STELE

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• The conducting system of pteridophytes consists of xylem and phloem
and associated parenchymatous cells, all of which are organized into a
stele.
• The term stele has been derived from a Greek word meaning pillar or
column.
• The concept of the stele as the fundamental unit of vascular system was
put forward by Van Tieghem and Douliot (1886) who proposed and
developed Stelar theory.
• According to the stelar theory the primary structure of the stem and root
were fundamentally similar in gross anatomy because both consisted of a
central core, the stele, surrounded by the cortex.
• The term stele was interpreted as the vascular tissue and conjunctive
tissues associated with them and the pith and pericycle (if present).
• The stele of stem was connected with that of leaf by a vascular
connection known as leaf trace.
Types of stele in pteridophytes
On the basis of the kind of stellar organization present in different
pteridophytes, an evolutionary sequence can be recognized among different
groups. The stele in pteridophytes can be differentiated into two major groups
1.Protostele: It is the most simplest and primitive type of stele. In protostele ,
the vascular bundle is a concentric solid mass and the central core of xylem is
surrounded by a layer of phloem and finally surrounded by a layer of pericycle.

Jeffrey (1898) named primitive type of stele as protostele. It is considered

primitive because it is present in some earliest known land plants that
appeared about 400 million year ago. The protostele exists in following forms:
Haplostele, Actinostele, Plectostele, Mixed Protostele, Protostele with mixed
pith, The radial protostele.
a) Haplostele: This is the most primitive type of stele. In this the xylem forms a solid,
smooth, and spherical central core which is surrounded by a continuous concentric layer
of phloem e.g. Lygodium, Selaginella. This particular type of stele has been regarded as
the most primitive among the different types.
b) Actinostele : This is the modification of the haplostele and somewhat more advanced in
having the central xylem core with radiating ribs and phloem is not concentric but
present in between the radiating ribs of xylem e.g. Psilotum . The protoxylem is present
at the tips of radiating arms.
c) Plectostele: In the stem of some species of Lycopodium the solid core of xylem gets
broken in a number of plate-like lobes, more or less lying parallel to one another. The
phloem alternates with xylem plates e.g. Lycopodium volubile. This specialized form of
protostele is termed as plectostele.
d) Mixed protostele: In this type, masses of xylem and phloem are uniformly distributed.
Scattered groups of xylem are embedded in the ground tissue of phloem e.g. Lycopodium
cernnum.
e) Protostele with mixed pith: In the centre there is parenchyma cells associated alongwith
the tracheids e.g. Lepidodendron. It may be derived from the transformation of the
tracheids into parenchyma and may be first step in formation of pith. Such type of protostele
is termed as mixed-protostele.
f) The radial protostele: This stele consists of radial vascular bundles enclosed by an
endodermis layer. This type generally found in root and stem of young stage of some species
of Lycopodium
2. Siphonostele: A kind of stele in which there is pith in the central
region is called a siphonostele.
This is the modified form of the protostele which has developed
central pith hence often called medullated protostele.
In this type, xylem is in the form of a hollow cylinder, enclosing a
well defined central pith in the centre and surrounded by concentric
phloem.
This type of stele is thought to have been evolved from a protostele
by conversion of treachery elements into parenchyma.

Origin of pith in siphonostele: There is a general acceptance that

siphonostele is evolved from a protostele. Two theories have been
proposed to explain the origin of pith:
Intra –stelar origin of pith: According to this view, pith has originated as a
result of transformation of tracheary elements of the central xylem core into
parenchyma. Thus, the pith is totally intra –stelar. This view is supported by
Boodle (1901), Gwynne – Vaughani (1908), Bower(1911). e.g. Osmunda regalis,
Botrychium ternatum where tracheids are scattered throughout the pith (mixed
pith).
Extra –stelar origin of pith: This hypothesis was supported By Jeffrey (1902,
1910, 1917). According to this view, protostele has transformed into
siphonostele due to migration of cortical cells into stelar axis. Openings such as
leaf and branch gaps, probably provided passage for invasion of parenchyma.
This view derives support from amphiphloic siphonostele that has two
endodermal layers, one delimits the stele from the cortex and the other from
the pith. Since endodermis is a structure peculiar to cortex, the pith is
considered as cortical in origin.
Types of siphonostele: The distributional patterns of xylem and phloem, the siphonostele is
classified into following two types:
a) Ectophloic siphonostele: In this type, the central pith is surrounded by concentric
cylinders of xylem and phloem e.g. Equisetum. The phloem is present only on the outer side
of xylem.
b) Amphiphloic siphonostele: In this type, there is pith in the centre . The concentric xylem
is surrounded on both the sides (externally and internally) with phloem cylinder which intern
is followed by pericycle e.g. Marsilea . In its simplest form, the siphonostele has no leaf gaps.
This type of siphostele is termed as cladosiphonic siphonostele e.g. some species of
Selaginella .On the other hand a siphonostele with leaf gaps is termed as phyllosiphonic
siphonostele e.g. Marsilea.
c) Eustele: Here the hollow vascular cylinder with central pith gets broken into a
number of collateral bundles arranged in a ring. This type is also known as
polyfassicular siphonostele e.g. Equisetum.
d) Solenostele: This is modified form of siphonostele which get perforated by a single
leaf gap. The solenostele may be either ectophloic or amphiphloic e.g. ferns.
e) Dictyostele: A dissected siphonostele is known as dictyostele. In this type, the cylindrical
stele is interrupted by numerous leaf gaps that overlap each other. In the cross section, the
stele appeared as discrete stands of or bundles each with internal xylem and concentric
phloem and is called meristele e.g. Pteris.
f) Polycyclic stele: This type of stele is most complex one among all pteridophytes. A typical
polycyclic stele possesses two or more concentric rings of vascular tissue in regular rings or
cylinders one within another .This may be a solenostele or a dictyostele. Two concentric rings
of vascular tissue are found in Pteridium aquilinum and three in Matonia pectinate.
HETEROSPORY

 Some Pteridophytes which produce two different types of

spores (differing in size, structure and function).

 Such Pteridophytes are known as heterosporous and the phenomenon is

known as heterospory.

 The two types of spores are microspores and megaspores.

 Microspores are smaller in size and develop into the male gametophyte.
Megaspores are large and develop into female gametophyte.

 According to Rashid(1976), only 9 genera of pteridophytes show

heterospory,i.e., Selaginella, Isoetes, Stylites, Marsilea, Pilularia,
Regnellidium, Salvinia, Azolla and Platyzoma.
The origin of heterospory can be better discussed on the
basis of evidences from paleobotany, developmental and
experimental studies.

Palaeobotanical evidences:
It has been suggested that heterospory arose due to degeneration of
some spores in a few sporangia. As more nutrition becomes
available to less number of spores, the surviving spore grow better,
hence increase in their size.

A number of heterosporous genera belonging to the Lycopsida,

Sphenopsida and Pteropsia were known in the late Devonian and
early Carboniferous periods.

Lepidocarpon, Lepidostrobus, Mazocarpon, Plaeuromeia,

Sigillariostrobiis (members Lycopsid) Calamocarpon,
of
Calamostachys, Palaeostachys (members of Sphenosida).
2. Evidences from Developmental Studies:

In heterosporous Pteridophytes, the development of micro and

megasporangia follow the same pattern.
They have identical organization but for their size. While in
megasporangia most of the spore mother cells degenerate but in
microsporangia only a few mother cells are
disorganize.
In Isoetes, are only 50-300 megaspores
megasporangium.
there in
megasporangium contains In onlySelaginella erythropus
one megaspore which is
functional.
In Marsilea,Salvinia and Azolla
the phenomenon of
heterospory becomes distinct after meiosis.
In Marsilea 64 microspores and 64 megaspores are formed after
meiosis in microsporangium and megasporangium respectively.
 Biological Significance of
The phenomenon
Heterospory:
of heterospory is of great biological significance on
account of the following facts:

(i)The development of the female gametophyte starts while the

megaspore is still inside the megasporangium.

(ii)Same is true of microspores i.e., they also start germinating into male
gametophytes while they are still inside microsporangium.

(iii)The female gametophyte derives its nourishment from the sporophyte

i.e., female gametophyte is dependent on sporophyte for its nourishment.

(iv) The dependence of female gametophyte on sporophyte for its nourishment

provides better starting point for the development of new embryo than an
independent green prothallus which has to manufacture its own food .
Seed Habit in Pteridophytes:
The adoption of heterospory and the retention and germination of a single
megaspore within megasporangium to form a female gametophyte, led to the
phenomenon of “seed habit”, a characteristic feature of the spermatophytes.
A seed is that ovule which contains an embryo developed as a result of
fertilization.

The origin of seed habit is associated with the following:

(i) Production of two types of spores (heterospory).

(ii) Reduction in the number of finally to one per

megaspores
megasporangium.
(iii) Retention and germination of the megaspores and fertilization of the
egg.

(iv) Continued development of the fertilized egg into the embryo while
still in situ.
From the above observations it is concluded that the
life history of Selaginella approaches towards seed
habit because of the following features:

1.The occurrence of the phenomenon of heterospory.

2.Germination of megaspore inside megasporangium.

3.Retention of megaspore inside megasporangium either till the

formation of female gametophyte or even after fertilization.

4.Development of only one megaspore per megasporangium for

example, in Selaginella monospora, S. rupestris, S.
erythropus etc.