
2. Division Coniferopsida
Order 1. Cordaitales –Families (3) 1. Ertophytaceae (Eristophyton)
2. Cordaitaceae (Cordaites)
3. Poroxylaceae (Poroxylon)

Order 2. Coniferales –Families (9) 1. Lebachiaceae (Lebachia)

2. Votziaceae (Voltziopsis)
3. Palissyaceae (Palissya)
4. Pinaceae (Pinus, Abies, Picea)
5. Taxodiaceae (Taxodium)
6. Cupressaceae (Cupressus)
7. Podocarpaceae (Podocarpus)
 8. Cephalotaxaceae (Cephalotaxus)
9. Araucariaceae (Araucaria, Agathis)
Order 3. Taxales- Family (1) Taxaceae (Taxus, Torreya)
Order 4. Ginkgoales.-Families (2) 1. Trichoptyaceae (Trichopitys)
2. Ginkgoaceae (Ginkgo)
3.Division Gnetopsida
Order 1. Gnetales.- Families (3) 1. Gnetaceae (Gnetum)
2. Welwitschiaceae (Welwitschia)
3. Ephedraceae (Ephedra)
Recently (1980) Taylor classified Gymnosperms in six divisions as
1. Progymnospermophyta,
2. Pteridospermophyta.
3. Cycadophyta.
4. Cycadophyta.
5. Ginkgophyta.
6. Coniferophyta.
Thus after reviewing the different theories and outlines of
classification of Gymnosperms adopted by different works from time
to time it shows that there are great controversies regarding the
classification of Gymnosperms. But even then the last and the most
important classification may be taken as correct one for studies.

Stewart (1983) placed Progymnosperpsida,

Gymnospermopsida and Gnetopsida as distinct classes under the
division Tracheophyta, the vascular plants of kingdom Plantae. These
three classes were further divided as-
Kingdom Plantae-
Division Tracheophyta
Class - 1. Progymnospermopsida
Order - A. Aneunophytales
B. Aracheopteridales
C. Protopityles
Class 2. Gymnospermopsida
Order - A. Pteridospermales G. Gimkgoales
B. Cycadales H. Cordaitales
C. Cytoniales I. Voltziales
D. Glossopteridales K. Coniferales
E. Pentoxylales L. Taxales
F. Gzekanowskiales
Class 3. Gnetopsida
Order A. Gnetales
B. Ephedrales
C. Welwitschiales.
Birbal Sahni (1920 a), based on morphological nature of ovule bearing
organ and axial or foliar nature of ovules divided gymnosperms into two
major groups- Stachyspermae ) ovules arise on the axial organ or stem)
spread over orders- Cordaitales, Ginkgoales, Coniferales, Taxales and
Cycadales and Cycadofilicales as Phyllospermae (ovules borne on
leaves).
Gymnospermae-
1. Phyllospermae 2. Stachyspermae
A. Cycadofilicales A. Cordaitales
B. Bennettitales B. Ginkgoales
C. Cycadales and C. Coniferales.
D. Taxales.
Christenhusz et al., (2011) proposed a new classification and linear
sequence of the extant gymnosperms based on molecular and
morphological phylogenetic studies. They divided all extant
gymnosperms into 4 sub classes, 8 orders and 12 families as follows-

Sub Class I. Cycadidae Order A. Cycadales

Family 1. Cycadaceae Family 2. Zamiaceae
Sub Class II Ginkgoidae Order B. Ginkgoales
Family 3. Ginkgoaceae
Sub Class III Gnetidae Order C. Welwitschiales
Family 4. Welwitschiaceae
Order D. Gnetales
Family 5. Gnetaceae
Order E. Ephedrales
Family 6. Ephedraceae
Sub class IV Pinidae Order F. Pinales
Family 7. Pinaceae
Order G. Araucariales
Family8. Araucariaceae
Family9. Podocarpaceae
Order H. Cupressales
Family10. Sciadopityaceae
Family11.Cupreassaceae
Family12. Taxaceae.
DISTRIBUTION

Certain groups of gymnosperms are entirely extinct, while others are

present in living as well as in fossil forms with primitive features. Still
there are some groups chiefly within living gymnosperms that extend
throughout the temperate, tropical and even in arctic zones. Most of the
gymnosperms are evergreen xerophytes.

The total number of living gymnosperms in the world is approximately

number seventy genera and 725 species. A total of 16 genera and 53
species were reported from India (M. B. Raizada and K. C. Sahni, 1960).

Maheshwari listed only 14 genera. This lesser number of representatives

is mainly due to their habitat as gymnosperms are mainly dwellers of
temperate regions in India and such climate was afforded only by the
Himalayas, They form extensive forests and grow luxuriantly in the
various Himalayan ranges that is why most of the gymnosperms are
distributed in eastern and western Himalayas besides in some other
regions of India.
There are six living orders of gymnosperms and out of them four are
represented in India. These include- Cycadales, Coniferales, Ephedrales
and Gnetales. The Cycadales are represented by 4 species of Cycas in
India.

Gymnosperms are very poorly represented in the Indian flora. In the vast
peninsular India they are represented by a few species of Cycas,
Podocarpus, and Gnetum. However, in the extra peninsular Himalayas and
in some extent in the connected ranges of Kashmir, Assam, and
Arunachal Pradesh gymnosperms represented only by conifers and
covering extensive tract of forest land.

Among gymnosperms, different groups of this category i.e.

Cycads, Conifers, Ginkgoales, and Gnetales the most frequently and
densely populated group is the Coniferales. This group as we know is
represented by Pinus, Cedrus, Abies, Larix, Picea, Cupressus, Tsuga,
Juniperous, Taxus, Araucaria, Thuja, Podocarpus, Cephalotaxus are common.

Conifers are found predominantly in the Himalayas and are particularly

rich in the north-west Himalayas (Uttarakhand, Kashmir, Himachal
Pradesh etc.).
Their distribution is generally governed by altitude and generally ranges
from 1800- 3300 meters asl. While some species of Pinus (P. insularis 700-
1,850 m asl and P. merkusii (150- 600 m asl.) are reported from Khasya
region of Assam and on the hillocks in East Bengal respectively).

Among Cycads only Cycas occurs in India and the genus is represented
by four species viz. C. circinalis, c. beddomei, C. pectinata and C. rumphii,
beside this another species C. revolute, which is a native of Japan is
commonly cultivated in Indian gardens.

Species of Zamia, Macrozamia, Encephalortos and Stangeria are exotics and

occasionally cultivated in Indian gardens. Similarly a few plants of Ginkgo
biloba, a native of China, occur in Indian under cultivation in gardens.

Gnetales are represented in India by a number of species of Ephedra and

Gnetum. Out of seven species of Ephedra only one (E. foliate) occurs in the
plains of Rajasthan and Punjab while rest six are confined to the north-
west Himalayan regions.
REPRODUCTION
Gymnosperms possess two different types of spores and hence refers as
hetrosporous. The microspores are smaller while another spore larger in
size called megaspore. These two kinds of spores on germination
produce two different kinds of gametophytes. The microspore or pollen
grains produce male gametophyte, while the larger megaspore produces
female gametophyte, bears two or more archegonia or female sex
organs. These spores are produced within the leafy structures or
sporangia that borne on sporophylls, spirally arranged along an axis to
form compact strobili or cones.
The microsporangiate or male strobili bearing microsporophyll
and microsporangia while the megasporangiate or female strobili bear
megasporophylls with ovules or microsporangia. The two types of cones
or strobili may be borne on same tree as in Pinus or on different trees like
in Cycas and Ginkgo. The microsporangium contains numerous small
microspores whereas the megasporangium contains only one larger
megaspore. Both the spores i.e. microspore and megaspores are haploid
and develop as a result of meiosis or reduction division in the respective
spore mother cells. They are the primary structures of the male and
female gametophytes respectively.
Male and female cones/strolili of Cycas(A,B), Ginkgo(C,D), Pinus (E,F), Taxus
(G,H), Ephedra (I,J), Gnetum (K,L) and Welwitschia(M,N)
In gymnosperms the gametophytes are endosporic i.e. they develop
within or inside the respective spore wall. In general the strobili or
cones are of varying shapes and sizes in different species. Their
position also varies from plant to plant. Among gymnosperms, the
microsporangiate or male cones are largest and arise singly at the apex
of male plant.

Female gametophyte or ovule: As per discoveries the ovules of

gymnosperms are without any covering or naked and are borne on
usually spirally arranged megasporophylls around a central axis. The
ovules are generally sessile. Among gymnosperms, ovules of Cycas are
the largest among the plant kingdom. In a ovule there is a
megasporangium or nuecllus encloses in a parenchymatous mass of
cells. The nuecllus encloses a single diploid megaspore mother cell that
undergoes meiosis and formed 4 haploid mother cells arranged in
linear tetrad form. Out of these 4 only one, usually the lower one
remains functional and the rest ones degenerates. The functional
megaspore enlarges and undergoes free nuclear division and resulted
into large number of free nuclei. This transforms into young
gametophyte that has developed within the megaspore.
Ovules is some gymnosperms
The nucellus now covered by a single massive layer or integument
grow around and leaving a small pore at one end known as micropylar
end. Thus the new megasporangium integumented is called ovule.
Now around each free nuclei a centripetal wall formation starts and it
continues till the whole female gametophyte becomes cellular.

In gymnosperms, a single ovule consists of one middle stony layer

covered by inner and outer fleshy layer. The apical region of the
nucellus forms a pollen chamber by degeneration consisting the semi-
germinated pollen grains or microspores. These megaspores remain in
the chamber till further growth, towards the micropylar end the female
prothallus develops two or more archegonia. Depending upon species
the archegonia have a short or long neck made up of 2, 4, and 8 cells, in
Cycas, Taxus and Biota respectively. This develops into megaspore
mother cell which further undergoes free nuclear divisions.
Afterwards nuclear divisions the cell wall formation starts as a result
of which the female gametophyte become a cellular structure. The
female gametophyte or prothallus get differentiated into upper
reproductive region, middle storage region and the lower basal
haustorial region.
 Ultimately most of the cells abort and only one remain and matures.
Generally in gymnosperms the female gametophyte has cellular tissue at
its lower end while on its upper end a few free nuclei remains without
wall formation and acts as eggs. After fertilization this apex end
however, becomes cellular (exceptional in Welwitschia).

Microspores: In gymnosperms the pollen grains or microspores are

unicellular and haploid structures. They differ in shape and sizes in
different groups of gymnosperms. They may be tetrahedral with a
definite polarity due to the thicker exine towards the base (e.g. Cycas) and
uniapertuate, may be almost spherical and uniapertuate (Ginkgo), may
be winged (saccate), uniaperturate with reticulate exine (Abies pindrow,
Cedrus deodara, Pinus roxburghii and P. wallichiana, Picea smithiana etc.)
whereas in Ephedra the pollen grains are inapertulate elongate with
palcate exine surface. E. foliate showing parallel ridges along the long axis
of grains the pollen grains possess two tiny (sac like) structures the
extremities however, in other species sacs are almost absent. (Figs. ovules
of gymnosperms, archegonia of gymnosperms). Based on studies it is
suggested there is a gradual reduction in the wings or sacs and
ultimately resulted in non-winged pollen grains as reported in Ephedra.
Pollen grains in gymnosperms
Male Gametophyte: The male gametophytes in gymnosperms are
endosporic in nature and show variations regarding their release from
sporangia, the number of male prothallial cells (they complete their
development partly in the microsporangium and partly in the pollen
chamber of the ovule), size and motility of male gametes and their time of
formation and discharge. In different groups of gymnosperms differs in
their morphology. In lower gymnosperms- cycadaceous micro
gametophytes there is one male prothallus cell that divides into a large
sterile cell or stalk cell and a body cell or a spermatogenous cell and a
tube nucleus. They are arranged in a linear row. The body cell again
divides into two multiciliate male gametes. In all Cycads the pollen tube
is formed and is more haustorial in nature than a sperm carrier. While in
Microcycas the stalk cell divides into 10 or 11 body cells or
spermatogenous cells. These all cells divide to produce 20 or 22
spermatozoids. Contrary to this in Ceratozamia there are 4 spermatozoids.
Generally in Gymnosperms the generative cell divides into stalk cell and
body cell except in Cycas revolute, where it is divided anticlinally.
In case of Coniferales, different families of this group show slight
variations in the sequence of micro- gametophyte development, although
the major event shows similarities.
 In Pinaceae family the nucleus of microspore divide twice by
periclinal walls and cut off three cells- two male prothallus cells and one
antheridial cell. The latter divides periclinally and forms a tube cell and a
generative cell. The grains are liberated at this four celled stage (two
prothallus cells, one tube cell and one generative cell). Again the
generative cell undergoes periclinal division and forms a stalk cell and a
body cell. All these cells lie in an axial row. The semi germinated pollen
grains may germinate immediately or after a month. The pollen tube may
start branching on entering the nuecllus, then the tube nucleus migrates
into one of the branches of pollen grain and moves to the tip, while the
generative cell remain within the spore wall and divides into stalk cell
and body cell between the pollination period of the year or in the
following spring, these two moves into the pollen tube. Simultaneously
about a week before fertilization, the body cell divides into two non-
motile, unequal size male gametes. There is difference in opinion
regarding the male gamete. According to some workers the two male
gametes are as two nuclei of binucleate sperm cells, whereas others
regard them as two sperm cells. Sometimes the generative cell divides
before pollination, in some genera e.g. Abies, Cedrus, Picea, and Larix etc.
In Abies the cell sometimes divides into two male gametes before
pollination or shortly after pollination.
However, Chawdhury (1960) have been reported equal male gametes in
Cedrus. The period between pollination and fertilization varies in different
members of Pinaceae. It is longest in Pinus ranges up to one year, up to
nine months in Cedrus, 4-5 weeks in Abies and a few days in Picea.
Similarly the number of prothallial cells also varies in different species of
gymnosperms.
In Abies balsamea formation of 3 or 4 prothallial cells have been reported
by Hitchinson (1915). While in Taxodiaceae the microgametes lack
prothallial cell and divides into a generative cell and a tube cell before
pollination. Further the generative cell divides into stalk cell and body cell
after pollination. In family Cuperessaceae too prothallial cells are absent,
here the pollen grains directly acts as antheridial cell and may divide into
generative cell and tube cell before pollination. Based on studies, in
Cupressus semipervirens the body cells divides into 4- 20 male gametes,
contrary to this in C. fusiberus only two male gametes are produced.
Among gymnosperms, the Araucariaceae are unique in producing many
prothallial cells the two non- motile male gametes are equal in size. In the
members of Podocarpaceae the number of male prothallial cells may be 1-
8 in Podocarpus, 3-6 in Dacrydium and 1-3 in Phyllocladus. While in
Pherosphaera the male prothallial cell is absent.
In Cephalotaxaceae there is no prothallial cell in any species of
Cephalotaxsus. Whereas in Taxus there are no male prothallial cells. The
male gametes are unequal in size. In ephedra the cells of male
gametophyte arranged in a single axial row including two prothallial cells,
a stalk cell, a body cell and a tube cell. Out of the two prothallial cells one
is without a cell wall and is thus called prothallial nucleus. After
pollination the body cell divides into two male gametes. Contrary to this in
Gnetum there is only one prothallial cell, a tube nucleus and a generative
nucleus. Further division takes place after pollination while at that time
the pollen grain contains only three nuclei. The generative nucleus divides
into two male gametes. In Welwitschia there is no prothallial cell and the
released grain contains a tube nucleus and a generative nucleus. Based on
above discussion on the general characteristics of male gametophyte in
gymnosperms following points emerge

1) The Cycadales, Ginkgoales, Coniferales and Ephedra among Gnetales all

have the same cellular cell organization of the micro gametophytes
including the tube cell, the stalk cell and the male gamete except one
difference for the formation of prothallial cell.
2) In case of Araucariaceae and Podocarpaceae formation of a very large
number of prothallial cells they developed secondarily at the base of the
microspore, may dealt as a later evolutionary development. This
condition is not reported in any living or fossil gymnosperms or nor
among lower heterotropous tracheophyes.

3) In another specific ontogenic feature is the orientation of the division of

spermatogenous cell which produces the stalk cell and the body cell, (e. g.
Ginkgo), Auricuriaceae, Podocapeceae and Cycas revoluta and in some
species of Ephedra. The division wall is anticlinal while in other Cycads
and Pinaceae the division wall is periclinal and the two cells are
superimposed.

4) Beside this in Microcycas the stalk cell which is sterile divides and gives
rise to additional spermatogenous cell or body cell.

5) While in some other gymnosperms i.e., Cupressus and sometimes in

Juniperous more than two male gametes are producedThis feature may
have related with the presence of large number of archegonial complex.
POLLINATION AND FERTILIZATION
In gymnosperms the medium of pollination is wind; it results in the
transfer of semi-germinated pollen grains on the micropyle of the ovule.
In most of the gymnosperms, the pollen grains are caught into a
pollination drop selected by micropylar end of the ovule. After drying of
pollination drop the microspores in semi- germinated stage are drown
into the ovule. Just after the drawing of the microspores in the micro
gametophyte the micropyle closes. A distinct pollen chamber is formed at
the apex of the nuecllus and receives the micro- gametophytes on
pollination as in some gymnosperms (Cycas, Ginkgo, Ephedra etc.)

While in Conifers and other gymnosperms the semi- germinated pollen

grains come in direct contact with the nuecllus beak. Contrary to this in
the palaeozoic gymnosperms now extinct the pollen chambers contained
liquid filled cavities in which the motile sperms were liberated due to the
dehiscence of the microspore wall. In general in all the living
gymnosperms, the microspores by division produce pollen tube as a
tubular outgrowth that grows through the nucellar tissue. In Cycads and
Ginkgo the pollen tubes mainly acts as the haustorial organs and grows
for long time (several months) into the nucellar tissue and absorb food
and supply it to the micro gametophyte at the grain end or at the lower
end of the pollen tube. The pollen tube bursts during fertilization and
liberation of multiciliate male gametes along with some liquid in the
cavity above the mega gametophyte. At that time the sperms swims to
the archegonial neck and enters into the archegonia and only one of
them fuses with the egg or oosphere and form the diploid zygote or
oospore.
In the conifers the pollen tube plays an important role of sperm
carrier. The male gamete along with stalk and tube nucleus migrate to
tip of the pollen tube. The tube grows through the nucellar tissue,
reaches to the archegonial tube enters through it and after bursting
liberates the male gametes. Among them, one fuses with the egg to form
a diploid zygote. This specific process of fertilization is termed as
Siponogamous.
However, in Cycadofilicales, Bennettitales and Cordaitales, the
extinct orders of gymnosperms did not produce pollen tubes and
sperms were liberated directly into the pollen chamber. This process is
known as Zooidogamous.
In case of Welwitschia the female gametophyte gives out tubular
prolongation that meets the pollen tips fertilization takes place after
the intervening wall dissolves. In Welwitschia and Gnetum there are no
archegonia.

EMBRYOLOGY: In different groups of gymnosperms embryogeny

differs in different stages. It also differs in living and fossil forms. In
living or present day gymnosperms the first phase in embryo
development is the free nuclear divisions except in Gnetum,
Welwitschia and Sequoia semipervirens. While it is completely absent in
angiosperms and other tracheophyta.

Just after free nuclear division, wall formation begins and the embryo
transformed into cellular form. Later it differentiated into a suspensor,
radicle, hypocotyl, plumule and cotyledons. When the shoot end of the
embryo is directed away from the micropylar end of the ovule, that
type of embryo development or embryogeny is called “Endosporic”.
Polyembryony is the characteristic and significant feature of
gymnosperms. This is possible as more than one archgonia are fertilized
and so more than one zygote are formed. These zygotes later developed
into embryos, but one of them succeeded in developing into a complete
embryo. Comparatively to Cycades, in Conifers there is a “Cleavage
Polyembryony”. As reported earlier that in conifers only four nuclei
formed, so in this case all the four cells of the young embryo separates
after wall formation and develop into 4 embryos, but only one completes
further development while others abort.
In Thuja no cleavage Polyembryony reported and only one
embryonial initial develops into an embryo. No free nuclear division is
reported in Sequoia semipervirens and the zygote divided first by a
transverse wall and forming two cells which divided further by other
longitudinal walls to form four cells. All these cells may function as
embryo initial and give rise to filamentous embryo.
In majority of conifers the zygote develops four free nuclei and
later because of further divisions and wall formation results in forming a
“Proembryo” with a four celled distal embryonal tier, middle suspensor
tier and upper rosette tier (Pinus, Cycas, Tsuga etc.)
While in other gymnosperms as in Abies, Picea and Larix the rosette tier
disappears at a later stage. In further developmental stages the lower
embryo tier develops into all the organs at a later stage. The suspensor
tier develops into additional embryos. While the upper most tier ends
are in open contact with the egg cytoplasm. This end is apparently active
in transmission of nutrients to the growing embryos.
The number of cotyledons varies in different species end even
in the same species. It is 10 in Pinus roxburghii, P. banksiana- 3-6, P.
contorta- 2-8, P. sabiniana-7-8.
Cleavage Polyembryony is not reported in Araucaria angustifolia
the member of Araucariaceae. In this species 32-45 free nuclei are
formed. Later by polar elongation wall formation is accompanied. The
cells at the distal end develop into enlarged cap cells. The central cells
develop into embryo and those situated towards the micropylar end
give rise to suspensor. There is complete absence of “Cleavage
Polyembryony”.
In case of Cuperessaceae the genus Actinostrobus shows
Cleavage Polyembryony and show somewhat different pattern of
development. Here 4 free nuclei are formed. Wall formation results in
forming 4 cells among which two vertically placed cells below
archegonial neck and two transversally placed cells below.
The lower cells divide once to form four embryonic initials while the
upper cells do not divide further. The lower ones are polarized in a
transverse plane and each of them forms a small distal initial cell and a
large suspensor cell and forms four small embryonal cells and four
large suspensor cells that elongate considerably.
Hence show cleavage polyembryony. The rosette tier and upper tier is
not formed in this case. The embryo has two cotyledons. The embryonal
cells are binucleate in the Podocarpaceae. While the number of
binucleate cells and the pro-suspensor cell varies with species and
genera. The embryo has two cotyledons.
Development of embryo in Gnetales is different than above mentioned
cases. Among Gnetales in Ephedra trifurea the zygote nucleus divides
into eight free nuclei of unequal size distributed unevenly in the
protoplasm. Three to five of these nuclei become enclosed individually
by irregular wall which become globular later. All these globular cells is
pro- embryo or is an example of proembryony. Later the globular pro-
embryo develops into an embryo with the massive embryonal mass at
the distal end and a number of suspensors. In Ephedra foliate
exceptionally there is only one suspensor tube.
 In Welwitschia embryo developed differently in this case the
zygote elongates and divides into different parts- an upper elongated
primary suspensor and a distal embryonal cell, later it divides to form an
apical wall. The outer mass of this apical mass are called inner cortical
ring and are situated adjacent to the primary suspensor. The outer
cortical ring is formed later containing elongate around the suspensor.
Thus the primary suspensor cell is now surrounded by two layers of 8- 16
cells. Later by division of these cells more layers are added around these.
At the distal end the cells form cap cells while those in the middle
develop into embryo.

Comparatively to this embryo development in Gnetum is quite different.

In this case free nuclear division is not found and the zygote divides and
develops into a two celled body that gives out elongated and tube like
suspensors. These suspensor cells may branch and all have a distal,
densely granulose embryonal cell. Further these terminal cells develop
into an embryo out of which only one reaches maturity. In Gnetum
embryo development is completed after the detachment of the seed.
 On the basis of comparative studies of embryonal development
in gymnosperm, it is concluded that there is no close relationship or
resemblances between the embryogeny of pteridophytes, gymnosperms
and angiosperms. This suggests parallel evolution among these groups
rather than evolved from a common ancestor. However, the
gymnosperms embryo share some common features with other
embryos-
1. Axial development of embryo.
2. Early determination of polarity.
3. A conspicuous meristematic distal pole.
These similarities point out the possibility of a common ancestry
of gymnosperms with other vascular plants.
Embryogeny of Sequoia semipervirens (A-E); Proembryos of Araucaria
angustifolia (F-G); Early embryogeny of Actinostrobus pyramidalis (H-K).
SEED FORMATION: After fertilization, the structure developing from
fertilized ovule and its consequent enlargement is known as seed. The
zygote develops into an embryo while the endosperm persist as an
nutritive tissue, whereas the nuecllus becomes disorganize ( or serves as
nurse cells for developing embryo) or it may remain in the form of dry
tissue at the micropylar end of the seed known as nucellar cup.

In gymnosperms the inner fleshy layer called the tegmen, may persist as
a thin layer of seed coat. The middle stony layer later changes into a
hard layer called the testa, which mechanically protected the female
gametophyte and the embryo. Development of seed may vary in
different species of gymnosperms. In Cycas and Taxus the outer fleshy
layer develops into scarlet red and fleshy outermost seed coat.

In Gnetum the seed develops before the embryo complete its

development. In Taxus a fleshy aril develops from the basal cup- shaped
structure. Except Cycas and Ginkgo, the seeds of all gymnosperms remain
dormant for some time. While in these two genera the seeds germinate
immediately, they lose their viability when fall on moist substratum.
In gymnosperms the seed represents
two sporophytic and one gametophytic
generation. Different parts of a seed of
gymnosperms represents different
generations.
1) The young embryo represents the
new sporophytic generation.
2 The seed coat represents the old
sporophytic generation and
3) The endosperm represents the
gametophytic generation.
In most of the genera of
gymnosperm the germination of seed is
epigeal means the cotyledons come
above ground except in Ginkgo where
the cotyledons remain embedded in the
endosperms. While in Ephedra trifurcate
Vivipary has been reported. All the
gymnosperms represent heterogenous
alternation of generation.
ECONOMIC INPORTANCE OF GYMNOSPERMS
Gymnosperms are of great economic importance in nature and have
many economic importance for human beings. It gives valuable wood,
resin, essential oils, gums, paper, turpentine, medicines, food,
ornamentals and miscellaneous items.
Gymnosperms are frequently used in parks, gardens because of their
evergreen habit and symmetrical appearance. The trees are used for
timber, building construction, resin, paper manufacturing etc. They are
also used in medicines, perfumes, varnishes, paints and essential oils.
While roasted seeds of Ginkgo are eaten at feast in China and Japan, to
promote digestion and diminishes the effect of drinking wine. Seeds of
Pinus gerardiana (chilgoja) used as dry fruit. Seed kernels of Gnetum ula
yield an oil for illumination and massage in rheumatism. Bark of Taxus
baccata is used as main ingredient of famous Bhatia tea.

FOOD: In some parts of India, Malaya, Philippines and Indonesia,

young succulent leaves of various species of Cycas are cooked and eaten
as vegetable. The famous “Sago” starch is obtained from the stem /
seeds of Cycas and used as food. This stem starch obtained from
Macrozamia spiratis is an important source of food for poultry, dairy
animals and pigs. The seeds of Cycas are used as paste and eaten as
cakes in Nicobar Island.
GREEN MANURE: Leaves of Cycas are rich in nitrogen and used as
green manure for rice, sweet potato and sugarcane.
MEDICINE: Leaf extract of Ginkgo biloba is useful in the treatment of
cerebral insufficiency and vertigo.
ORNAMENTAL: Ginkgo biloba and Cycas species are grown as an
avenue tree and in gardens also for beautification. These trees are
preferred especially due to their slow growth, evergreen nature and
beautiful symmetry.
TIMBER: Conifers and Taxales are most important genera of
gymnosperms significantly important to produce high quality, straight
grained, light colored, high weight and strong wood in comparison to
their weight. They are suitable for making cabinets and furniture due
to their strength and durability. The wood of Abies is light and termite
free. It also has pleasant scent smell and used for packing cases, match
wood, wood wool, aircraft work, plywood, light camp furniture and
also used as household materials. Juniperus wood is fragrant, reddish
brown and rarely damaged by insects. Cedrus wood is also durable,
oily, fragrant, insect repellant and rot resistant.
The wood of Taxus is strong, oily, elastic, close- grained, fragrant and
very durable with smooth glossy surface. Beside this, wood of Araucaria
canninghana used for plywood manufacture.
RESIN: Conifers exudated resins, this help the wood resistant to decay.
Conifers are the major resin yielders of the world. These resins evaporate
their oil and became harder which makes them invaluable in paints,
varnishes, paper sizing, medicines and liquors industries.
CANADA BALSAM: A resin obtained from Abies balsamea which has a
very high refractory index approximately that of glass. Due to this
property it is extremely suitable as amounting medium for microscopic
objects and as cements for uses in optical work.
ESSENTIAL OILS: All conifers young branches and adherent leaves
provide essential oils. Himalayan Cedar oil (Cedrus deodara) and Red
Cedar Wood (Juniperus virginiana) are used cleaning tissues in histological
work and also use with the oil immersion lens of the microscope. The oil
obtained from Cedrus atlantica possess medicinal properties and used
against bronchitis, tuberculosis, skin diseases and gonorrhea. The
essential oils are used extensively in preparation of deodorants, room
sprays, disinfectants, perfumery and medicine etc.
FATTY OILS: Many conifer seeds are rich in fatty oils. The oil from
the seeds of Pinus cembra and Torreya nucifera is edible and also
used for paints. The Tail Oil obtained as a by product from
sulphate process of cooking conifer wood for making Kraft paper is
used in paints, soaps, linoleum, emulsifiers etc.
PHARMACEUTICALS: The leaves of Taxus baccata are used in
asthma, bronchitis, hiccough, epilepsy and for indigestion. Taxol
(from Taxus brevifolia) is found effective against ovarian cancer,
breast cancer, and melanoma and colon cancer. Ephedra is the
source of a valuable drug Ephedrine obtained from E. equisetina, E.
gerardiana, E. major, E. sinica, E. intermedia and E. nebrodensis. It is
used against cold, respiratory disorder and hay fever. An aromatic
beverage, known as Mormon tea is also brewed from the species of
Ephedra in south western United State.
AMBER: It is a fossil, water insoluble tree resin which was secreted
by the now extinct pine, (P. succinifera). It is yellow, brown to black,
hard and brittle with an aromatic odor.
GLOSSARY
1. AMBER: It is a fossil, water insoluble tree resin, secreted by now
extinct pine (P. succinifera), yellow, brown and black in color and hard,
brittle with aromatic odor.
2. ARIL: Fleshy and hairy outgrowth of seed or fertilized ovule regarded
as modified outer integument.
3. EPIGEAL: Describing germination of seed where cotyledons are
raised above the ground surface by considerable elongation of
hypocotyl.
4. EMBRYO: Young individual formed after fertilization.
5. ENDOSPORIC: Gametophyte describing a gametophyte that
develops within the spore.
6. GAMETOPHYTE: vegetative structure representing gamete
producing generation in the life cycle of plant. It arises from germination
of haploid spore.
7. GENERATIVE CELL: One of the cells found in pollen tube of seed
plants. In gymnosperms it gives rise to a body cell and a stalk cell.
8. GREEN MANURE: Any fast growing, inexpensive crop sown
towards the end of season to be ploughed into the soil after a while. It is
green so that it may increase soil manure on decomposition.
9. GUM: Any substance that swells in water to form gels or sticky
solutions.
10. HETROMORPHIC ALTERNATION OF GENERATION: Type of
alternation of generation where morphologically different
gametophytic and sporophytic generations alternate with each other to
complete the sexual life cycle of organism.
11. HETEROSPORY: Plants producing different types of spores by a
species.
12. HYPOCOTYL: Region of stem that is derived from part of the
embryo lying between cotyledons and radicle.
13. HYPOGEAL: Type of seed germination in which cotyledons
remain buried in the ground even after germination.
14. LIVING FOSSIL: Any present day species that resembles some
characteristics to extinct organism (only Ginkgo biloba).
15. OVARY: The swollen, rounded basal portion of angiospermic
carpel that contains ovule.
16. POLLEN CHAMBER:A cavity at the micropylar end of nucellus in
some gymnosperms into which pollen grains settle after the
pollination and mature there prior to germination.
17. PRO- EMBRYO: Term referring to young plant individual formed
after fertilization not before its differentiation into embryo and
suspensor tissue.
18. POLYEMRYONY: Condition of formation of more than one embryo
in one ovule which may develop by division of one fertilized zygote
(cleavage polyembryony) or may develop asexually from somatic tissue
along with sexual embryo (adventitive polyembryony).
19. PROTHALLUS: Free living gametophyte of certain lower vascular
plants. Female gametophyte of gymnosperms is sometimes termed a
prothallus.
20. SPOROPHYLL: A modified leaf that bears sporangia
21. SPOROPHYLL: Individual, usually of diploid phase of life cycle that
is formed from the fusion product of two gametes by mitosis and
throughout and reproduces by spores which germinate in haploid in life
cycle.
22. SYPHONOGAMOUS: When the male gamete along with tube and
stalk nucleus migrate to the tip of the pollen tube, which grows through
the nucellar tissue and reaches to archegonial neck and pierces through
it, bursts and then liberates the male gametes out of these one fuses with
egg to form a zygote (e.g. Pinus).
23 TEGMEN: In a zygote the inner fleshy layer in gymnospermic seed
may persist as a thin layer of seed coat.
24. TESTA: The middle stony layer changes into a hard layer called
testa in gymnosperm seeds.
25 VIVIPARY: Phenomenon of differentiation of young plants or
bulbils at the floral axis (instead of flowers) phenomenon of
germination of seeds or spores in situ on the maternal plant even
before its release, common in mangrove plants.
26. XYLEM: A type of vascular tissue which serves mainly to
translocate water and solutes.
27. ZOOIDOGAMOUS: Some extinct orders of gymnosperms did
not produce pollen tube and the sperms were liberated directly into
the pollen chamber (e.g. Cycadofilicales, Bennettitales and
Cordaitales).
REFERENCES
1. Botany For Degree Students, Gymnosperms, (1990), Vol. V, P. C.
Vashishta, S. Chand And Company Ltd., Ram Nagar, New Delhi-
110 055.

2. A Text Book of Botany, Vol. II (thirteenth Edition), S. N. Pandey, S. P.

Misra, P. C. Trivedi (2016), Vikas Publishing House Pvt. Ltd. , E-28,
Sector- 8, Noida-201301 (U. P.) India.

3. Botany For Degree Students (Revised edition) (1963), A. C. Dutta,

(Sixth edition), Oxford University Press, Ground floor, 2/11 Ansari
Road, Daryaganj, New Delhi 110002, India.

A Text Book of Botany, (2016-17), (Fifth edition), Prof. V. Singh, P. C.

Pandey D. K. Jain, Rastogi Publications, Gangotri Shivaji Road, Meerut,
250002.
 SUGGESTED READINGS

1. DICTIONARY OF BOTANY (1988). P. K. Garg, Academic (India)

Publishers, 31/1 Rajendra Nagar New Delhi-110066,
2. https: //www.Google.co.in.
3 Wikipedia.
5. Yourlibrary.com
6. www.Biologyreference.com.
7.www.biologydiscussion.com