There are ferns in most New Zealanders’ backyards and local environments.

Ferns are green flowerless plants with divided leaves that tend to grow in damp,
shady areas. The developing leaves of most ferns uncoil from a koru.

Fossil fern
Ferns are an ancient group of plants. By about 350 million years ago, ferns are seen
in the fossil record. This makes ferns older than most land animals and far older than
dinosaurs!

Ferns are ancient plants

Ferns are an ancient group of plants. From the fossil record, scientists consider
that land plants emerged from the water around 475 million years ago. By about
400 million years ago, vascular plants had separated from non-vascular plants,
and soon after this, ferns separated off. By about 350 million years ago, some of
the major families of ferns are seen in the fossil record. This makes ferns older
than most land animals – some invertebrate animals were on land by this time –
and far older than dinosaurs!

Ferns come in a variety of shapes and sizes, from the very small, like the kidney
fern, to the 20 metre tall tree fern. Most ferns share the same basic structure.

Parts of a fern
Dr Patrick Brownsey shows us the 3 major parts of a fern: the rhizome (the
underground stem), the leafy frond and the sporangia (the reproductive structure).

The structure of a fern

Ferns have 3 major parts – the rhizome, the fronds and the reproductive
structures called sporangia. The characteristics of each of these 3 parts of the
fern plant are used for classification and identification.

The rhizome is the stem of the fern plant. It comes in 3 basic forms:

 An erect rhizome, which is a solid mass that gives rise to a tuft of fronds.
You can see this type of rhizome on a king fern or a crown fern.
 A laterally growing rhizome that creeps along or under the ground. It may
even climb up a tree. Hound’s tongue and thread ferns are examples of a
fern with a creeping rhizome.
 A vertical rhizome. This can grow into a short or a tall trunk. The trunk of
the ponga (silver fern) is a vertical rhizome.

Fern structure
Ferns come in a variety of shapes and sizes and this interactive explores the
diversity of form in New Zealand ferns. The fern body consists of 3 major parts – the
rhizome, the fronds and the sporangia.

Find out more about the sporangia in this related interactive.

The fronds are the leaves of the fern. There is usually a stalk (the stipe) with a flat
blade (the lamina), often divided into segments. The frond may be simple and
undivided or it may be divided into a number of divisions (called pinnae). New
fronds are produced from the rhizome. They are tightly coiled into a spiral (called
a fiddlehead or koru), and these slowly uncoil as they mature. Fronds have a dual
function. They are there for photosynthesis but they are also there for
reproduction.

The spores grow inside casings called sporangia. These are found on the
underside of fronds. Not every frond has sporangia underneath it. Fronds that
have sporangia are called fertile fronds. In the vast majority of ferns, the
sporangia are found in clusters (called sori). These are the brown, black or
orange patches that you see on the underside of fronds. When the sporangia
break open, they release the spores.

Underlay of a fern
Ferns come in a variety of shapes and sizes and this interactive explores the
diversity of form in New Zealand ferns.

The fern body consists of 3 major parts – the rhizome, the fronds and the sporangia.
This interactive explores the sporangia.

Find out more about the upper layer of ferns in this related interactive.

Ferns are unique

Ferns are unique amongst land plants in that they have 2 separate living
structures in their reproductive cycle – the sporophyte and the gametophyte.
Fern life cycle
The life cycle of ferns is different from other land plants as both the gametophyte and
the sporophyte phases are free living. This interactive illustrates the alternation of
generations in ferns.

The leafy fern plants we see in the bush that produce spores are sporophytes.
When the spores are released by the sporangia, if they land in a hospitable
environment, they can grow into a tiny plant – the gametophyte. This
inconspicuous, short-lived plant has 2 sets of reproductive organs –
the antheridia(male) and the archegonia (female). In suitably moist
conditions, fertilisation takes place either on the same gametophyte or an
adjacent one. Fertilisation gives rise to a new sporophyte.

No other land plant has these 2 separate independent living stages. This is a
unique characteristic of ferns

Ferns come in a variety of shapes and sizes and this interactive explores the
diversity of form in New Zealand ferns.

The fern body consists of 3 major parts – the rhizome, the fronds and the sporangia.
This interactive explores the sporangia.

Find out more about the upper layer of ferns in this related interactive.
Transcript
Frond dissection
Fronds can be simple and undivided like the leather leaf fern, or pinnate (once
divided) like the thread fern. More commonly, fronds are bipinnate (silver fern) or
tripinnate (hen and chickens fern).

Image acknowledgements: University of Waikato, Steve Attwood, Public domain and

Museum of New Zealand Te Papa Tongarewa

Sori position
The sori may be located on the edge of the pinna or away from the pinna margin.

Image acknowledgement: Museum of New Zealand Te Papa Tongarewa

Sorus
Each sorus is a cluster of sporangia. The shape and position of the sori are
important for identification of ferns. They provide the main features for identifying the
different genera. Sori can be round, oval, oblong or considerably elongated. They
may occur on the edge of the pinna or away from the edge. With a 10x hand lens,
you’ll be able to see that the sori are composed of numerous, small, round bodies
that are the sporangia.

Image acknowledgement: Museum of New Zealand Te Papa Tongarewa

Indusium
A flap of tissue that protects the sori in some ferns. This can take a variety of forms.
When spores are mature and ready for release, the indusia usually shrivel or bend
backwards to expose the sporangia. Occasionally, if the indusia completely cover the
sporangia. they may tear irregularly.

Image acknowledgement: Museum of New Zealand Te Papa Tongarewa

Sporangium
The reproductive structures on the underside of the frond. Each sporangium is a
capsule that contains spores. They are usually aggregated into clusters called sori.
The position and arrangement of the sporangia are very important for the
identification of ferns. Fronds that have sporangia on their underside are fertile, and
those that don't are sterile.

Image acknowledgement: Museum of New Zealand Te Papa Tongarewa

Spore
A single cell. Spores are produced in capsules called sporangia. Most ferns produce
64 spores in each sporangium. Sporangia are aggregated into clusters called sori.
When mature, the spores are released from the sporangia. Once released, the
spores germinate readily on contact with damp soil.

Ferns stand out among garden regulars for their lack of flowers and seeds. Botanically, they
belong to the division of non-flowering plants known as Pteridophyta. With more than 10,000
known species of modern ferns, these plants display tremendous diversity in size, form and
cultural needs. Even so, some general structural and reproductive characteristics distinguish
ferns from all other plants.
Wild ferns grow in a forest. (Image: Jill Lang/iStock/Getty Images)

Stemlike Rhizomes
Instead of the stems common to most plants, ferns have rhizomes. Usually growing
underground or right along the surface, these horizontal stems hold the vascular system that
transports water and nutrients. Fern leaves spring up from the upper side of rhizomes while
roots grow on the underside. Clump-forming ferns have a crownlike rhizome, and other types
spread. Ostrich fern (Matteuccia struthiopteris), for example, hardy in U.S. Department of
Agriculture plant hardiness zones 3 through 7, colonizes eagerly as its running rhizomes
sprout leaves along their length. Tree ferns' rhizomes grow erect. The trunklike, 28-inch-
diameter rhizome of soft tree fern (Dicksonia antarctica, USDA zones 9 through 10) takes the
plant to a height up to 30 feet.

New fiddle head ferns sprout

from a forest floor. (Image: Jill Oliver/Hemera/Getty Images)
Leafy Fronds
A fern frond begins at the rhizome and ends at the leaf tip. The base stalk, called a stipe,
holds the leafy portion known as the blade. Fern leaves can be whole, such as the straplike
leaves of bird's nest fern (Asplenium nidus, USDA zones 11 through 12). They can also be
divided and subdivided into complex leaf forms, right down to 1/16-inch fronds of aquatic
ferns. Many fern fronds unfurl in a classic fiddlehead shape known as a crozier. Some are
edible; some are not. Small hairs and scales on some fronds reflect light and create colorful
foliage. The fronds of Christmas fern (Polystichum acrostichoides, USDA zones 3 through 9)
remain evergreen.

A close-up of a bird's nest fern.

(Image: Picheat Suviyanond/iStock/Getty Images)

Spores and Sporangia

Rather than seeds, ferns produce tiny spores. The spore-producing capsules, known as
sporangia, form in clusters called sori. Often mistaken for scale insects, sori appear in
distinctive shapes, patterns and places that identify different fern species. In some ferns, a
leaf section rolls over and protects the sori. Other species have protective, disclike flaps
called indusia. Kidney-shaped indusia cover the round sori of autumn fern (Dryopteris
erythrosora, USDA zones 5 through 8) and mature in colorful stages from pink to red. When
the time is right, fern sporangia catapult their spores onto the wind. A single fern produces
multitudes of spores in a lifetime, but few survive.
Sporangia form on the underside of fern leaves. (Image: Victorburnside/iStock/Getty Images)

Distinctive Reproduction
Spores are just the first step in fern reproduction. The spores that find hospitable conditions
form simple, heart-shaped, 1/2-inch-wide plants called prothallia, which lie on soil. These
inconspicuous plants form male and female organs. If moisture is present, fertilization can
occur, and new ferns eventually result. Some desert ferns can reproduce without water or
fertilization, but most ferns demand moisture. Characteristic habitats for survival and
reproduction are moist, humid, shaded areas that retain dampness year-round. Few ferns
tolerate direct sunlight, low humidity, boggy conditions or deep shade. Evergreen species
tolerate the lowest light levels. The more sun exposure, the more water ferns require.

The young leaves of a fern open up. (Image: Linjerry/iStock/Getty Images)

Characteristics of Pteridophyta |
Botany
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The Pteridophyta, commonly called vascular cryptogams,

are represented by more than about 10,000 species, and
are characterized by the following peculiarities:
1. Members of this group are most primitive living vascular plants
such as Selaginella, Lycopodium, Equisetum etc., and also fossil
vascular plants such as Rhynia, Horneophyton, Asteroxylon, etc.

2. Most of the plants prefer to grow in cool and shady places while
some are xerophytic, e.g., Selaginella rupestris, and many occur in
aquatic conditions like Marsilea, Salvinia, Azolla, etc.

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3. Plant body is sporophytic and differentiated into roots, stem and

leaves. In some plants an intermediate stage between root and stem,
i.e., rhizophore, is present as in Selaginella.

4. Young sporophyte is partially or completely dependent on

gametophyte for some time, but at maturity it becomes
independent.

5. The fibro-vascular cylinder, consisting of xylem, phloem, etc., is

present in the stem of these plants.

6. In lower members, the stele is protostele, as in Lycopodium,

Selaginella, etc., but it is siphonostele in Marsilea (amphiphloic),
Equisetum (ectophloic), and dictyostele as in Pteris, Aspidium,
Polypodium, Dryopteris, etc.

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7. Vascular supply to the leaves takes place by leaf traces through
leaf gaps of the vascular cylinder of stem.

8. Leaves may be simple, small and sessile, as in microphyllous

types such as Selaginella, Lycopodium, etc., or very large, petiolate
and megaphyllous types as in the members of Filicinae.

9. Plants reproduce by the spores formed in sporangia. Sporangia

develop either on the ventral surface or in the axil of leaves.

10. Plants may be homosporous, i.e., all the spores are of one type
as in Equisetum, or heterosporous, i.e., two types of spores are
present (microspores and megaspores) as in Selaginella

Classification of Pteridophyta: 4
Classes | Botany
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In this article we will discuss about the classification of

pteridophyta.
1. Class: Psilotopsida:
The members of the class Psilotopsida show close resemblance in
fundamental characteristics to the Silurian and Devonian members
of Rhyniopsida (e.g., Rhynia, Cooksonia), Zostero- phyllopsida (e.g.,
Zosterophyllum) and Trimero- phytopsida (e.g., Trimerophyton,
Psilophyton). Psilotopsida includes only two living genera viz.,
Psilotum and Tmesipteris.
Characteristic Features of Class Psilotopsida:
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1. The plant body is a rootless sporophyte that differentiates into a

subterranean rhizome and an aerial erect shoot.
2. Branching is dichotomous in both subterranean rhizome and
aerial shoot.
3. The large rhizoids borne on the rhizome absorb water and
nutrients from the soil.
4. On the aerial shoots, spirally arranged scale-like (e.g., Psilotum)
or leaf-like appendages (e.g., Tmesipteris) are borne.
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5. Stele is protostelic or siphonostelic with sclerenchymatous pith.

6. Secondary growth is absent.
7. Bi- or trilocular sporangia are borne in the axils of leaf-like
appendages.
8. Mode of sporangial development is of eusporangiate type.
9. Spores are of equal sizes and shapes i.e., homosporous.
10. The gametophytes are non-green, cylindrical, branched and
subterranean. They grow as saprophytes with an associated
endophytic fungus.
11. Antherozoids are spirally coiled and multi- flagellated.
2. Class. Lycopsida:
This class has a long evolutionary history and is represented both by
extant and extinct genera. This group first originated during the
Lower Devonian period of Palaeozoic Era (ca 390 my).
This class is represented by five living genera
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— Lycopodium, Selaginella, Phylloglossum, Styhtes, and Isoetes,

and fourteen extinct genera
— Asteroxylon, Baragwanathia, Protolepido- dendron,
Lepidodendron, Sigillaria etc.
Salient Features of the Class Lycopsida:
(a) The sporophyte plant body is differentiated into definite root,
stem and leaves.
(b) The sporophytes are dichotomously branched.
(c) The leaves are usually small and micro- phyllous.
(d) The xylem in stem exarch.
(e) Sporangia are borne singly on the ada- xial (upper) surface of
the sporophylls.
(f) The spores may be of either one type i.e., homosporous (e.g.,
Lycopodium) or two types i.e., heterosporous (e.g., Selaginella).
(g) The spores develop into independent gametophyte.
3. Class: Sphenopsida:
This class is represented by only one living genus (Equisetum) and
about 18 extinct forms (e.g., Calamites, Annularia etc.). This group
originated during the Devonian period of Palaeozoic Era, attained
their maximum development in the Carboniferous period. Sub-
sequently, the group became less prevalent and at present is
represented by only a single genus (Equisetum).
Salient Features of the Class Sphenopsida:
1. The stems and branches are jointed with nodes and internodes.
The internodes are with longitudinal-oriented ridges and furrows.
2. The leaves are extremely reduced and borne in whorls at the
nodes of,aerial branches and stems.
3. Branches arise in whorls.
4. The sporangia develop on a peltale appendage called
sporangiophore. Sporangial walls are thick.
5. Most of the” members are homosporous including Equisetum.
However, some extinct forms were heterosporous (e.g., Catamites
casheana).
6. The gametophytes are exosporic and green.
7. Antherozoids are multiflagellated.
8. The embryo is without suspensor and is exo- scopic in nature.
4. Class: Pteropsida:
This group of pteridophytes is commonly known as ‘ferns’. The
Pteropsida differs from other classes in possessing raised leaves
(mega- phylls). This is the largest and highly evolved group of
pteridophytes and is represented by about 9,000 species which
show a wide range of distribution. The Pteropsida are known from
as far back as the Devonian period of Paleozoic Era.
Salient Features of the Class Pteropsida:
1. The sporophytes are usually perennial in nature and
differentiated into roots, stem and spirally arranged leaves.
2. Most of the members grow in moist and shaded habitats, either
epiphytic or terrestrial. A few are aquatics.
3. Mostly, the rhizomes are short and stout.
4. The leaves are large (megaphylls), pinnately compound and
described as frond, except Ophioglossum (simple leaf).
5. The rachis is covered with brown hairs (ramenta). Leaf trace is
usually C-shaped with adaxial curvature.
6. Young fronds show circinate vernation (coiling of leaves), except
Ophioglossum.
7. The stele in Pteropsida shows a wide variety of types, e.g.,
protostele, siphonostele,
solenostele, dictyostele and polycyclic stele.
8. Most ferns are homosporous, but a few aquatic members are
heterosporous.
9. Sporangia are borne at the tips or at the margin of the pinnule or
to the abaxial surface of the fronds.
Classification of Pteridophyta |
Botany
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In this article we will discuss about the classification of

pteridophyta.
i. Eusporangiate type, in which the sporangium develops from a
group of superficial initials, e.g., Selaginella, Equisetum.

ii. Leptosporangiate type, in which the sporangium develops from a

single initial, e.g., Marsilea.

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a. Spores develop into a multicellular, gametophytic body called

prothallus.

b. Two sex organs, i.e., antheridia and archegonia develop on the

prothallus.

c. Fertilization takes place with the help of water, and results into
the formation of a diploid zygote. It develops into a structure
bearing roots, stem and leaves. This structure is called sporophyte.

d. Plants show clear alternation of generations.

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18 Important Characters of
Gymnosperms – Explained!
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Some of the most important characters of gymnosperms
are as follows:
1. Habit:
The living gymnosperms are woody, evergreen (except Larix and a
Taxodium) perennials grow as trees or shrubs. Tallest trees are
Sequoia sempervirens (366ft) and S. gigantia (342ft).

2. Occurrence:
The living members are founding in colder regions of earth where
snow (not rain) is the source of water. Only the members of
cycadales and gnetales thrive in warm dry climate.
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3. The dominant plant body is sporophyte (2n) which may be

dioecious or monoecious. Gametophytes are inconspicuous and
endosporic i.e. develop with the spores.

4. Sporophyte differentiated into — root, stem and leaves.

Root system:
Tap root system is exarch and diarch to polyarch. Besides tap root,
coralloid roots (in cycads) and mycorrhizal root (in coniferals)
present.

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Stem: Erect, generally branched (Cycas is un-branched)

Leaves:
Dimorphic i.e. 2 types, foliage and scale leaves.

5. Xerophytic Traits:
Gymnosperms are xerophytes in nature due to presence of thick
bark, thick hypodermis, thick cuticle, scales leaves, sunken stomata,
transfusion tissue, etc. In some cases leaves modified into needle-
like, scale like or small leathery. These are the adaptations to
combat water stress in air and colder regions.
6. Xylem:
Composed of xylem parenchyma and tracheids with bordered pits.
Vessels are absent (exceptin Gnetales).

7. Pliloem:
Composed of sieve cells and phloem parenchyma but companion
cells absent.

8. Heterospcry:
The gymnosperms are heterosporous, means 2 types of spores
produced i.e. haploid microspores and megaspores. Microspores
produced within micro-sporangia while megaspores produce within
megasporangia (nucellus) of ovules. Both types of sporangia are
formed on special leaf-like structures called sporophylls
(microsporophylls and megasporophylls).

9. Cones or Strobili:
Sporophylls are spirally arranged along an axis to form compact
cone or strobili i.e. male or pollen cones and female or seed cones
but in Cycas female cone is loosely arranged called lax.

10. Ovules:
Naked, sescile, geneiallyorthotropous, and unitegmic or bitegmic
(in Gnetum).

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11. Pollination is anemophillous i.e. by wind.

12. Fertilization is siphonogamous i.e. male gametes carried to

female gametes by means of a pollen tube. Double fertilization, a
feature unique to angiosperms, is absent in gymnosperms, but
found in Ephedra. After fertilization, Zygote develops into embryo
and ovules become seed.

13. Endosperm or female gametophyte formed before fertilization

and is always haploid. (But triploid in angiosperms.)

14. Embryo development is meroblastic i.e. develops from a small

part of zygote.
15. Most members show polyembryony i.e. development of more
than one embryo, but only one survives at the end.

16. Cotyledons 2 (in Cycas) or many (in Pinus 2-14).

17. Cambium form secondary wood which is monoxylic (soft and

porous) or pycnoxylic (hard and compact)

18. Connecting Links:

The lower gymnosperms like Cycadales resemble pteridophytesin
absence of vessels and wood fibers whereas higher gymnosperms
like Gnetales and Coniferales resemble angiosperms.

Gymnosperms: Characteristics, Types

and Economic Importance
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In this article we will discuss about:- 1. Characteristics of

Gymnosperms 2. Types of Living Gymnosperms
3. Economic Importance.
Characteristics of Gymnosperms:
Gymnosperms are, those seed plants in which the seeds remain
exposed over the surface of the megasporophylls because the latter
are not folded to form pistils.

1. Gymnosperms are a small group of seed plants which are

represented by only 900 living species.

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2. Gymnosperms are more ancient than the angiosperms. They

formed dominant vegetation on earth some 200 million years back
in mesozoic era. Today they are dominant only in cold areas, where
instead of rain; snow is the source of water.
At other places they have been replaced by angiosperms. In warmer
areas only a handful of gymnosperms can be observed, e.g., Cycas
(like C. circinalis in South India), Araucaria (native of South
America, New Zealand and East Australia, like A. heterophylla.

3. All gymnosperms are perennial and woody, forming either

bushes or trees. Some of these are very large and live for thousands
of years, e.g., Sequoia sempervirens (tallest gymnosperm of 111.6 m)
Zamia pygmia is smallest (26 cm).

4. Flowers are absent. Two types of sporophylls, microsporophyll’s

and megasporophylls are usually aggregated to form distinct cones
or strobili, pollen cones (male cones) and seed cones (female cones)
respectively.

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5. Seeds do not occur inside a fruit. They are naked.

6. A distinction of ovary, style and stigma is absent.

7. Ovules are orthotropous and sessile. Each ovule is surrounded by

a 3-layered integument.

8. Female gametophyte contains archegonia.

9. Pollination is direct as a stigma is absent and the pollen grains

directly reach the micropylar ends of ovules. Pollination is usually
accomplished by wind (anemophily).

10. Male gametophyte produces only two male gametes or sperms.

Generally one of them is functional.

11. An external water is not required for transport of male gametes.

Instead, a pollen tube is formed by the male gametophyte for
effecting fertilization (siphonogamy).

12. Seeds contain a food laden tissue or endosperm for future

growth of embryo into seedling. The tissue represents the female
gametophyte.
13. Like pteridophytes, xylem does not possess vessels except in
some geophytes. Phloem is without companion cells and sieve
tubes. Sieve cells are not arranged end to end in rows.

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14. Vascular tissues are arranged into vascular bundles just like
angiosperms. Vascular bundles of stem are open so that secondary
growth is quite common.

Types of Living Gymnosperms:

Living gymnosperms are divided into four groups— Cycadopsida,
Ginkgopsida, Coniferopsida and Gnetopsida. Ginkgopsida is
represented by maiden hair tree (Ginkgo biloba).

Gnetopsida contains Gnetum, Ephedra and Welwitschia. These are

nearer to flowering plants in having flower like arrangement of
sporophylls and possessing primitive vessels in xylem. Plants of
cycadopsida and coniferopsida are commonly called cycads and
conifers respectively.

i. Cycads:
It is smaller group of gymnosperms which have palm-like habit and
fern-like foliage. Leaf bases are often persistent. In stem the
vascular tissues constitute only a narrow patch though secondary
growth occurs. Plants are dioecious, that is, microsporophyll’s and
megasporophylls develop on separate plants. The sporophylls are
aggregated to produce strobili or cones (exception megasporophylls
of Cycas).

Megasporophylls are often leaf-like and possess ramenta. Sperms

are motile. The plants are mostly xerophytic and occur in warm
weather. Their population is, however, decreasing due to extension
of agriculture and severe competition from xerophytic angiosperms.
Cycads are grown for their ornamental appeal, e.g., Cycas circinalis,
C. revoluta.

Cycas (Fig. 3.18):

Cycas is also called living fossil because it possesses a number of
characters of extinct pteridosperms and cycads. The gymnosperm is
an evergreen small palm-like or tree-fern like sporophyte that
occurs in some tropical and subtropical areas.

It reaches a height of 0.4 m to 20.0 m. Cycas has an un-branched

columnar stem (rarely branched) which is covered by spiral bands
of persistent rhomboidal leaf bases. The top bears a crown of leaves.

Leaves are of two types, large green foliage and small brownish
scale leaves. Scale leaves develop in spiral rows alternating with
foliage leaves. They are covered with ramental hair. Foliage leaves
are large (1-3 m), petiolate and unipinnate. Base is broad. Petiole
bears two rows of spines. Petiole continues into rachis. Rachis has
two lateral grooves from which arise 50- 200 pairs of sessile, linear-
lanceolate pinnae or leaflets.

Leaflet apex is spiny while the margin can be straight or revolute

(e.g., C. revoluta, C. beddornei). Young leaves show circinate ptyxis.
Roots are of two types, normal and coralloid.

Coralloid roots are irregular, negatively geotropic, dichotomously

branched coral like roots which do not possess root hairs and root
caps. Coralloid roots have a symbiotic association with blue-green
algae like Nostoc and Anabaena species. Sago grains occur in the
stem cortex.

Vegetative reproduction occurs by fleshy bulbils which arise

adventitiously in the crevices amongst persistent leaf bases.
Sexually Cycas is dioecious with distinct male and female plants.
The plants bear terminal cones. Male cone is oval, thick, 20-80 cm
long stalked structure which replaces the growing point of the plant.
Further growth of male plant is carried out by a lateral bud, so that
stem axis of male plant is sympodial.

Male cone has a number of densely crowded spirally arranged

microsporophyll’s. Each microsporophyll has a flat fertile proximal
region and a distal bent sterile region or apophysis. The fertile
region bears a number of microsporangia or pollen sacs arranged in
sori. Microspores or pollen grains are boat-shaped.

Female cone is loose. It consists of a rosette of brown

megasporophylls. Each megasporophyll bears 2-12 reddish ovules
in the middle fertile part. The terminal sterile part may show
remains of foliar structure. Ovules of Cycas are the largest, 6-7 cm
in diameter. Similarly, egg or oosphere is the largest in plant
kingdom.

Pollination is anemophilous. Before pollination, pollen grain

develops three cells prothallial cell, generative cell and tube cell.
Pollination brings pollen grains inside the ovules. The tube cell
grows and forms a pollen tube. Pollen tube is haustorial in function
in Cycas. Generative cell divides into sterile stalk cell and large
fertile body cell. Body cell produces two sperm mother cells.

The sperms of Cycas (upto 300 pm) are the largest in the biological
world. They are top-shaped with a number of flagella present in 5-6
grooves on the pointed end. Seed is oval to rounded orange-red
structure which generally has a fleshy outer sarcotesta for attracting
birds.
ii. Conifers:
They comprise more than 500 living species of cons bearing
gymnosperms which have withstood competition from
angiosperms. The plants are evergreen with dense and massive
vascular tissues and non-motile gametes. Conifers are usually
monoecious. Cones or strobili are compact and woody. Both the
types of cones are borne on the same plant. Unlike cycads, they are
not produced on the tips of main branches.

Conifers have monopodial growth where the main axis of trunk

rises straight from base and reaches up to the tip. The branches do
not compete with the main axis. Stem branches are of two types,
long and dwarf. Growth of the long branches is indefinite. The lower
long branches are longer than the upper ones. This gives a conical
or ex-current appearance to the plant just like a ‘Christmas Tree’.

Leaves are mostly borne on the dwarf branches. The dwarf branches
have a limited or definite growth. Leaves have thick cuticle and
sunken stomata. They are needle like (e.g. Pinus), small flat and
leathery (e.g., Araucaria) or scale like (e.g. Thuja). All these leaf
characters help conifers in conserving water and reduce
transpiration in dry conditions associated with cold habitat.

The roots are often associated with fungal symbionts or

mycorrhizae so as to overcome difficulty in absorbing water and
obtaining inorganic nutrients from organic debris.

The plants usually possess an aromatic antiseptic sticky semifluid

substance called resin. It is secreted by special channels or ducts
present in wood, bark, leaves, roots, etc. Resin seals the places of
injury and protects the plants from attack of microbes as well as
insects.

Araucaria (a tall and majestic conifer of South America, New

Zealand and East Australia) grows in warm weather in plains as well
as foot hills.

Other conifers form vast and dominant woodlands in mostly north

temperate regions of Europe, Asia, North America, etc. They are
abundant on the upper timber-line zone of mountains including the
Himalayas, e.g., Pinus (Pine), Cedrus (Cedar), Picea (Spruce), Abies
(Fir), Juniperus (Juniper), Thuja, Larix, Cupressus.

Conifers are dominant constituents of north temperate

flora due to the following reasons:
1. In the temperate areas, conifers have an advantage over
angiospermic trees. While the angiospermic trees shed their leaves
during autumn-winter period, the conifers remain evergreen. They
continue to manufacture their food during this period when other
plants are in a state of hibernation because of the absence of leaves.
2. Conifers have a number of xerophytic characteristics which help
in conserving water. They are, thus, able to tide over the winter
period when the soil becomes frozen and water availability is very
little. Mycorrhizae allow them to get water and minerals.

3. Enzymes of conifers are functional even at —35°C at which

temperature they become inactivated in other plants.

Pinus (Fig. 3.18):

Pinus or Pine is a coniferous gymnosperm. It is a large tree having
pyramidal or ex-current shape like a Christmas tree. The plant body
is a sporophyte having stem, leaves and roots.

The main stem is straight. It may reach a height of 10-50 metres.

The main stem shows monopodial branching. Stem branches are of
two types, long and dwarf. The dwarf branches possess 1-5 needle-
like foliage leaves which are surrounded at the base by a sheath of
scale leaves.

Pinus has a horizontally spreading tap root system with peg-like

downwardly directed roots present at intervals. Finer roots are of
two types— normal (with root hairs and root cap) and mycorrhizal.
The mycorrhizal roots occur near the soil surface. They are devoid
of root hairs and root cap.

The sporophytic plant body does not multiply vegetatively. It

produces micro- and megasporophylls in two types of cones, male
and female. Male or pollen cones occur in clusters sub-terminally
on lower long branches. Each male cone has a short stalk, a central
axis and a number of spirally arranged microsporophyll’s.

A microsporophyll bears two oblong, parallel microsporangia on its

lower surface. Pollen grains are dispersed by air currents. They form
yellow clouds in the pine forests. A pollen grain has two air sacs or
wings for making it light.

The female or seed cones develop in groups of 2-6 on upper long

branches of the tree. Each female cone has a long stalk and a central
axis on which are borne a number of spirally arranged paired scales.
The lower of the pair is called bract scale while the upper scale is
ovuliferous scale or megasporophyll. The ovuliferous scale bears
two ovules towards the basal region on the upper side.

The female cones open in the year of their formation for pollination.
Pollination is anemophillous (by air) and direct. After fertilization
the ovule matures into a seed.

Part of the upper surface of the ovuliferous scale is peeled off along
with the seed to form its wing. A female cone takes about 26 months
for reaching maturity. It then opens to release winged seeds which
are dispersed by air. After falling on a suitable soil, each seed gives
rise to a new plant.

iii. Ginkgoales:
They are primitive gymnosperms which possess deciduous leaves
with furcate venation, two types of branches, unisexual plant, catkin
like male inflorescence with each microsporangiosphore having 2-
12 microsporangia, multi ciliate male gametes and clusters of
megasporangiate structures each with a long stalk and two ovules.
Ginkgoales are represented by a single species, Ginkgo biloba.

Ginkgo biloba:
Ginkgo biloba (Fig. 3.19) has not changed for the last several
millions of years since its appearance in triassic period. It is also
called living fossil. The plant has survived due to interest shown by
horticulturists. It has natural immunity to several plant diseases.
The plant is a tall tree of upto 30m height. Leaves are fan-shaped.

They are deeply bilobed on long shoots. The leaves are entire or
sinuate on dwarf shoots. Leaves possess furcate venation. Ginkgo is
dioecious like cycads. Dwarf shoots of female plants bear distinct
megasporangiate structures.

Each has a stalk that ends into two ovules. Ovule has a 3-layered
integument, micro Pyle and a pollen chamber. Endosperm of
roasted seed is edible. However, horticulturists prefer to grow only
male plants as the female plants give unpleasant smell.

Economic Importance of Gymnosperms:

Some of the Major Economic Importance’s of
Gymnosperms are given below:
1. Edible Seeds:
Seeds of Pinus gerardiana are edible. They are eaten after roasting.

2. Timber:
Gymnosperms possess softwood. The same is used in preparation of
light furniture, plywood, packing cases, match sticks, railway
sleepers, etc.

3. Paper:
A number of gymnosperm woods are used in the manufacture of
paper. They include Picea, Pinus, Larix and Abies.

4. Fibre Boards:
Needles of Pinus and other conifers are used in making fibre boards
that are used in making packing cases.

5. Linoleum:
Saw dust is employed in making linoleum and plastics.

6. Resin:
Resin is a semifluid secreted by special tubes which contains
terpenes, resin acids and esters. It solidifies on exposure to air.
Therefore, it plugs the places of injury. It helps in sealing female
cones after pollination, scale leaves around leaf bases and apical
buds. Resin retains water. It is antiseptic and toxic to pests.

Therefore, it prevents microbial and insect attack. Resin is

commercially extracted and distilled to obtain turpentine and rosin.
Rosin is used in water proofing, sealing joints and preparation of
writing paper. Turpentine is used as solvent in paints, polishes and
wax. It is employed medicinally in removing pains, curing
bronchitis and expelling worms.

7. Ephedrine:
Drug ephedrine is obtained from Ephedra, a gymnosperm. The drug
is used in curing respiratory ailments, including asthma.

Gymnosperms: Classification and

Main Orders | Botany
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Let us learn about Gymnosperms. After reading this

article you will learn about: 1. Classification of
Gymnosperms 2.Main Orders of Modern Gymnosperms.
Classification of Gymnosperms:
It was generally accepted, and expressed in schemes of
classification, that the Gymnospermae formed a group coordinated
with the Angiospermae. From time to time suggestions have been
made for splitting the Gymnospermae into a number of major
groups taxonomically equivalent one to another and to the
angiosperms.

Coulter and Chamberlain (1910) divided the gymnosperms into

seven orders. Some of the orders are quite extinct and not found in
present day.

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These orders are as follows:

1. Cycadofilicales-extinct

2. Bennettitales-extinct

3. Cycadales-Mesozoic to present day

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4. Cordaitales-extinct

5. Ginkgoales-Palaeozoic to present day

6. Coniferales-Palaeozoic to present day

7. Gnetales-Recent.

Chamberlain (1934) however, divided the gymnosperms into two

large groups A. Cycadophyta and B. Coniferophyta.

In the Cycadophyta, sporophylls are in cones.

They include three orders:

Cycadofilicales (Pteridospermae)-extinct

Bennettitales (Cycadeoideales)-extinct

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Cycadales-Mesozoic to present day.

The Coniferophyta is divided into four orders:

Ginkgoales-Palaeozoic to present day.

Cordaitales-extinct

Coniferales-Palaeozoic to present day

Gnetales-Recent.

Prof. B. Sahni (1920) divided the gymnosperms into two

large groups and gave them the terms:
1. Phyllospermae and

2. Stachyospermae.

1. Phyllospermae:
They comprise of the Pteridosperms and the Cycadophyta
(Cycadales and Bennettitales). The seeds are being inserted on the
modified leaves.

2. Stachyospermae:
This group includes the orders of Coniferophyta of Chamberlam.
They are more or less microphyllous plants with seed inserted on
the stems.

Raizada and Sahni (1938) have summarized the

classification of gymnosperms as follows:
Cycadophytes:
Pteridospermae (Cycadofilicales)-Carboniferous. Extinct.

Cycadeoideales (Bennettitales)-Mesozoic. Extinct.

Cycadales-Mesozoic to present day.

Pentoxylales: Jurassic
Coniferophytes:
Cordaitales-Palaeozoic. Exinct

Ginkgoales-Palaeozoic to present day.

Coniferales-Palaeozoic-Mesozoic to present day

Gnetales-Recent.

The characteristic features of these orders are as follows:

1. Pteriodspermales (Cycadofilicales):
Trees or small plants with fem-like leaves. Sporangia borne on
frond-like leaves, more or less modified, but never formed a cone.

2. Cycadeoideales (Bennettitales):
Trees with stems covered with an armour of persistent leaf bases.
Male sporophylls frond-like forming a loose crown. Female
sporophylls found in a cone-like structure, no longer leaf-like.

3. Cycadales:
Plants with palm-like habit, male and female sporophylls grouped
into cones, except in the genus Cycas where the female sporophylls
are collected into a loose crown. Male cells (gametes) motile.

4. Pentoxylales:
Low shrubs. Female inflorescence consisting of numerous cones
spirally attached on a peduncle, each bearing numerous ovules on a
central axis. No megasporophylls, ovuliferous scales or inter-
seminal scales. Male flowers terminally borne on dwarf
shoots. Unilocular sporangia terminate short branchlets of the
appendages or microsprophylls.
5. Cordaitales:
Large trees with flat strap-shaped leaves. Fructifications in cones.

6. Coniferales:
Large trees or shrubs. Leaves needle or scale-like, sometimes
flattened, male cells never motile. The male flowers are arranged in
more or less compact clusters or catkins and female flowers are
usually clustered to form a cone.
7. Ginkgoales:
Medium sized trees. Leaves flattened and lobed in various ways
with dichotomous venation. Male and female flowers collected into
strobili.

8. Gnetales:
Small trees or shrubs or climbing shrubs. Opposite leaves,
dicotyledonous embryos and compound cones in both male and
female.

Modern taxonomy divides the gymnosperms into six or seven

classes or orders.

These classes or orders are as follows:

1. Pteridospermae (Cycadofilicales)

2. Cycadinae (Cycadales)

3. Bennettinae (Bennettitales)

4. Cordaitinae (Cordaitales)

5. Ginkgoinae (Ginkgoales)

6. Coniferae (Coniferales)

7. Chlamydospermae (Gnetales)

From the view point of their phylogeny the Gnetales or

Chlamydospermae are placed intermediate between the
gymnosperms and the angiosperms. Though the order includes
three families yet recently several authors raised the families to the
rank of orders because of the divergence in the characters of the
genera belonging to them. The orders are—Ephedrales, Gnetales
and Welwitschiales.
Economic Importance of Gymnosperms:
Economically the gymnosperms are of very great importance in
providing timber and wood pulp and they are planted in many parts
of the world under schemes of forestation and reafforestation. The
Coniferales are the most important order of forest trees in the
economy of civilized man. Their timber is of universal importance.
Coniferous timber is the cheapest and the best.

It is none to second in durability, lightness, elasticity and fineness.

The family Pinaceae in particular is of great importance as far as the

production of timber and resin is concerned. In the coniferous-pine
forests of Himalayan region-Northern India the resin is collected on
commercial scale. Resin and turpentine are a source of immense
revenue to India.

Cedrus deodara (Deodar) is the strongest of the Indian coniferous

woods and is in great demand for construction work and furniture.
The railway sleepers are made from deodar wood.

The Government of India is keeping in view to manufacture

newsprint out of fir and Himalayan spruce to save a lot of foreign
exchange. Large quantities of fir and spruce are available in
Kashmir, Himachal Pradesh and Tehri-Garhwal region.

The annual revenue from the sale of pine resin from the
Government owned coniferous forests of Kashmir, Himachal
Pradesh, Uttar Pradesh, West Bengal and Assam is immense and
millions of rupees. The revenue derived from the sale of coniferous
timber of Government owned forests of above mentioned states
adds much to the economy of country.

Main Orders of Modern Gymnosperms:

The modern gymnosperms are commonly grouped under
four orders:
1. Cycadales;

2. Ginkgoales;

3. Coniferales and

4. Gnetales

The Cycadales and the Ginkgoales include living members which

have a long, fossil history and can be regarded as ‘living fossils’.
Ginkgoales in the past (early Mesozoic) were represented by widely
distributed group of plants, but now the order is represented by a
single species Ginkgo biloba.

In the similar way Cycadales flourished well during the Mesozoic

and are now represented by nine well defined genera that are
confined to limited areas in the tropical and subtropical countries of
the world.

The Coniferales from the most conspicuous order of the living

gymnosperms and include the plants like Pinus, Cedrus, Abies,
Juniperus, Cupressus, Biota, etc.

Some conifers are the world’s tallest and long-lived plants, e.g.,
Sequoia gigantea lives for 4,000 years and grows very tall (300 to
400 feet in height). The conifers are cosmopolitan and widely
distributed throughout the northern and southern hemispheres of
the globe and form extensive forests.

The Gnetales are represented by three living genera, e.g., Gnetum,

Ephedra and Welwitschia.

Genus Welwitschia is monotypic and represented by W. mirabilis.

This is found in deserts of South-West Africa. Ephedra (30 – 40
species) has many species that are distributed in tropical and
temperate regions of Asia, Africa and South America.

Besides, the coniferous forests provide a cool and soothing climate

suitable for health. Several health resorts and sanatoria run in the
hills under the shadow of pine trees where hundreds of people and
patients go every year for recreation and treatment.

The pith of Cycas revoluta yields sago, and thus, the plant is called
the sago palm. The seeds of Pinus gerardiana are edible and known
as chilgoza. They are generally found in the dry arid zones of
Himalayas. The famous drug known as epedrine is obtained from
different species of genus Ephedra. The young leaves and strobili of
Gnetum are eaten as vegetable.

Evolution of Gymnosperms:
However, the gymnosperms are not in such an
advantageous position because:
(i) They lack vegetative means of reproduction by means of cuttings,
layering, etc. and are slow growing.
(ii) Limited means of dispersal (only wind and by man) and their
failure to grow under varied habitats.

(iii) Absence of vessels in xylem and of companion cells in the

phloem.

(iv) Absence of bisexuality which reduces the chances of self-

fertilisation and lot of pollens are wasted, as wind-pollination is the
main source of carrying pollen grains,

(v) Unprotected ovules and seeds.

(vi) Their decline may also be due to the running out of the gene
potential for environmental adaptation and appearance of certain
harmful mutations and chromosomal alterations.

(vii) Economically too they are less important in comparison of

angiosperms.
The gymnosperms are the most ancient seed plants that originated
during the late Palaeozoic (i.e., 350 million years ago from the
upper Devonian to the present) era, but flourished well during the
Mesozoic. Their long evolutionary history is full of diverse lines of
evolution within the group, during which many organisms
flourished and then became extinct.

Most of them had fernlike general appearance and foliage but

possessed the primitive type of seed. The seeds were found attached
to the pinnatifid fronds, e.g., Medullosa noei.

Gymnosperms vs Ferns

Many people know what ferns are. Not by how their life cycle
progresses but by their general appearance. But many don’t
know what gymnosperms are. This group of plants is rather more
technical but it is very surprising to know that they are just one
of those familiar tree plants and shrubs that you happen to see
around.

Ferns are plants that do not bear flowers. They do not have any
seeds as well. In this regard, their mode of reproduction is
through spores. Gymnosperms on the other hand have seeds,
although they are not put inside an ovary. Hence, they procreate
using those seeds. Examples of such are the conifers and cycads.

In terms of family classification, ferns are grouped within the

division Pteridophyta whereas gymnosperms have four divisions
namely: Pinophyta (conifers), Ginkgophyta, Gnetophyta and
Cycadophyta. The first division is regarded as the biggest of them
all that specifically includes pines trees.

Moreover, it has been noted that ferns are the bigger group of
plants that is comprised of almost 20,000 different species
whereas gymnosperms only have less than 1,000. There are 700
to 900 species of gymnosperms to be more exact.

In more detail, the life cycles of ferns and gymnosperms really

differ. Ferns’ cycle is usually described as an alternation of
several generations that usually start in a sporophyte (diploid
cells) producing many spores through meiotic cell division. The
latter grows further through mitotic division to form
gametophytes. These gametophytes create gametes (usually
sperms and eggs altogether). The more motile sperm will then
look for an egg to fertilize while still being affixed to the
prothallus. After fertilization, the resultant product is a new
diploid cell that matures through a second round of mitotic
division hence making the actual sporophyte ‘“ the fern.

Gymnosperms are also termed as sporophytes that create spores.

Their mode of reproduction slightly varies depending on the
division to which the gymnosperm belongs. For example, cycads
have more mobile sperms that can go immediately towards the
egg in the ovule by swimming. Conifers on the contrary have
flagella-less or ‘tail less’ sperms that are transmitted to the eggs
using a certain pollen tube. Also, there is a difference in the
gametophytic life cycle stages between ferns and gymnosperms
wherein ferns have complex liberally surviving gametophytes
unlike gymnosperms.

Summary :
1.Ferns are flowerless plants that do not have any seeds whereas
gymnosperms do have seeds of their own.
2.Ferns are grouped in one division whereas gymnosperms have
four different divisions.
3.Ferns have a bigger number of species compared to
gymnosperms.
4.Ferns have free-living gametophytes whereas gymnosperms
don’t.

Read more: Difference Between Gymnosperms and Ferns |

Difference Between | Gymnosperms vs
Ferns http://www.differencebetween.net/science/nature/differ
ence-between-gymnosperms-and-ferns/#ixzz5iyEbvnEX

Extant gymnosperms comprise four distinct lineages of

seed plants with prevalent sexual reproduction. Characters and
processes involved in sexual reproduction of gymnosperms are
described in this chapter in a comparative approach. Phylogenetic
evidence suggests monophyly for this group and some characters
involved in sexual reproduction are shared between taxa, although
remarkable exceptions are always found. Breeding systems varies
between basal and derived taxa, prevailing dioecy in Ginkgoales
and Cycadales, monoecy in conifers and unfunctional gynodioecy
in Gnetales; in any case outcrossing mating systems are promoted.
These plants have long life spans and long reproductive cycles
characterized by a delayed gametogenesis. Male reproductive
structures are simple and morphological patterns are similar between
groups with exceptions in Gnetales and Taxaceae, whereas female
structures could be simple or compound showing an astounding
diversity. Gametogenesis is a slow process that usually ends until
pollination. Female gametogenesis consists of a coenocytical
megagametophyte development, followed by cellularization and
differentiation of two or more archegonia. Exceptions to this pattern are found in Gnetales
and Cupressaceae, where unique processes inside seed plants
are found. Male gametogenesis involves meiosis to form tetrads of microspores and
distinct number of subsequent meiosis before sperm differentiation. Sperm could be
free nuclei or celled with or without mobility. Anemophilous pollination is common,
but pollen transportation by insects is present in two lineages. Pollination mechanisms
involve secretion of pollination drop and distinct pollen morphologies. Fertilization is
predominantly siphonogamous but in basal gymnosperms, pollen tube development is
haustorial. Embryogenesis usually begins with acellular stages and the number of
mitosis before cellularization varies between groups. Polyembryony is feasible to
occur by two distinct processes but only one embryo reaches maturity. Modest
knowledge has been achieved relative to genetic basis of processes related with sexual
reproduction and morphological diversification of reproductive structures in
gymnosperms, warranting future research in these areas.
Introduction
All extant non-flowering seed plants are comprised in a group without
taxonomic rank: the gymnosperms. This group includes four divergent
lineages –Ginkgoales, Cycadales, Gnetales and Coniferales orders- with
morphologies highly similar to the fossil record from Mesozoic and a
worldwide distribution. Sexual reproduction is characteristic for this group,
since a few cases of asexual reproduction have been documented. Opposite to
angiosperms, reproductive structures of gymnosperms are typically unisexual,
although occurrence of bisporangiated cones have been reported for gnetales
and some conifer species. Reproductive units are called cones due to its conic
shape or strobilla; the female is the ovulated (seed) cone or the megasporangiated
strobile, while the male is the pollen cone or microsporangiated strobile. Cones
are considered simple when the structures that bear gametes are directly attached
to the central axe of the cone, or compound when the gametes bearing structures
borne at the axil of a bract, making up clustered complexes.
The term gymnosperm –naked seed- was assigned to this group because
the seed is not covered by a fruit as in angiosperms, although in some
lineages the seed is not properly naked, instead is covered by distinct
structures with apparent homology. Different molecular phylogenies support
the monophyly of this group, although due to deep divergence times and
lineage extinctions, taxa included in gymnosperms are highly differentiated
morphologically and the phylogenetic relationships between the four orders
are unstable. In the past, misinterpretation of morphological characters lead
to relate Gnetales with angiosperms and recently some molecular data place
Gnetales as sister group of all seed plants, challenging the monophyly of
gymnosperms [1, 2]. Even though, the majority of molecular evidence
supports gymnosperms as a natural group [3, 4]. However, lack of agreement
Sexual reproduction in gymnosperms 3
in relationships between the four basal branches difficulties to reconstruct
evolutionary history of morphological characters and processes involved in
sexual reproduction. Functionally unisexual reproductive structures,
unitegumented ovules, presence of three cell generations in seed, and
anemophilous pollination syndromes are common features for the group,
although exceptions to these patterns will be described in this chapter.
Life cycle
An overview of Pinus (Pinaceae) life cycle, illustrates the reproductive
process in gymnosperms, though variations to these pattern are found in
different species. Adult trees commonly bear both pollen and ovulates cones
(dioecius) and the age for first reproduction ranges from 6 to 20 years. Male
cones carry microsporophylls enclosing the microsporangium. This diploid
tissue originates pollen (microspores) through meiosis of diploid
microsporocytes (pollen mother cells). After a series of unequal cell divisions
the shed pollen contains four haploid cells (1n), including the generative cell
that in turn, originates sperm through mitosis. Whereas in female cones,
ovules develop adaxially to ovuliferous scales, each with only one
integument enclosing a megasporangium or nucellus (2n), which in turn
contains a megasporocyte or megaspore mother cell that undergoes meiosis
to produce four megaspores, of which three degenerate. The functional
megaspore divides into a multicellular megagametophyte (1n) containing two
or three archegonia each with one egg. Ovules produce a pollination drop
trough a small opening of integument (micropyle), where pollen arrives.
Pollen grains germinate developing a pollination tube that penetrates both
nucellus and archegonium to release its gametes in to the egg. Polyembryony
is feasible to occur due to embryo multiplication or multiple archegonia
fertilization, although, only one embryo will develop until maturity. Embryo
develops emerging from the seed as a seedling with multiple cotyledons [5].
Dioecious breeding systems without sexual dimorphism are common in
“basal” gymnosperms –Ginkgoales, Cycadales and Gnetales-, since male and
female structures borne in separate trees that are indistinguishable till they bear
such distinct reproductive structures. It seems that in Ginkgo (Ginkgoaceae) some
chromosomal differences are associated with sex determination, although
cultivated trees occasionally are monoecious [6]. In Gnetales abortive ovules
often are found in male cones therefore these plants are referred as gynodioecious
but, because ovules in hermaphroditic structures are unfunctional, also could be
referred to as dioecious plants [7]. Opposite, inside conifers monoecious systems
are common, with exception of Taxaceae family where exclusively dioecious
populations are found [8]. Inside Pinaceae, Araucariaceae and Podocarpace
4 Alejandra Vázquez-Lobo
families, some species have dioecious breeding systems. This phenomenon seems
to be related with ambiental factors more than with a genetic determination of
sex, because at different ages or ambiental conditions male or female trees might
develop both cone types.
Prevalence of monoecious breeding systems in basal gymnosperms and
anemophilous pollination syndromes promote outcrossing as mating system
in this group. In conifers, endogamy is avoided through different mechanisms
such spatial or phenological separation of female and male cones. In many
conifer species male cones are restricted to low branches whereas the females
growth only at highest branches (helping also to seed dispersal by air).
Phenological separation includes species where young trees produce only
pollen cones and older trees ovulated cones and species in which distinct
reproductive structures are mature at different seasons. Moreover there is
evidence of genetic post-fertilization systems to avoid the development of
autogamous embryos as in Pinus, where a genetic overdominant lethal
system stops the development of embryos produced by self-fertilization [9].
Reproductive cycle, from ovule differentiation to seed maturity, occurs
over a time span of one or two years. Exceptionally in Ephedra
(Ephedraceae) reproductive cycle is shorter than a year and some species of
Cycadales, Pinaceae and Cupressaceae delay until three years to complete the
reproductive cycle [6]. Trees can live many years, indeed the oldest living
trees are conifers, such as Pinus longaeva, Pinus aristata and Picea abies
(Pinaceae) that reach estimate ages of 8000 years old.

Perbanyakan tumbuhan paku dengan spora merupakan

metode perbanyakan yang paling umum dilakukan untuk
mendapatkan tanaman baru dalam jumlah besar. Meskipun
demikian, perbanyakan dengan spora hasilnya seringkali
tidak sesuai harapan. Keberhasilan perkecambahan spora
dipengaruhi oleh berbagai faktor, antara lain media tumbuh,
kemasakan spora, air, kelembaban, aerasi, dan derajat
keasaman (pH). Media tumbuh yang baik akan
menyediakan lingkungan yang baik pula bagi
perkecambahan spora serta pertumbuhan bibit (Jones,
1987; Toogood, 1999).

Introduction
Pteridophytes, also known as ‘vascular cryptogams’ and
‘ferns and fern allies’, comprise about 12 000 species of
vascular plants that do not produce flowers or seeds,
reproducing instead via the production of spores. Pteridophytes
occur in most terrestrial habitats on earth and
are also present in some aquatic communities. They are an
important part of the ground vegetation in many forest
communities and, with about one-third of the species
growing on the trunks and branches of trees, they are also
an important component of many epiphytic plant communities.
Some species are very beneficial to humans, but
the group also contains some of the most important weed
species in the world.
Life Cycle
Pteridophytes are characterized by a life cycle that usually
involves an alternation of two free-living generations –
sporophyte and gametophyte – with the sporophyte the
larger phase of the life cycle. Nonvascular plants like
mosses and liverworts also have an alternation of generations,
but in these organisms the gametophyte generation is
generally the dominant phase. In seed plants, the
gametophyte is no longer free-living but remains enclosed
in tissues on the sporophyte and there is a progressive
reduction in the size through various gymnosperm groups
such that in flowering plants (angiosperms) the gametophyte
generation is reduced to just a few cells in the
germinating pollen grains and the ovules.
The conspicuous phase of the pteridophyte cycle is the
sporophyte, which is how most people observe the plants in
nature. These are usually perennial. Sporangia are
produced on the leaves of sporophytes (sometimes in
specialized cone-like strobili). In true ferns, these are
commonly on the leaf undersurface and are often clustered
into discrete units called sori. Within each sporangium,
specialized cells undergo a series of mitotic (structural)
divisions followed by meiosis (sexual division) that results
in production of spores with half as many chromosomes as
in the original sporophyte. The more advanced ferns
usually have 64 spores per sporangium, but more primitive
ferns and fern allies may have hundreds or even thousands.
At maturity, the sporangium dries and ruptures, dispersing
the spores into the air.
When a spore lands on a suitable substrate, it
germinates, the cells dividing and forming first a filament
and eventually usually a heart-shaped gametophyte (sometimes
other shapes in some groups) that is the same species
as the sporophyte but appears very different. Gametophytes
are often moss-like in appearance and are quite
small, usually less than 1 cm wide at maturity, but are often
fairly easily located in nature near adjacent sporophytes.
Although in a few genera gametophytes can be long-lived,
in most ferns their lifespan is usually much less than a year.
They are the sexual phase of the life cycle in that they
produce multicellular sex organs at maturity on the side
away from the light. The more or less spherical antheridia
(male gametangia) are produced among the rhizoids
towards the base of the plant and at maturity they pop
open to release motile flagellated spermatozoids. Archegonia
(female gametangia) are usually produced at the
opposite end near the notch region, and are flask-shaped
structures containing a single egg cell. A film or droplet of
free-standing water is necessary in order for the spermatozoids
to swim to an archegonium of the same or a
different gametophyte. The neck cells of the archegonium
spread at maturity and the spermatozoid swims down the
archegonial canal to fuse with the egg, effecting syngamy
(fertilization) and forming a zygote with twice the number
of chromosomes as the gametophyte. This zygote grows
and develops into a new sporophyte, completing the cycle,
while the maternal gametophyte withers away.
In a typical pteridophyte, each gametophyte is potentially
bisexual, producing both antheridia and archegonia.
Because the eggs and spermatozoids of an individual all
grew from a single spore and are thus genetically identical,
potentially these plants can become self-fertilized in a way
that renders the resultant sporophyte entirely homozygous
(having only one kind of allele for each gene locus) for its
entire makeup. Various mechanisms exist to promote
cross-fertilization: the gametangia often mature at different
times; the genome may have deleterious alleles that are
Article Contents
Introductory article
fatal to homozygous individuals (genetic load); and there
may exist mating factors that prevent successful selffertilization.
Some ferns also produce pheromones known
as antheridiogens, in which the first spore to germinate at a
site becomes a female gametophyte and exudes a substance
causing later-germinating spores to develop into male
gametophytes.