There occur variation in the actinosteles.

In the stems of some species of

Lycopodium, e.g., L. clavatum and L. volubile, when seen in a cross section the
xylem occurs in the form of small parallel bands alternating with the floem plates.
This specialised form of actinostele is usually termed as plectostele (Figure
25.10C)
In Lycopodium cernuum, the xylem when seen in a cross section appears in the
form of irregular groups that are embedded in the ground of mass of phloem. This
type of actinostele is called the mixed protostele (Figure 25.10D)
In certain pterydophytes, e.g., Hymenophyllum demissum, the protoxylem of the
protostele is surrounded by a complete ring of metaxylem
(1) Siphonostele: A kind of stele in which there is present a pith in the central region is called
a siphonostele or medullated protostele. This type of stele is said to have been evolved
from the protostele, and represents a stage in evolutionary advance. In siphonostele,the
vascular tissue are arranged n the form of a hollow cylinder, with a distinct pith in the
centre. It is found inthe stems of most members of the Filicophyta
(a) Origin of siphonostele: There is a general acceptance that the siphonostele has
evolved from protostele. Two theories have been proposed accounting the
phylogenetic origin of the pith: intrastelar theory and extrastelar theory.
Intrastelar theory, which is supported by Boodle (1901), Gwynne-Vaughan (1908),
Bower (1911), Petry (1914) and others, holds that pith originates by metamorphosis
of the inner vascular elements into parenchyma. Support for this theory is furnished
by a number of plants in which Botrychium virginianum, Botrichium lunaria and
Osmunda regals in which the central region of the protostele consists of both
tracheids and parenchyma cells.
According to the extracelar theory which was put forth by Jeffrey (1892, 1907,
1917), the pith is extra stelar in origin. The theory holds that the pith originated as a
result of the invasion of cortical tissues into the stele through the leaf gaps and branch
gaps in the course of phylogenetic development of the vascular plants. According to
this theory the pith is cortical in nature
(b) Types of siphonostele: According ti the distribution of the vascular tissuesm the
siphonostele has been classified into following two types:
(i) Ectophloic siphonostele
(ii) Amphiphloiv siphonostele
In the ectophoic siphonostele, the xylem is in the form of a hollow cylinder
surrounding a pith with the pholem occuring only outside the xylem (Figure 25.10E).
It is found in Equisetum and some ferns like Osmunda and Schizaea
In the amphiphloic siphonostele, the xylem forms a hollow cylinder enclosing the pith
with the phloem occuring both on the inner side and outer side of the xylem (Figure
25.10F). It is found in the ferns like Adiantum and Marsilea.
In the lower vascular plants, e.g. Lycopodium and Selaginella the leaf gaps are
absent. In these plants the stele shows a continuous ring of tissue. A siphonostele
which has no leaf-gap is termed cladosiphonic siphonostele. In ferns large leaf gaps
are found in the siphonostele. Such a siphonostele with the leaf gaps is known as
phyllosiphonic siphonostele.
Leaf-traces and leaf gaps. In the vasluar plants the vascular system is
continuous throughout the plant body and consequently there are connections in
between the vascular stands of leaves and stem. At each node of the stem, the
vascular tissues depat from the stem and pass out into the base pf the petiole as the
leaf trace leaving a break in the vascular tissue of it. Such an interruption in vascular
tissues of the stem is called a leaf gap. The leaf traces continue up through the petiole
and are broken into fine strands thus constituting the vein system of the leaf.
In some ferns there may be leaf gaps which do not overlap. A cross section
through the internode shows a continuous ring of vascular tissues. This particular type
of siphonostele in which there is no overlapping of gaps is termed solenostele. It may
be extophloic solenostele (Figure 25.10G) or amphiploic solenostele (Figure 25.10H).
In some ferns , the leaf gaps are large and extend vertically so that they do not
overlap. As a result of overlapping of the leaf gaps the vascular tissues of the stem
appear as scattered series of vascular bundles. This type of siphonostele is known as
dictyostele or dissected siphonostele (Figure 26.10I). In this type the vascular strands
are interconnedted and form acylindrical meshwork. When seen in a cros section this
type of stele shows the presence of separate strands called meristeles. Each such
strand or meristele is concentric in structure consisting of a central strip of xylem
 surrounded by phloem. Dictyostele is found in Pteris, Pteridium, Polypodium, and
Dryopteris.
In some pterydophytes, the gaps occur in the sele which are not associated with
the leaf-traces. Such gaps have been termed perforations. Therefore a dictyostele
having such gaps or perforations is termed perforated dictyostele.
In certain pterydophytes complex type of stelar structure is seen in which two or
more concentric rings of vascular tissues are present. This type of stele is known as
polycyclic stele (Figure 25.10J, K). In Pteridium aquilinum there are two concentric
ring of vascular tissues in which the inner ring forms a siphonostele and the outer ring
forms a dictyostele consisting of numerous meristeles (Figure 25.10K). In Matonia
pectina there are three rings of vascular tissues.

HETEROSPORY AND SEED HABIT IN PTERIDOPHYTES

The occurrence of two kinds of spores in the same plants is called the heterospory. Of the
two kinds of spores, the smaller are called the microspore and the larger are termed as
megaspore and are produced within the microsporangia and megasporangia respectively.
The production of two kinds of spore in heterosporous plants is difenitely related with a
differentiation in sex for from a microspore develops a male gametophyte and from a megaposre
of female gametophyte. The heterospory thus is associated with the sexual differentiation of
gametophytes. The phenomenon of heterospory is common in eight genera of pteridophytes.
There are Sellaginella, Isoetes, Stylites, Marsilea, Pilularia, Regnellidium, Salvinia, and Axolla.
Advantage of teherospory: In a homosporous pterydophyte the spores germinate on the
soil to produce gametophytes that are independent of the sporophytic plants. In doing so the
gametophyte of the homosporous pterydophyte with much vegetative embryo which is
dependent upon it. In a heterosporous pterydophyte each spore germinates within the sporangium
and give rise to a prothallus. The gametophyte which is greatly reduced, is retained wholly or
largely within sthe spore wall depending entirely upon the food deposited in the spores. As the
gametophytes of heterosporoud pteridophyte obtain their nourishment from the rarent
sporophyte, the are independent of external conditions than are the free living gametophytes of
hoosporous pterydophytes.
Heterospory and Seed Habit
The differentiation of spores into microspores and megaspores and their dependence on
the parent sporophytes for the nutrition are the certain features in the life cycle of Selaginella
which have been considered as the exxential pre-requisites for the formation of seeds
characteritstic of Spermatophytes. It is generally agreed that the seed plants arose from the
heterosporous vascular plants that insted of discharging the megaspore acquired the habit of
retaining it within the megasporangium.
In the seed bearing plants there are two-kinds of spore: microspores and megaspore
which grwo to form male and female gametophytes respectively. In these plants the single
megaspore (embryosac) is not shed from the megasporangium but is retained within it while still
attached to the mother plant. It germinates inside the megasporangium (nucellus) producing the
much reduced female gametophyte bearing the archegonia. Later the nucelus and the
gametophyte are protected by a covering or integument and the whole structure is known as an
ovule. The female gametophyte gets the nutrition for its development from the parent plants and
thus does not need to produce by its own effort. Aftre fertilisation the zygote whithin the ovule
gives rise to an embryo, the rest of the gametophytic tissue to the nutritive tissue or endosperm
and the integument thickens to form a seed-coat. This entire structure, i.e., the integumented
ovule is known as seed. It is detached from the parent plant and germinates to form a new plant.
Thus we find that for the production of seeds the following pre-requisites are essential
1. Production of two types of spores (heterospory)
2. Megasporangium (nucellus) does not open and the single megaspore within it germinates
to form the female gametophyte
3. Nucellus becomes invested by a coring or integuent which late forms a seed-coat
4. Within the nucellus if formed a linear tetrad of four haploid megaspore as a result of the
reduction division of the functional megaspore mother cell. Out of these four megaspores
the lowermost gives rise to the female gametophyte whereas the rest degenerate
5. The male gametes reach the egg by means of a tubular outhgrowth of the male
gametophyte known as pollen-tube
6. Fertilisation and formation of embryo take place within the megasporangium
7. The embryo udergoes a resting period
Selaginella exhibit a remarkable approach to the seed habit characteristic of the
spermatophytes because of the following feature:
1. The heterospory occurs in almost all the species of Selaginella
2. In most species only one functional megaspore mother cell is produced which by
reduction division produce four haploid megaspore. In some species, e.g. S. Rupestris
and S. Monospora, only one megaspore is formed in each megasporagium and this
single megaspore in not shed but germinates to form the female gametophyte
3. The fertilisation and development of embryo in both the species take place while the
megaspore is enclosed with the megasporangium
Therefore, it becomes evident that Selaginella has considerably advanced towards the
seed habit in a few species but its approach to the trus seed is not complete due to the
following features:
(a) The megasporangium lacks an integument or covering
(b) The retention of the megaspore permenantly within the megasporangium has not
become established
(c) After the development of the embryo there is a lack of resting period.