Useful Notes On Anthocerotopsida Order-Anthocerotales (4994 Words)

21/10/2017 Useful Notes on Anthocerotopsida Order-Anthocerotales (4994 Words)
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Useful Notes on Anthocerotopsida

Order-Anthocerotales (4994
Words)
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Here are your useful notes on Anthocerotopsida !
The class Anthocerotopsida (Anthocerotae) consists of a single order, the

Anthocerotales and a single family, the Anthocerotaceae, 6 genera and
301 species. According to Muller (1940), Reimers (1954) and Proskauer
the order Anthocerotales includes two families, (i) Anthocerotaceae and
(ii) Notothylaceae. The latter includes a single genus, i.e., Notothylas.
http://www.yourarticlelibrary.com/biology/useful-notes-on-anthocerotopsida-order-anthocerotales-4994-words/6919 1/23
However, according to top bryologists, there is only one family, i.e.,

Anthocerotaceae. About five or six genera are included in this family.
Image Courtesy : upload.wikimedia.org/wikipedia/commons/c/ce/Phaeoceros_laevis.jpg
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These genera are—Anthoceros, Phaeoceros, Aspiromitus, Notothylas,

Dendroceros and Megaceros. Four genera are universally recognised,
they are—Anthoceros, Megaceros, Dendroceros and Notothylas. This
group differs in many respects from the other Bryophyta.
However, the group is placed intermediate between Hepaticopsida

(Hepaticae) and Bryopsida (Musci). The group is considered to be very
important from the point of view of its morphology, because of its
intermediate position between the two important groups, the
Hepaticopsida and Bryopsida.
The most characteristic features of the group are as follows: The

gametophytic plant body is thalloid and dorsiventral. The rhizoids are
simple and smooth walled. Tuberculate rhizoids and ventral scales are
altogether absent.
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The tissues of the thallus are not differentiated. Air chambers and air-
pores are absent. Each cell of the thallus possesses a large chloroplast
and a conspicuous pyrenoid within it.
The antheridia are endogenous, i.e., they arise from the hypodermal cells
of the thallus on the dorsal side of it. The antheridia are developed within
the antheridial chambers, singly or in groups on the dorsal side of the
thallus.
The archegonia are found in sunken condition on the dorsal side of the
thallus.
The sporogonium arises from the dorsal side of the thallus. It is

elongated and cylindrical in structure. It consists of foot, meristematic
region and capsule. It possesses intercalary meristem, and continues its
growth throughout the growing season. The wall of sporogonium
contains chlorophyll. The central sterile portion is columella, which is
surrounded by sporogenous tissue and spores. The elaters are also
present.
The sporogenous mass develops from amphithecium and arches over the
columella.
Family-Anthocerotaceae:
There are five genera (i.e., Anthoceros, Phaeoceros, Aspiromitus,

Dendroceros and Megaceros) in this family.
Characteristic features:
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The sporogonium (capsule) is linear and vertical. Generally the stomata

are present on capsule wall. The archesporium arises from
amphithecium. The elaters are four-celled, smooth or thick-walled and
with or without thickening bands. The genus Anthoceros has been
discussed here in detail.
Genus ANTHOCEROS:
Habitat and distribution:
About 200 species of this genus are found throughout the world in
temperate and tropical regions. The species are moist and shade loving.
About 25 species have been reported from India, by various workers. The
three common Himalayan species are, Anthoceros himalayensis, A.
erectus and A. chambensis.
The above mentioned species are common in various hilly regions such
as, Mussoorie, Kumaon hills, Chamba valley and other places, 5,000 feet
to 8,000 feet. Some species have been reported from South India. All the
species are found in very shady and moist places. They are found in the
hollows of the moist rocks in dense patches. According to Cavers (1911)
some of the species grow on decaying wood. The species, are not at all
drought resistant.
External structure of the thallus:
The thallus is small, prostrate, dark green and dorsiventrally

differentiated. The thallus is lobed and the lobes are somewhat divided.
The mid-rib is not found. The dorsal surface of thallus of A. laevis is
smooth, whereas it is velvet like in A. crispulus (Bhardwaj, 1950) or it
may be with spines and ridges as in A. fusiformis.
In every case the smooth walled, simple rhizoids are found. The ventral
scales and tuberculate rhizoids are altogether absent. The thalli are dark
green, because of the presence of the Nostoc colonies, which may be
easily seen with the help of lens from the underside of the thallus.
Internal structure of the thallus:
The anatomy of the thallus is quite simple. Internal to the upper and
lower epidermis there are simple, parenchymatous cells. The cells of
parenchyma are isodiametric and uniform. Air chambers and air pores
are absent. Each cell contains a big chloroplast which possesses a single
pyrenoid in its centre. The chloroplasts are lens shaped. The chloroplasts
of the superficial cells are longer than the chloroplasts of the other cells.
According to McAllister (1914, 1927) the pyrenoids of Anthoceros are

quite different in structure from those of Chlorophyceae. On the ventral
side of the thallus certain intercellular mucilage cavities are found. These
cavities open by small openings, the slime pores on the ventral surface of
the thallus.
Sometimes, the colonies of blue green algal form-Nostoc, are found in the
mucilage cavities. There is no symbiotic relationship between these
Nostoc, colonies, and thalli. According to Peirce (1960) these colonies,
however, do harm to the thalli. The nucleus lies in the close vicinity of the
chloroplast, near the pyrenoid. Sometimes the chloroplast enfolds the
nucleus within it.
Apical growth:
The apical growth of Anthoceros is a controversial topic, whether it takes

place by a single apical cell or by a group of apical cells. According to
Smith (1955) the apical growth of the thallus is initiated by a single apical
cell.
According to Leitgeb (1879) the apical growth in this genus takes place by
several marginal apical cells. According to Mehra and Handoo (1953) the
apical growth in Anthoceros erectus and A. himalayensis takes place by a
group of cells situated in the depression at the apex of the thallus.
Reproduction:
The reproduction takes place by means of (1) vegetative and (2) sexual
methods.
1. Vegetative reproduction:
The vegetative reproduction takes place by various ways.
(a) By progressive growth and death of thallus:
The vegetative propagation takes place by progressive growth and death

of the older part of the thallus reaching dichotomy. But this method is
not so common in Anthoceros as in Riccia and Marchantia.
(b) By tubers:
In certain species of Anthoceros, the thallus becomes thickened at several

places on the margins. Such marginal thickenings are called, the tubers.
These tubers are perennating structures. They survive in the drought
conditions. On the advent of the favourable conditions, they develop into
new thalli. The tubers are formed in A. laevis, A, tuberosus, A hallii, A.
pearsoni and A. himalayensis.
(c) By gemmae:
In some of the species of Anthoceros, the gemmae have been found. The
gemmae have been recorded from the species, A. glandulosus, A.
formosae, etc. Each such gemma develops into a new thallus.
(d) By persistent growing apices:
According to Campbell the thalli of the species A. pearsoni and A.

fusiformis become completely dried up in summers, leaving growing
apices with adjacent tissues. These apices face the drought conditions.
On the approach of favourable conditions, these apices develop into new
thalli.
2. Sexual reproduction:
The species of Anthoceros may be homothallic (monoecious) or

heterothallic (dioecious). Some of the homothallic species are- A.
fusiformis, A. punctatus, Kashyap (1915, 1929) has recorded, A.
himalayensis as a dioecious (heterothallic) species, but Mehra and
Handoo (1953) have reported the same as monoecious (homothallic) but
protandrous. The heterothallic species are, A. pearsoni, A. halli, A.
erectus and others. The sex organs, i.e., antheridia and archegonia are
found embedded in the tissues of the dorsal side of the thallus.
Development of antheridium:
The antheridia are produced singly or in groups in the antheridial

chambers. The development is endogenous. Though the antheridium
develops from a superficial cell, yet it is enclosed within the antheridial
chamber which does not open out by any opening.
A dorsal superficial cell of the thallus, situated near the growing apex
divides periclinally giving rise to two daughter cells. According to Cavers,
Campbell and Haupt the superficial cell divides transversely and not
periclinally. The upper daughter cell acts as roof initial and the lower one
as antheridial initial.
Eventually, a mucilage filled space appears in between the roof initial and
antheridial initial. This mucilage cavity enlarges in size and ultimately
becomes the antheridial chambers. The roof initial is nothing to do with
the development on the antheridium.
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It divides and redivides several times anticlinally and periclinally giving

rise to a two layered roof of the antheridial chamber. Simultaneously, the
antheridial initial develops into a single antheridium or in a group of
antheridia. A single antheridium develops in A. pearsoni and sometimes
in A. himalayensis.
According to Mehra and Handoo (1953), in A. erectus a number of

antheridia develop in an antheridial chamber. Here, the antheridial
initial divides many times anticlinally producing many cells and each cell
thus produced, develops into an antheridium. The further development is
as follows:
The antheridial initial divides twice by vertical walls intersecting each

other at right angles, giving rise to four cells. This is followed by another
transverse division giving rise to two tiers of four cells each. The four
cells of the upper tier divide transversely giving rise to eight cells, the
octant stage.
All of the cells of octant stage divide periclinally giving rise to eight outer
primary jacket cells and eight inner primary androgonial cells. The four
cells of the lower tier develop into a stalk of the antheridium consisting of
the four rows of the cells.
The so formed androgonial cells divide repeatedly. The last generation of

these cells are androcyte mother cells. According to Bagchee (1924) each
androcyte mother cell divides diagonally producing two androcytes. Each
androcyte metamorphoses into a spindle like biciliate antherozoid.
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Structure of mature antheridium and its dehiscence:
The mature antheridium is stalked and club shaped. The stalk of the
antheridium may consist of the mass of cells, e.g., A. laevis, or it may
consist of the four rows of the cells, as in A. erectus and A. punctatus. The
antheridium proper is covered by a single layered jacket. Inside the
jacket, there are numerous androcytes which are to metamorphose into
antherozoids.
On the maturation of the antheridium, the roof of the antheridial

chamber disintegrates, with the result the antheridia are exposed to
outside. Soon after, the antheridia absorb water and burst at their apical
ends, giving way to the antherozoids to move outside. Sometimes the
androcytes come out in the form of an opaque mass at the opening of the
antheridium. Within few minutes they metamorphose into the
antherozoids.
The antherozoids:
The antherozoid is spindle like and biciliate. The cilia are attached to the
anterior end of the body. Sometimes just near the attaching point of the
flagella to the body, the blepharoplast is visible. The antherozoids swim
in the water by the lashing moment of their flagella.
Development of archegonium:
The archegonia are found embedded in the thallus. They remain in direct
contact of the vegetative cells of the tissue of the thallus lateral to them.
They do not possess jacket sterile cells around them. The development of
archegonium begins from a single superficial cell. This cell becomes

prominent and acts as archegonial initial.
According to Mehra and Handoo (1953), this has been established that
the archegonial initial functions directly as a primary archegonial cell.
Formerly it was believed (Campbell) that it divides, and produces two
cells, the primary archegonial cell and primary stalk cell. The archegonial
initial first divides vertically, producing three jacket initials which
surround an axial cell.
The axial cell divides transversely, producing a cover initial and a central
cell. Thereafter, the central cell divides by a transverse wall, giving rise to
a primary canal cell and a primary venter cell. The primary canal cell
divides repeatedly, producing a linear file of 4-6 neck canal cells. The
primary venter cell divides once transversely, giving rise to two cells, a
ventral canal cell and an egg (oosphere).
Comparatively, the neck canal cells are narrower than the venter canal
cell and the egg. Eventually, the three jacket initials also divide by
transverse walls. The further development of the jacket layer is not at all
clear. Since the cells on the lower face of the egg have been derived from
the archegonial initial they cannot be treated as a part of archegonium.
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Structure of archegonium:
On the maturation of the archegonium, the venter canal cells and neck
canal cells become gelatinized. Thus a mature archegonium is flask-like
in shape, without neck canal cells and with an egg (oosphere) in its
venter. At the top of the neck of the archegonium there are four cover
cells, which become separated from the archegonium, as soon as the
gelatinization of the venter and neck canal cells is over.
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Fertilization (syngamy):
Prior to fertilization, the cover cells become detached from the

archegonium, and the neck canal cells become gelatinized. Through the
medium of water, the antherozoids enter the mouth of archegonium. The
antherozoids are attracted chemotactically. Ultimately, one lucky
antherozoid penetrates the egg, and the fertilization is effected. The male
and female nuclei unite to each other, producing a zygote (oospore),
zygote is 2n, and this is the beginning of the sporophytic stage.
Development of sporogonium:
After fertilization, the zygote secretes a cellulose wall around it and

enlarges in size still it completely fills the venter of the archegonium. The
first division of the zygote is vertical. But according to Pande (1932) and
Bhardwaj (1950), in certain cases this divides transversely.
With the result of the first vertical division, the daughter cells are
produced, which are subjected to a transverse division producing four
cells of equal or unequal size. When the cells are unequal in size,
definitely the cells towards the neck of the archegonium are larger cells.
These cells again divide vertically, developing eightcelled embryo, four

cells in each tier. The upper tier of four cells divides transversely. This
way the three tiers of four cells each have been produced. The lowermost
tier produces the foot, the middle tier produces partly the foot and
mainly the seta and the upper-most tier produces the capsule.
The cells of the lowermost tier divide regularly or irregularly many times,
producing the bulbous foot. The foot is haustorial in nature, which
absorbs food from the tissue of the gametophyte.
The uppermost tier of four cells, divides once or twice transversely,

producing two or three tiers of cells. Now the cells divide periclinally,
giving rise to an outer layer the amphithecium and the central mass of
cells the endothecium. In the young sporogonium, the columella consists
of four vertical rows of the cells, the endothecium.
In the young sporogonium, the columella consists of four vertical rows of

the cells, but later on it is made up of sixteen rows of cells. According to
Bhardwaj (1958) in A. gemmulosus the columella consists of 36 to 49
vertical rows of the cells. The amphithecium divides periclinally
producing the outer sterile layer of the jacket initials and an inner
sporogenous tissue, the archesporium.
The jacket initials divide again and again periclinally producing the 4 to 6
layered wall of the capsule. The outermost layer develops into the single
layered epidermis. The epidermal cells are cutinized. The stomata may or
may not develop on the epidermal layer. The rest of the layers, beneath
the epidermis develop in normal chlorenchyma. These chlorophyllous
cells, however, help in synthesizing the food.
The layer of archesporium developed by periclinal division overarches

the columella. The sporogenous layer may be one to four celled in
thickness in its further development. In Anthoceros hawaiensis it
remains one celled in thickness; in A. pearsoni and A. himalayensis it
may become two, three or even four cells in thickness.
In younger stages all the cells of sporogenous tissue remain somewhat

rectangular in shape. Later on, the sporogenous tissue becomes
differentiated into two types of cells, i. e., (i) the sporocytes (spore
mother cells) and (ii) the sterile cells (pseudoelaters).
The sporocytes or spore mother cells undergo the reduction division,

each producing a tetrad of four spores. These spores are haploid. The
spores, which develop in the upper part of the capsule, mature first. Each
spore contains a nucleus and a chloroplast.
The sterile cells soon divide obliquely or transversely producing three to

five celled pseudoelaters. On maturity, the pseudoelaters lose their
protoplasm. The walls of the elaters may be smooth, irregularly
thickened or with spiral thickenings. Such thickenings, vary from species
to species. The pseudoelaters help in the dehiscence of the spores and
behave like true elaters. In earlier stages their function seems to be
nutritive.
The sporogonium and its dehiscence:
The capsules arise from the thalli in the form of small horny structures.
Usually they are two to three centimeters long. But in some species they
range even from five to fifteen centimeters in their height and because of
their horny appearance, the species are called ‘hornworts’. The mature
sporogonium consists of a bulbous foot and a projecting, slender and
erect capsule. There is a meristematic zone above the foot, instead of
seta.
The bulbous foot consists of parenchyma. It remains penetrated in the

thallus and acts as haustorium. The superficial cells of the foot are
palisade like. The space in between the foot and the capsule is occupied
by a meristematic zone. The cells of this zone divide throughout unless
and until the food is completely exhausted. This way, the capsule
increases in length. The capsule increases in height even after
maturation.
The cells developing from the meristem become differentiated into jacket
layer, columella and archesporium. Thus, the upper part of the
sporogonium matures first, and the basal part remains young. The
capsule does not mature at the same rate in all parts of it. The mature
spores liberate from the upper part of the capsule whereas, in the basal
part still the cells are in embryonic condition.
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The main capsule consists of many important parts. The central region of
the capsule is occupied by a sterile columella. According to Campbell
(1924), in A. fusiformis the columella acts as water conducting tissue.
However, this function of columella in other species is not very certain.
The main function of columella is to give the mechanical support to the
sporogonium. It also helps in the dispersal of the spores.
The columella remains surrounded by sporogenous tissue. In the region

just above the foot the archesporium is single layered and too young. The
tip region of the sporogonium possesses mature spores and elaters. The
wall of the capsule consists of the four to six layers of the
parenchymatous cells. The outermost layer is epidermis, which is
interrupted by stomata at several places. The epidermal cells are
cutinized.
The stomata open in the intercellular spaces of the chlorophyllous cells.

Usually each cell possesses two chloroplasts. The process of
photosynthesis takes place by means of chloroplasts and stomata. This
way, the organic food is synthesized for the sporogonium. However, the
sporogonium remains dependent upon the thallus for the supply of water
and other nutrients throughout its life.
On maturation, the tip of the sporogonium becomes black or dark-brown

in colour. The capsule divides at this stage. The tip of the capsule shrivels
up by losing water. The dehiscence of the capsule is more or less
dependent upon the loss of water. This way the dry atmosphere helps in
the dehiscence of the capsule.
The dehiscence begins from the tip region of the capsule. At first a small
longitudinal slit appears, which widens and enlarges. The pseudoelaters
are hygroscopic in nature. They begin to twist after the exposition of the
internal mass the outside. Due to this twisting of the pseudoelaters the
pressure is exerted on the jacket layer, and it bursts liberating the spores
in the atmosphere.
According to species, the dehiscence takes place by means of one to four

longitudinal slits. The valves of the capsule curve backward and
ultimately, on drying up they become twisted around each other. The
liberated spores are dispersed by wind from one place to another.
The spore:
In earlier stages the spores are found to be arranged in tetrads. After

being separated from each other they are dispersed. Each spore is
somewhat spherical and possesses two wall layers. The outer wall layer is
exine and the inner wall layer is intine.
The intine is smooth and thin, whereas the exine is somewhat thick and
ornamented. The colour of the mature spores varies from species to
species; this may be yellow, brown, dark brown or black. Each spore
possesses a single nucleus, a colourless plastid, few oil droplets and food
material within it.
Germination of spore:
After their liberation from the sporogonium the spores undergo a period
of rest prior to germination which ranges from few weeks to few months.
The exine of the spore ruptures, and the intine comes out in the form of a
germinal tube or protonema of variable length.
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Development of young gametophyte:
The chlorophyll present in the chloroplast of the spore passes in the

germinal tube along with oil droplets and food material. Thereafter the
germinal tube divides transversely at its apical end. This division is
followed by another transverse division. Soon after, the so formed two
cells divide by longitudinal intersecting walls and the quadrants are
resulted.
At the terminal end a growing point with an apical cell appears which by
producing several segments develops into the young gametophyte. Soon
after mucilage slit develops on the ventral side of the thallus. Some of the
marginal cells of the young thallus develop into smooth walled rhizoids.
The Nostoc sp. penetrates the thallus through the mucilage slits, which
later on form the colonies of the same.
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Inter-relationships of Anthocerotopsida:
The Anthocerotopsida differ from other bryophytes in a number of

respects. These differences are:
(a) Among Anthocerotopsida the cells are with large chloroplasts and
each chloroplast contains a pyrenoid.
(b) In this group the development of antheridia takes place from

hypodermal cells on the dorsal side of a gametophyte.
(c) The archegonia of this group are almost completely embedded in the
gametophyte.
(d) The growth of sporophyte is indeterminate because of a meristematic

region continually adding to the base of capsule.
Because of the differences mentioned above the class has been placed in
between Hepaticopsida on one hand and the Bryopsida on the other.
The other features of special interest of Anthocerotopsida are:
(a) The gametophytes are thallose, somewhat lobed or radially dissected,

and sometimes show a tendency of dichotomous branching.
(b) The gametophytes are always dorsiventrally differentiated and

possess numerous smooth-walled rhizoids on the ventral surface.
(c) The ventral surface is devoid of scales and mucilage hairs.
(d) Lateral margins of most of the genera of this group (except of

Dendroceros) are more than one cell in thickness.
(e) There is no internal differentiation of tissues.
(f) The ventral portion of a thallus has mucilage-filled intercellular

cavities on the dorsal side and opening to the surface by narrow slits.
These cavities generally contain Nostoc (a blue green alga) colonies in
them.
(g) The pyrenoids of Anthocerotopsida are not homologous with those of

green algae (chlorophyceae) since they consist of a crowded mass of 25 to
300 disc or spindle shaped bodies (McAllister, 1914, 1927).
(h) The growth of a thallus is initiated by a single apical cell with two
cutting faces (except in Dendroceros where there are three cutting faces).
(i) The vegetative reproduction by death and decay of thalli is less

frequent in this group than in Hepaticopsida. However, tuber formation
is frequent in this group.
(j) Most of the species are homothallic, but some are heterothallic
(Proskauer, 1948). In heterothallic species the sex determination is
genotypic, that is, two spores of a tetrad develop into male and two into
female gametophytes.
(k) Anthocerotopsida and Hepaticopsida are fundamentally different in

the fact that in the former an antheridial initial is the inner daughter cell
produced by periclinal division of a superficial daughter cell of the
gametophyte. This suggests that Anthocerotopsida are derived from
ancestors in which antheridia developed from superficial dorsal cells.
(l) In Anthoceros and some other genera of this group, the antheridial
initial may divide vertically into two or four daughter cells, each of which
develops into an antheridium.
(m) The development of the primary antheridial cell into the antheridium
proper is similar as in Sphaerocarpales and Marchantiales.
(n) In Anthocerotopsida, the spermatogenesis is much like that of other

bryophytes and involves a metamorphosis of androcytes into biflagellate
sperms on antherozoids.
(o) In Anthocerotopsida the archegonial initial functions directly as a

primary archegonial cell instead of dividing into primary archegonial cell
and primary stalk cell as found in other bryophytes.
(p) The first division of a zygote is vertical but cases have been found
(Bhardwaj, 1950; Pande, 1932) where the transverse division occurs.
(q) The amphithecium divides periclinally, where the outer layer

functions as the initial layer of the jacket and the inner layer as the
archesporium.
(r) One unique feature of Anthocerotopsida is that cells of a capsule do

not mature at the same rate and that cells in the basal portion of a
capsule remain embryonic even after those in the apical portion are fully
mature. This feature is not found in other bryophytes.
Affinities of Anthocerotopsida:
This group exhibits similarities with green algae, Hepaticopsida (liver

worts), Bryopsida (mosses) and Psilophytales of Pteridophyta.
(A) Features common with green algae:
The green algae (Chlorophyceae) have been considered a group from

which the bryophytes and pteridophytes are believed to have originated.
(i) Usually each cell of gametophyte has a single large chloroplast of a

definite shape.
(ii) The pyrenoids are present in the chloroplasts of the cells of

gametophyte. The presence of pyrenoids is the characteristic of cells of
green algae only
(iii) The pyrenoids of Anthocerotopsida and green algae are similar in

function and form starch grains in their peripheral region.
(iv) The outline and branching of the gametophyte is similar in both

cases.
(v) The presence of biciliate or biflagellate (both flagella of whiplash

type) antherozoids.
(B) Features common with liverworts (Hepaticopsida):
Many taxonomists include Anthocerotales in Hepaticae.
(i) The gametophyte is thallus-like.
(ii) The smooth-walled rhizoids are found in the members of

Anthocerotopsida as well as in the members of Jungermanniales of
Hepaticopsida.
(iii) The apical growth of thallus is similar in both.
(iv) Archesporium giving rise to spores and sterile cells in both groups.
The sterile cells with spiral bands are found in Megaceros of
Anthocerotopsida which exhibit the similarity with many members of
Hepaticopsida.
(v) Differentiation of the amphithecium and endothecium by periclinal

walls is similar to that of many Hepaticae.
(C) Features common with the Bryopsida (mosses):
(i) Presence of central columella in both groups.
(ii) Large reduction of the sporogenous tissue in both Anthocerotopsida

as well as Bryopsida groups.
(iii) Presence of functional stomata (e.g., in Funaria).
(iv) Differentiation of archesporium from the inner amphithecium as in

Sphagnales. This feature shows link between Anthocerotopsida and
Bryopsida.
(v) The developmental stages of embryo are quite similar. The early
divisions are very much alike.
(D) Features common with the Pteridophyta:
(i) The presence of sunken sex organs is common in both groups.
(ii) The presence of similar vegetative structure of the gametophyte in

Anthoceros and Fern.
(iii) The presence of highly developed sporogonium with photosynthetic

tissue indeterminate growth and functional stomata.
The abovementioned facts support that the Anthocerotopsida are a

distinct but synthetic group of plants. It forms a connecting link with
liverworts and mosses on one hand and with pteridophytes on the other.
There is also a remote connection with green algae (Chlorophyceae).
Campbell (1928) suggested, “The fact that the primary sporogenous
tissue in the Anthocerotales always arises from the amphithecium, while
in all other liverworts it is developed from the endothecium, would seem
to be a radical difference”.
Campbell is also of opinion that the sporophyte of Anthoceros with its

assimilatory system with stomata and continued growth shows the close
alliance to the independent, rootless dichotomously branched sporophyte
of the primitive fossil group, Psilophytales. Mehra (1957) suggested that
both Anthocerotopsida and Psilopsida arose from the common
Anthorhyniaceae stock.
Biological significance of Anthocerotopsida:
The sporophytes of Anthocerotopsida exhibit probable lines of biological

progress as follows:
(i) The presence of elaborate ventilated assimilatory system suggests the

beginning of the physiological independence of the sporophyte.
(ii) The discontinuation of continuous sporogenous tissue by the growth

of sterile cells between the spore mother cells which suggests the
beginning of the formation of sporangia.
(iii) The establishment of a well developed sterile columella from the

central endothecium suggests the beginning of the formation of
conducting system, and the initial stage in the formation of superficial
sporangia.
(iv) The presence of intercalary meristematic zone suggests the beginning

of the indeterminate and continuous growth of the sporophyte.
Summary:
Systematic position and distribution:
Division-Bryophyta; class Anthocerotopsida (Anthocerotae); Order-

Anthocerotales; Family- Anthocerotaceae; Genus-Anthoceros. 25 species
are found in India; cosmopolitan in distribution.
Habit: External features:
Moist shaded places. Thallus thin, dorsiventral with a broad indistinct

mid rib; tendency towards dichotomous branching; rhizoids-unicellular,
only smooth walled; unbranched scales are absent.
Internal structure:
All cells photosynthetic; apical growth is either by a single apical cell or a

group of apical cells.
Sexual organs:
Monoecious and dioecious; antheridia endogenous, inside closed

antheridial cavities on the dorsal surface of thallus, stalk is long, either
slender or massive, body jacket is single layered, antherozoid is biciliate;
archegonia-embedded in the dorsal surface of the thallus, neck is made
up of 6 vertical rows of cells, venter is single layered thick, cover cells are
2 to 4, neck canal cells are usually 4.
Sporophyte:
Amphithecium gives rise to jacket of capsule and archesporium;

endothecium gives rise to columella; archesproium gives rise to
sporocytes and pseudoelaters; foot here is bulbous; seta is absent;
capsule is long and cylindrical and its wall is four to six-layered and
persistent; pseudoelaters without thickenings; columella present, arched
over by spore sac; dehiscence of spores occur when capsule wall splits
into 1 to 4 valves, which remain united at the tip, pseudoelaters help in
spore dispersal.
Young gametophyte:
Two spore wall layers; size of spore is 0.025 to 0.05 mm; germinal tube
forms at the time of spore germination.
Home ›› Anthocerotopsida
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Useful Notes On Anthocerotopsida Order-Anthocerotales (4994 Words)

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Useful Notes on Anthocerotopsida

Here are your useful notes on Anthocerotopsida !

The class Anthocerotopsida (Anthocerotae) consists of a single order, the

However, according to top bryologists, there is only one family, i.e.,

Image Courtesy : upload.wikimedia.org/wikipedia/commons/c/ce/Phaeoceros_laevis.jpg

These genera are—Anthoceros, Phaeoceros, Aspiromitus, Notothylas,

However, the group is placed intermediate between Hepaticopsida

The most characteristic features of the group are as follows: The

The sporogonium arises from the dorsal side of the thallus. It is

There are five genera (i.e., Anthoceros, Phaeoceros, Aspiromitus,

The sporogonium (capsule) is linear and vertical. Generally the stomata

Habitat and distribution:

External structure of the thallus:

The thallus is small, prostrate, dark green and dorsiventrally

Internal structure of the thallus:

According to McAllister (1914, 1927) the pyrenoids of Anthoceros are

The apical growth of Anthoceros is a controversial topic, whether it takes

The vegetative reproduction takes place by various ways.

(a) By progressive growth and death of thallus:

The vegetative propagation takes place by progressive growth and death

In certain species of Anthoceros, the thallus becomes thickened at several

(d) By persistent growing apices:

According to Campbell the thalli of the species A. pearsoni and A.

The species of Anthoceros may be homothallic (monoecious) or

The antheridia are produced singly or in groups in the antheridial

It divides and redivides several times anticlinally and periclinally giving

According to Mehra and Handoo (1953), in A. erectus a number of

The antheridial initial divides twice by vertical walls intersecting each

The so formed androgonial cells divide repeatedly. The last generation of

Structure of mature antheridium and its dehiscence:

On the maturation of the antheridium, the roof of the antheridial

archegonium begins from a single superficial cell. This cell becomes

Prior to fertilization, the cover cells become detached from the

After fertilization, the zygote secretes a cellulose wall around it and

These cells again divide vertically, developing eightcelled embryo, four

The uppermost tier of four cells, divides once or twice transversely,

In the young sporogonium, the columella consists of four vertical rows of

The layer of archesporium developed by periclinal division overarches

In younger stages all the cells of sporogenous tissue remain somewhat

The sporocytes or spore mother cells undergo the reduction division,

The sterile cells soon divide obliquely or transversely producing three to

The sporogonium and its dehiscence:

The bulbous foot consists of parenchyma. It remains penetrated in the

The columella remains surrounded by sporogenous tissue. In the region

The stomata open in the intercellular spaces of the chlorophyllous cells.

On maturation, the tip of the sporogonium becomes black or dark-brown

According to species, the dehiscence takes place by means of one to four

In earlier stages the spores are found to be arranged in tetrads. After

Development of young gametophyte:

The chlorophyll present in the chloroplast of the spore passes in the

The Anthocerotopsida differ from other bryophytes in a number of

(b) In this group the development of antheridia takes place from

(d) The growth of sporophyte is indeterminate because of a meristematic

The other features of special interest of Anthocerotopsida are:

(a) The gametophytes are thallose, somewhat lobed or radially dissected,

(b) The gametophytes are always dorsiventrally differentiated and

(c) The ventral surface is devoid of scales and mucilage hairs.

(d) Lateral margins of most of the genera of this group (except of

(e) There is no internal differentiation of tissues.

(f) The ventral portion of a thallus has mucilage-filled intercellular