Pteris:

Plants are usually terrestrial or epilithic. Rhizome is short, erect or short-creeping, slender to
massive, scaly generally towards apex, scales basifixed, brown or black, linear lanceolate, margin
entire or variously lobed. Solenostelic or dictyostelic or sometimes polycyclic. Stipes (petioles)
arise close together, well developed, adaxially grooved, colour straw or brown, generally scaly at
base, naked upwards, vascular bundles either one (in a cross section ‘U’ or ‘V’ or omega-shaped),
or less often 2 C-shaped vascular bundles, or in large species sometimes with two larger adaxial
and a semi-circle of smaller abaxial ones.; rachis (mid-rib) grooved on upper side, sometimes
winged. Lamina rarely simple, 1-2 rarely up to 5-pinnate, ultimate divisions often pinnately lobed
or pinnatifid, lamina occasionally pedate or palmate, never finely dissected, texture herbaceous or
coriaceous, glabrous or less often hairy rarely scaly when mature; lowest pair of pinnae often
forked near the base on basiscopic side resulting in a long pinnule with its lobes similar to other
pinnae, however there are many species that do not show such a forking of the lowermost pinnae
(or pinnule) eg. the very common Pteris vittata, P. excelsa etc., terminal pinna similar to lateral
ones; leaf axis rarely winged, adaxially grooved, the grooves with few exceptions continuous on
axes of different order; larger divisions sessile or short (rarely)–stalked, very often of an elongate-
oblong to lanceolate-linear,usually symmetric, upper divisions lobed-confluent. Stomata
polocytic and anomocytic. Veins free or anastomosing to form a series of narrow areolae along
the costa or costules; areolae with or without included veinlets; costa and costules usually with
short setae on the upper surface, Sori marginal (actually submarginal as the leaf margin curls back
to form the indusium), linear; pseudoindusia formed by reflexed pinna margin. Spores brown,
tetrahedral trilete rarely bilateral monolete, non-perinate, exine smooth or tuberculate or
verrucose, the two surfaces have different types of ornamentation, a prominent collar-like ridge
girdles the spores,. perine absent. Chromosome base number x=29.
Gametophyte Generation:
The spore, which is the first stage of the gametophyte generation, is a living cell, protected by a
thick wall and is the main source of population dispersal. The spores of Pteris have an exine with
a species-specific ornamentation and are believed to retain viability long after shedding but
germinate readily under appropriate moist conditions. Germination is preceded by swelling of the
contents of the spore by absorption of water and the spore opens. A small colourless protuberance
the rhizoidal initial emerges out of the spore opening and is soon followed by another
protuberance full of chloroplasts, the protonemal filament initial. Through cross-walls
perpendicular to the long axis of the filament, a uniseriate protonemal filament composed of
barrel-shaped 2-7 chlorophyllous cells bearing one or a few rhizoids at the base, is formed. The
terminal cell of the protonemal filament undergoes divisions by vertical and transverse walls, at
right angles to each other resulting in a group of four cells, which by further divisions in two
planes form an anterior plate of cells leading to the formation of the spatula stage. An abrupt
change in the plane of cell divisions results due to formation of a prothallial plate (meristem). The
stage at which the meristem is formed differs in various species. In some cases a wedge-shaped
ephemeral apical meristematic cell may precede the characteristic pluricellular meristem.
Depending upon the position of the meristem and when it is established, the young prothallus is
asymmetric (lopsided) for alonger or shorter period. Further growth leads to the formation of a
cordate prothallus with a central cushion 3-5 or 5-10 cells thick. It is surface-living, green and
photosynthetic. The shape of the adult prothallus varies in various species of the genus Pteris. It
is tapering-bilobed type (narrowed gradually towards the base) in P. excelsa, P.
pseudoquadriaurita and P. stenophylla) but broad-winged with an almost posterior end in
P.vittata and P. wallichiana.
Archegonia are borne on cordate meristic prothalli invariably below the notch. Rhizoids are also
very abundant on the cushion. No trichomes or glands of any kind are present on the prothalli of
Pteris. Antheridia are produced from early stages of development of the prothallus and these are
present either on the central cushion below the archegonia or on the wings. Three kinds of
gametophytes are generally produced: (i) large-sized and generally fast-maturing cordate prothalli
with symmetrical wings, initially archegonial but later may bear antheridia (ii) relatively smaller
cordate prothalli, narrow below, usually protoandrous and during subsequent archegonial phase
hardly producing any fresh antheridia, (iii) slow-growing ameristic prothalli exhibit arrested
growth remain small, non meristematic and irregularly shaped but loaded with antheridia.
Sex organs: Development of Antheridia: The ontogeny of the antheridium in ferns has been a
subject of controversy. In the antheridial initial three walls are formed successively and according
to the classical concept (before 1951) the first and third wall were described as ‘funnel-shaped’
and the second as hemispherical and periclinal lying over the first wall. Davie in 1951 refuted the
classical concept and considered successive walls to be transverse initially and later bulge
downwards or upwards. Stone (1962) thought the first and in some cases, the third wall to be
funnel-shaped. Much emphasis was laid on the axis of the nuclear division and the differentiation
in cytoplasm prior to cytokinensis which determined the position in which each new cell wall is
deposited and in interpreting the formation of a funnel-shaped wall. Verma and Khullar (1965)
considered the first and third walls to be either straight transverse or slightly concave or deeply
concave and retaining their shape except for minor displacements. The second wall is
hemispherical, lying over the first wall with its convexity more or less running parallel to the
periphery of the antheridium in order to enclose the dense contents of the androgonial cell within
An antheridium inducing substance known as antheridogen or antheridiogen has been isolated
from the gametophytes (especially in Pteridium aquilinum) and from the culture medium in
which the gametophytes were grown. This substance stimulates and hastens the precocious
formation of antheridia by 3-4 weeks. Seven types of gibberellins had the same effect as
antheridiogen and so it is believed that the antheridiogen is a gibberellin. An extract from
predominantly archegonia bearing- prothalli exhibited more antheridium inducing activity than an
extract from juvenile, predominantly antheridium-bearing prothalli.
Archegonia: A superficial cell on the cushion acts as the archegonial initial. It divides by a
periclinal wall into an outer smaller primary neck cell and an inner cell. The latter divides again
by a periclinal wall cutting off a central cell and a basal cell, thus forming a tier of three cells. The
primary neck cell divides by two longitudinal walls at right angles to each other to form four
cells. These are the initials for four tiers of neck cells. By further successive periclinal divisions
of the neck cells, an elongated neck is organized. The central cell first cuts off a primary neck-
canal cell and then divide again to form a venter canal cell and an egg. The neck canal cell
divides to form two neck canal nuclei. At maturity the archegonium has a neck of four rows of
cells with 4-5 cells in each tier; an egg, a venter canal cells and two neck canal nuclei are present
in the center. Before fertilization all cells above the egg disintegrate. Spermatozoids are attracted
by the species-specific fluid oozing out of the opening of the archegonia, and fertilize the egg.
Embryogenesis: Although more than one egg on each prothallus is fertilized only one develops
into a young sporophyte, (A natural population control!). It is not known whether it is the first
formed or the subsequently developed archegonia that form the sporophyte. The first division in
the fertilized egg is by a cross-wall to the long axis of the archegonia followed by another at right
angles to the first resulting in the formation of a quadrant. Differentiation into the organs of the
young sporophyte may occur as early as the quadrant or octant stage. The outer anterior part of
the quadrant gives rise to the first leaf, the inner anterior to the shoot apex, the inner posterior to
the foot and the outer posterior to the first root.
Departures from the normal sexual life cycle (as described above) can result in the cytological
and/or morphological alternation of generations being interrupted or modified. Such examples
include apogamy, apospory and vegetative reproduction. Apogamy and apospory occur in ferns
often as sporadic aberrations, compounded or combined with other developmental errors.

Apogamy: (apo=without; gamy=marriage). In many homosporous ferns (about 10%) a

sporophyte is produced without gametic fusion (fertilization). This is known as apogamy. Many
species of the genus are apomictic and Pteris cretica, is the earliest and well known example. In
apomictic forms archegonia are not developed and any cell of the prothallus directly gives rise to
the sporophyte. In an apomictic life-cycle the genetic consequences of sexuality, recurrent gene
segregation and recombination are averted. In such cases the function of meiosis in halving the
number of chromosomes is neutralized by events during the premitotic mitoses which remains
incomplete at a particular stage, when a failure of anaphase separation followed by the formation
of restitution nuclei produce spore-mother-cells with double the number of chromosomes. Such a
shift in the breeding system could be through a suitable mutation in a sexual taxon or due to the
interaction of the two genomes of a species combined in a hybrid brought together in cross
fertilization. Apogamy can also be induced in laboratory conditions by sugar or ethylene
treatment.
Apospory: In apospory the gametophytic tissue is produced by the sporophyte without the
intervention of spores. The prothalli so produced are usually functional but may have a somewhat
different morphology from those of the same species produced through spores. It occurs
sporadically in ferns but can also be induced. Aposporous prothalli bear normal sex organs and
since they have the same chromosome number as the parental sporophyte tissue they provide a
means of inducing polyploidy.
Reproductive Biology:
Although homosporous ferns may have bisexual gametophytes, there is no evidence that these
gametophytes characteristically self-fertilize. It was earlier believed that due to the close
proximity of the gametangia on the prothallus, it underwent self-fertilization, where the sperms
from the antheridia swim to fertilize the egg. In contrast, electrophoretic data indicate that inter
gametophytic matings occur frequently. However studies have indicated that many homosporous
ferns possess mechanisms that promote out crossing in natural populations. Noteworthy of these
mechanisms is the production of the hormone antheridiogen (discussed above), which results in
the production of effectively unisexual gametophytes, thus promoting intergametophytic mating.
The sequence of gametangia formation on mersitic prothalli reveal that some species of Pteris are
either exclusively or mostly inter gametophytic in their mating system. Intra and inter
gametophytic mating or a combination of both the mating systems may be found in the
polyploids. Another interesting aspect is the inability of monoecious prothalli, upon isolation, to
bear a sporophyte. Through genetic analysis the existence of a weak self-incompatibility system
has been demonstrated. Several factors including self-incompatibility and genetic load play a role
in determining the infertility of monoecious prothalli in isolation or in pair-wise combinations to
produce sporophyte. The fern breeding systems are diverse rather than stereotyped. Several
questions on the genetic and epigenetic controls of their sexuality remain to be answered.