EVOLUTION OF STELE IN PTERIDOPHYTES

INTRODUCTION:

Stele is the central cylinder or core of vascular tissue in higher

plants and pteridophytes. It consists of xylem, phloem, pericycle and medullary
rays and pith if present.

The term stele has been derived from a Greek word meaning rod
or column. Van Tieghem and Douliot (1886) introduced this term and put
forward the stelar theory. The theory suggests that the cortex and stele are
the two fundamental parts of the shoot system. Both these components (stele
and cortex) separated by endodermis.

Tieghem and Duoliot recognized only three types of steles. They

also thought that the monostelic shoot were rare in comparison of polystelic
shoots.

But now it is an established fact that most shoots are monostelic and
polystelic condition rarely occurs. The stele of the stem remains connected
with that of leaf by a vascular connection known as the leaf supply.

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 EVOLUTION OF STELE IN PTERIDOPHYTES

TYPES OF STELES:

There are several types of stele found in pteridophytes

which are:

1. Protostele
2. Siphonostele
3. Solenostele
4. Dictyostele
5. Polycyclic stele
6. Eustele
7. Polystele

Protostele:

Jeffery (1898), for the first time pointed out the stelar theory
from the point of view of the phylogeny. According to him, the
primitive type of stele is protostele.

In protostele, the vascular tissue is solid mass, and the central core
of the xylem is completely surrounded by the strand of the phloem. This
is the most primitive and simplest type of the stele.

 Pith is absent in protostele i.e., it is non- medullated.

 Majority of the pteridophytes show protostelic

conditions in their rhizome, stem or roots.

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 EVOLUTION OF STELE IN PTERIDOPHYTES

There are five forms of protostele:

1. Haplostele
2. Actinostele
3. Plectostele
4. Mixed Protostele
5. Mixed protostele with pith
a)Haplostele:
This is the most primitive type of protostele. Here the
central solid smooth core of xylem remains surrounded by uniform
layers of phloem
E.g., Selaginella, lycopodium.

b) Actinostele:
 Xylem is star shaped

 The phloem is not present in continuous

manner.
 Phloem occurs as separate patches between arms of the xylem E.g.,
Psilotum

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 EVOLUTION OF STELE IN PTERIDOPHYTES

c) Plectostele:
This is the most advanced type of protostele. Here the central core of the
xylem is divided into number of plates arranged parallel to each
other.The phloem alternates the xylem.

D. Mixed protostele:
Xylem divided into several units or groups.
Each xylem units are scatteredly arranged inside the ground mass
phloem.
E.g., Lycopodium cernuum

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 EVOLUTION OF STELE IN PTERIDOPHYTES

E)Mixed protostele with pith;

Here the xylem elements are mixed with the small patches
of parenchymatous cells of the pith. This type is found in primitive
fossils and living ferns. They are treated to be transitional types in
between protosteles and siphonostele.
E.g., Lepidodendron.

Siphonostele:

This is the modification of protostele. A stele in which the

protostele is medullated or with pith at the centre is known as
siphonostele. Such stele contains a tubular vascular region and a
parenchymatous central region i.e., the central core of pith is surrounded
by the xylem.

 It is advanced than protostele.

Origin of siphonostele:

There are different views among the scientists regarding the origin of
siphonostele. But they all agreed that, the siphonostele is originated from
protostele by the formation of pith in the centre. Here, the centrally placed
xylem core is replaced with parenchymatous pith.

Different stages of changing protostele to siphonostele can be observed

in the T.S at different levels in Osmunda.

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 EVOLUTION OF STELE IN PTERIDOPHYTES

There are two views regarding the origin of pith in siphonostele.

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 EVOLUTION OF STELE IN PTERIDOPHYTES

Jeffery’s theory:

According to this theory, the pith is formed as a result of the

invasion of cortical parenchymatous cells into the stele. The invasion of pith
occurs through the leaf gap or branch gap.

Thus pith and cortex are homogenous structures according to this

theory.

This theory is not accepted by most of the authors since in many

pteridophytes there is stele without leaf gaps but having siphonostele.

Boodle’s theory:

According to the theory proposed by boodle(1901), and Gwynne

Vaughan, the siphonostele has been evolved from the protostele by a
transforamation of the inner vascular tissue into parenchyma.

A siphonostele may be of the following types:

a) Ectophloic siphonostele:
In this type of siphonostele, the pith is surrounded by concentric
xylem cylinder and next to xylem the concentric phloem cylinder.

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b) Amphiphloic siphonostele:

In this type of siphonostele the pith is surrounded by the vascular

tissue . The concentric inner phloem cylinder surrounds the central pith. Next
to the inner phloem is the concentric xylem cylinder which is immediately
surrounded by outer phloem cylinder.

E.g., Marsilea

Solenostele:

In solenostele, pith is found with one leaf gap. It may be ectophloic

or amphiphloic solenostele.

a) Ectophloic Solenostele:

Xylem is surrounded by only on the outer side by phloem

E.g., Osmunda.

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 EVOLUTION OF STELE IN PTERIDOPHYTES

b) Amphiphloic Solenostele:

In the centre, pith is found. Xylem is surrounded on both sides of

phloem.

e.g., Marsilea rhizome.

Dictyostele:

Solenostele that is broken into a network of separate vascular

strands are called dictyostele. This breaking up of stelar core is due to the
presence of large number of the leaf gaps.

Each such separate vascular strands is called meristele. Each meristele

is of protostelic type. The dictyostele with many meristeles looks like a
cylindrical meshwork.

E.g., Pteris.

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Polycyclic stele:

This type of stelar organization is the most complex one

amongst all vascular cryptogams(pteridophytes). Such type of steles are
always siphonostele in structure.

A typical polycyclic stele possesses two or more concentric rings of

vascular tissue. This may be a solenostele or a dictyostele. Two concentric
rings of vascular tissue are found in pteridium aquilium

a) Polycyclic solenostele

b) Polycyclic dictyostele

Eustele:

According to Brebner (1902), there is one

more modification of the siphonostele known as eustele. Here the vascular
system consists of a ring of collateral or bicollateral vascular bundles situated
on the periphery of the pith.

E.g., Equistem.

 It is the modification of ectophloic siphonostele.

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 Splitting takes place because of the overlapping

leaf gaps.

Polystele:

More than one stele in the axis of the pteridophytes is present known as
polystele. It is type that must have derived from protostele because each
stele shows protostelic condition.

E.g., Selanginella.

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Other modification of siphonostele:

1. Cladosiphonostele: No leaf gaps present. It is the simplest type

of siphonostele. And found in several species of selaginella.

2. Phyllosiphonic siphonoste: Siphonostele that remain perforated

by smaller or larger leaf gaps caused by leaf traces. Member
Filicophyta.

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CONCLUSION:

In the vascular plants, the stele is the central part of the root or stem
containing the tissues derived from the procambium. These include vascular
tissue, in some cases ground tissue(pith) and a pericycle, which, if present,
defines the outermost boundary of the stele. Outside the stele lies the
endodermis, which is the innermost cell layer of the cortex.

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REFERENCE:

 Foster, A.S. and Gifford,E.M.(1974)comparative morphology of

vascular plants

 Gifford, Ernest M and Foster, Adriance S (1988) Morphology and

evolution of vascular plants.

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