ANGIOPTERIS
The Marattiopsida contain the single order Marattiales and family Marattiaceae,
consisting of six genera: Angiopteris, Christensenia, Danaea, Eupodium, Marattia,
and Ptisana .
Angiopteris is a genus of huge evergreen ferns from the family Marattiaceae, found
throughout the paleotropics from Madagascar to the South Paci c islands.The family
Angiopteridaceae has seven genera and nearly 145 species. All are tropical plants
occurring in humid forests. Of the seven genera Marattia and Angiopteris are widely
distributed.
Angiopteris ourishes in the tropical forests and is represented very well in the
eastern hemisphere. The number of species in Angiopteria is variable. While some
people-recognise only one variable species (A. evecta) others recognise over a
hundred species. Angiopteris evecta has been introduced and naturalised in Hawaii,
Jamaica, and parts of Central America, where it has become an invasive weed in
lower elevation drainage A. evecta is a common species found in India.
CLASSIFICATION:
Kingdom: Plantae
Clade: Tracheophytes
Division: Polypodiophyta
Class: Polypodiopsida
Order: Marattiales
Family: Marattiaceae
Genus: Angiopteris
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Economic Importance:Angiopteris are cultivated as ornamentals and also as food
(from edible stems). Perfumes and oils are extracted from these, and it is also an
alcoholic drink extracted from stem starch..
Sporophyte of Angiopteris:
Morphology of the plant/ External features:
The sporophytic plant body consists of an upright, tuberous, conical, eshy
rhizomatous stem. The stem is quit thick and inhabit the plant resembles a tree fern.
The stem is often called a caudex or trunk which may be a foot or two in height and
almost the same girth
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The upper surface of the rhizome bears a crown of graceful, stately leaves. The leaves
are 5-6 metres long in a luxuriously growing plant, with a petiole as thick as a man’s
arm. The leaves are typically bi-pinnately compound. The venation is of the open
dichotomous type. Pinnae are glabrous (smooth).
It is very typical of Angiopteris and other marattiaceous ferns to have a pair of thick
eshy stipules at the base of the leaf. When the leaves fall off, these persist with the
leaf bases and form a protective armour around the stem. Leaves pinnately divided,
pulvinate (enlarged or swollen at attachment point of lea ets) in living genera, and
with well-developed, eshy stipules (appendages at leaf base); sporangia
eusporangiate, in sori, or more or less coalescent in synangia (clusters);
homosporous; mostly massive, eshy ferns.The genera are distinguished mainly by
the disposition of the sporangia (spore-producing structures), which occur on the
lower side of the leaves. Angiopteris has separate sporangia.
The base of the petiole and the stipules together appear like a horse’s hoof. Roots are
produced from the under surface of the rhizome at the base of each leaf. While leaves
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are deciduous, roots are perennial. They are thick and have a mycorrhizal association.
The branching of the root may or may not be profuse.Roots are endogenous in origin
and pierce the stipules during the course of growth. Root hairs are peculiar in being
multicellular. At the margins of the pinnae on the abaxial surface are borne, the sori.
There is no distinction into foliage leaves and sporophylls. The sori occupy a near
terminal position on the dichotomously branched veins.
In some species (A. evecta), at the base of the pinnae and the petiole there are
swollen articulations (joints) which are apparently the regions of abscission. The
pinnae are long (more than 10 – 12 cm long), dorsiventrally attened and have a long
drawn tip. The margin is serrate. A. evecta feature a large, erect, woody rhizome with
a wide base supported by thick roots. The fronds are deltoid, pinnate, 5–8 metres (16–
26 ft) long, with spreading lea ets.
Angiopteris is unique among ferns in having explosively dispersed spores, thought to
be caused by the cavitation of an airspace between spore layers.The
basal chromosome number for this genus is 2n=80. The type species is Angiopteris
evecta.
Internal Structure/Anatomy:
Stem/Rhizome:
A transverse section of the stem of Angiopteris shows an outer epidermis, a broad
cortex and a central region of vasculature. The cortex is parenchymatous and
mucilage ducts are seen. The vascular cylinder of the rst formed part of a stem is
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protostelic. At the level of the third or fourth leaf the stele changes into a siphonostele
and then to a dictyostele.
A number of concentric rings of meristeles which, in a dissection are seen to be part
of a series of complex and irregular mesh works lying one within the other, yet
interconnected by 'reparatory strands'. The whole system may be described as a
highly dissected poly- cyclic dictyostele.
Although each meristele is surrounded by an endodermis, in the adult state the
endodermis is completely lacking.
These concentric circles represent the funnel shaped zones of anastomosing bundles.
Mattenius (1986) who made a detailed study of vasculature in Angiopteris
observes. “The vascular bundles form funnel shaped zones with the lower ends in
the stem, and their upper portions continued into the leaves as leaf traces.
Segments of the outer zone pass into the leaves as leaf traces and the gaps thus
left are lled by segments of the next inner zone”.
According to Campbell (1911), the concentric, vasculature of the stem is built up by
the union of leaf traces.
A very weak secondary development may occur in some cases. Tracheids have
scalariform pitting.
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Petiole:
In Angiopteris petiole shows much the same structure as in the stem. There is
palisade parenchyma below the upper epidermis and a loose spongy parenchyma
above the lower epidermis. The leaf trace may be C-shaped.
Leaf:
Lamina shows a prominent midrib. There is a palisade parenchyma below the upper
epidermis and a loose spongy parenchyma above the lower epidermis. Leaves are
generally hypostomatic.
Root:
A transverse section shows an epidermis, cortex and stele. The cortex has
parenchyma as well as sclerenchyma. There is a conspicuous endodermis encircling
the stele. Stele is an exarch, polyarch, actinostele. In the young roots a mycorrhizal
fungus is present.
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Reproduction:
Sporophyte reproduces vegetatively as well as by spore production. Dormant buds
are formed on the rhizome where the stipules join the petiole. When the stipules are
ultimately shed from the plant, the buds develop into new individuals.
Spore Producing Organs:
Spores are produced in sporangia which aggregate to form the son. The sporangia as
has already been said, lie at the back of the veins. The sporangia in the sori are
arranged in two linear rows. They may exhibit a certain degree of lateral fusion.
But this is not suf cient to call it a synangium. The synangial nature in Angiopteris is
not as evident as in Marattia. In fact, many people regard the sporangia of
Angiopteris completely independent. There is no typical induciuim. Rarely indicial
hairs may be present.
The rst indication of a sporangial development is the accumulation of cytoplasm in
certain epidermal cells lying in localized regions on the abaxial surface of the pinnae.
This forms the receptacle, from which sporangia develop. There is usually a single
sporangial initial, which undergoes periclinal division to form upper and lower cells.
While the outer cell contributes to the jacket, the inner cell gives rise to the
archesporium. However a part of the jacket (basal portion) is derived from the
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surrounding cells. Hence the development is of the eusporangiate type. A tapetum
differentiates from the innermost wall layer.
A nearly mature sporangium has a multilayered jacket enclosing a mass of spore
mother cells, which undergo reduction division and produce haploid spores. Spore
output varies from 1450-7500, per sporangium. All spores are of the same type. All
the sporangia in a sorus mature together (simple sorus).
Dehiscence of the Sporangium:
In a mature sporangium there is an arch of cells in the wall, with slightly thickened
end walls. This (annulus) extends from side to side across the top of the sporangium.
By the shrinking of these cells and those in the wall of the sporangium away from the
centre of the sorus, the wall is torn open towards the centre. Spores come out in a
mass. They are wind dispersed.
Gametophyte of Angiopteris:
Structure and Germination of Spores:
Germination of the spores is rapid, occurring within a few days of being shed, and
they develop directly into a massive dark green thalloid prothallus, which is
mycorrhizal and is capable of living for several years. An old prothallus may be
several centimetre long and may resemble closely a large thalloid liverwort .The
prothallus is monoecious but, while the antheridia occur on both the upper and lower
surfaces, the archegonia are con ned to the lower surface only, where they occur on
the central cushion along with rhizoids. Both types of gametangia are sunken beneath
the surface of the prothallus and the antheridium is large and massive. The
archegonium has a large ventral canal cell (except in Danaea) and a neck canal cell
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with two nuclei. The antherozoids are coiled and multi agellate, as in other ferns.
The structure and development of antheridium and archegonium are similar to that of
Ophioglossum. In Marattia the ventral canal cell is distinct and not transitory as in
Ophioglossum. Interesting feature rarely found else where and presumably associated
with its massive structure.
Angiopteris is homosporous. The spores are minute and usually tetrahedral in shape.
They germinate a few days after shedding. During the rst month there is
considerable enlargement in the size of the spore together with the appearance of
chlorophyll (Eames 1964). The rst division of the spore is transverse.
Of the two cells, the smaller one forms the rst rhizoid. The outer wall of the spore
ruptures facilitating the exit of the rhizoid. The larger cell forms a cluster of cells in
which is differentiated an apical cell. Soon a plate of cells is formed. Subsequent
growth by marginal divisions results in the formation of a thalloid structure.
The Mature Pro-thallus:
The pro-thallus is a deep green, aerial (surface living), lobed or cushioned, thalloid
structure . There is a de nite growing point. The thallus has dorsiventral symmetry.
Rhizoids are produced from the ventral surface and help in anchorage and absorption.
In spite of the presence of chlorophyll, a mycorrhizal association is always present.
To what extent this association is helpful to the thallus is doubtful.
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All the cells in the pro-thallus are parenchymatous.
Reproduction:
Prothalli of Angiopteris are monoecioius, with archegonia towards the cushioned
region and antheridia scattered all over. In structure and development antheridia are
similar to those of Ophioglossum. Antherozoids are multi-ciliate.The archegonia also
are similar to those of Ophioglossum but have a shorter, broader neck. The neck is
almost sunk in the gametophylic tissue. There is a single, (bi-nucleate) neck canal
cell. The venter canal cell is also large and conspicuous. Fertilisation is similar to that
in other pteridophytes.
Embryogeny:
The rst division of the zygote is transverse to the long axis of the archegonium.
According to Eames (1964) quadrant divisions are obscure and the young organs are
not de nitely traceable to these. Lang (1923) has reported the development of
suspensor from the epibasal half in some individual cases of A. evecta.Thus, the
embryogeny may be endoscopic. In any case it is evident that the typical embryonal
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parts like stem, root and cotyledon arise from the hypo-basal half of the embryo. No
de nite foot is formed. Thus as a contrast to ophioglossaceae, the embryo has a
reverse orientation. The primary root is formed only after the embryo has attained a
considerable size. Only one sporophyte is normally formed on a pro-thallus.
Phylogeny of Angiopteris:
Angiopteris resembles both Ophioglossum and true ferns. While in habit and general
appearance it is close to the latter, in important structural features it appears to be
closer to the former.
Pinnately branched leaves showing circinate venation recall what is seen in true
ferns. The presence of stipules shows resemblance to Ophioglossum. But it has to be
pointed out that the stipules of Angiopteris are thick and paired unlike those of
Ophioglossum.
Anatomically Angiopteris occupies a position intermediate between Ophioglossum
and true ferns. While the bulk of soft tissue present in the stem is suggestive of
Ophioglossum, the presence of some mechanical tissue is indicative of true ferns.
In eusporangiate development, and output of spores Angiopteris is de nitely like
Ophioglossum. But the soral organisation not seen in Ophioglossum, is a true fern
character.
Embryogeny bears many resemblances to Ophioglossum. Within the family,
Angiopteris with no synangial arrangement is regarded as primitive when compared
with Marattia and others which show a synangial organisation.
References:
https://www.biologydiscussion.com/pteridophytes/angiopteris-occurrence-and-
phylogeny-botany/73641
https://uou.ac.in/sites/default/ les/slm/MSCBOT-503.pdf
WYONA DIETY LOBO
242B0T05
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