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Dr. Renu Jangid
 Stewart and Rothwell(1993) proposed six evolutionary events
that took place in ovule evolution after the onset of
heterospory:
1. Degeneration of three megaspore and formation of a single
functional megaspore in megasporangium.
2. Retention of the functional megaspore in the
megasporangium (nucellus) until embryo development.
3. Formation of endosporic megagametophytes within an
indehiscent megasporangium.
4. Elaboration of the apical part of nucellus to receive
microspores or pollen grains.
5. Formation of an integument which delimited a micropyle.
6. Formation of pollen tube or pollen tube like structure from
endosporic microgametophyte.
 In geological time, the evolution from
homospory to heterospory. During Late
silurian & Early Devonian, only
homosporous plants were present. Middle
devonian saw the advent of heterospory.
 Two kinds of spores were produced in
different sporangia. The microspors were
the more numerous and remained
approximately the same size, wherease the
megaspores numbered less and ultimately
to one per megasporangium.
 Most pteridophytes are homosporous and
monoecious (the spores produced by any
particular individual are all of equal size and
develop into gametophytes bearing both
archegonia and antheridia). With such an
arrangement, the probability of self-fertilization is
relatively high and evolutionary progress is
correspondingly slow.
 It may be prevented, or made less probable, if the
sex organs on a prothallus ripen at different
times(e.g. in fern prothalli, the antheridia
commonly ripen before the archegonia), or if there
is some incompatibility mechanism: but, in any
case, cross fertilization can only occur if there is
sufficient water for the spermatozoids to swim
from one prothallus to another.
 Some pteridophytes have escaped from this
gametophytic limitation by becoming
endosporic, i.e. the prothallus is retained
within the swpore wall, scarcely protruding
from it, even at maturity when the spore
cracks open. The prothalli of such
pteridophytes are invariably dioecious, i.e.
the antheridia and archegonia are on different
prothalli, thus completely preventing self-
fertilization. Such pteridophytes are also
heterosporous, i.e. the spores are of two
sizes megaspore and microspore.
 Seed is a ripened or fertilised ovule. A seed (ovule)
may be defined as an integumentted indehiscent
megasporangium.
 The ovule of gymnosperm consists of three basic
parts: (a)Outer protective layer-Integument,
(b)Middle protective layer- Nucellus, and
(c)Innermost female gametophyte in which female
gamete and later embryo develops.
 The evolution of ovule, from free sporing,
heterosporous condition involved:-
1.Reduction in number of fertile(functional)
megaspores(where only one is functional).
2.Elaborat of nucellar tissue (apex of the
microsporangium) to receive male gametophyte.
3.Differentiation of additional protective layer
intequment.
 Two theories are known which use fossil evidence to
show how seed and integument evolved.
 1. Nucellar Modification Concept (Andrews,1961):-
According this theory the Lagenostomalean
integument is derivable from the nucellus or
sporangium wall by its vascularisation and growth
and loobing of the apex. These concept makes use of
coenopterid fern megasporangium which exhibitstwo
megaspore in the upper half of the sporangium. The
lower half is sterile and is supplied by a single
vascular strand.Andrews opines that a further
reduction in the megaspore(from two to one) took
place. The surviving megaspore increase in size and
sinks towards the basal sterile part.
 The proliferation of the sporangial wall took place
and was accompanied by the division of the basal
vascular strand into two, one each going on the
two sides of megaspore in the nucellur tissue. The
fused nucellus and integument of lyginopterid
seeds gives support to this theory.
 The first stage of development is represented by
Rhynia type branching telome, some of them being
fertile bearing terminal sporngia.
 Gradual reduction of some axes and corymbose
clustering around one terminal sporangium gave rise
to a central sporangium encircled by a ring of
vascularized sterile telomes.
 Fusion of encircled telomes at the base would cover
the sporangium with vscularized integument.
 Concurrent to these changes, the tip of the
sporangium is modified for pollen reception and
reduction of number of megaspores from many to
one would give rise to an evolved seed.
 The integument may fuse with the nucellus (as in
lyginopterids) or may remain separate.
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