RICCIA

Hepaticopsida
• Divided into four orders (Cavers, 1910, Campbell, 1936).

i. Sphaerocarpales
ii. Marchantiales
iii. Jaungermanniales
iv. Calobryales.
Classification

Division: Bryophyta
Class : Hepaticopsida
Order : Marchantiales
Family : Ricciaceae
Genus : Riccia
Distribution and Habitat of Riccia
• Riccia is generally a normal Liverwort.
• Riccia, the most widely distributed genus of family Ricciaceae.
• Species of Riccia are found in the tropical and subtropical regions of the
world.
• During the rainy season it is seen or grown on the land or the layer of a
wet substrate.
• Among all of the Bryophtes, the species of Riccia is seen all over the
world and it is well distributed.
• The Total species of Riccia is 130, out of these all are terrestrial except
one and they are grown all over the world.
• Riccia fluitans - floating Riccia , aquatic.
• They are terrestrial thallus and the gametophyte is the main plant of life
cycle.
External Structure of Plant body-
the Gametophyte
• Main plant body is gametophytic, thalloid (thallus like).
• Riccia generally prostrate, dorsiventral, dichotomously branched thallus.
• The dichotomous branching of the thallus is quite close to each other.
• It is deep green. However, the aquatic species possess light green
thallus.
• The thallus of a Riccia is dichotomously branched and collectively make
a structure like as a rose. This is called the rosette shape.
Notches:
• The upper part of a thallus is green in color and it
has a thick midrib.
• Each branch of thallus has a conspicuous median
longitudinal groove.
• The groove is present on the dorsal side of each
branch of the thallus.
• The top point of a thallus has a thin part which is
called apical notch.
• In this part the development of thallus is occur.
Scales:
• The ventral surface of the thallus bears a row of scales along the margin of the
thallus. The scales are violet coloured and multicellular.
• They are arranged close to each other towards the apex of the branch.
• On the other hand, the scales are quite apart from each other away from the
notch.
• The scales are found in one row towards the apex of the branch.
• But they are found in two rows in the portion away from the apex.
• Thus the older parts of the thallus bear two rows of the scales.
• In Riccia crystalline the scales are either absent or rudimentary.
Rhizoid:
• The rhizoids are also found on the ventral surface of the thallus in the middle.
• They are unicellular.
• Rhizoids absorb the nutrients and water from substratum.
• The rhizoids are two types. Some are smooth walled and some are with
tuberculate walls.
• The tuberculate rhizoids possess the peg-like infolding peeping.
• The simple or smooth walled rhizoids have no such infoldings.
Internal Structure(Anatomy)
• The transverse section (T.S.) of a thallus shows two distinct regions of cells
internally under microscope.
1. Dorsal region: a layer with chloroplast which is a photosynthetic region.
2. Ventral region: a colourless parenchyma layer which is storage region.
Dorsal Surface/ photosynthetic
region/ Assimilatory Zone:
• The dorsal surface has a cell layer with
chloroplast which is organized in vertical rows.
• Each cell layers has a midrib space which is
filled with air is called air chamber.
• This region is photosynthetic. It prepares
carbohydrates.
• The upper cell layer of dorsal surface is called
upper epidermis.
• The epidermis of the thallus is discontinuous
and contains air pores. The airpores function
for gas exchange.
• The epidermis is single layered .
• However the epidermis is continuous in aquatic
species.
Ventral surface/ Lower Storage
Region:

• The lower part of a thallus i.e. ventral

region consists of simple colourless
parenchyma.
• No intercellular spaces are found in this
region. Cells are compactly arranged.
• The cells of this region make the storage
tissue. They are filled with starch grains.
• The ventral region has a single layered
epidermis.
• Epidermis gives rises to several unicellular Ref: Vashishta Page
rhizoids and multicellular scales.
46
Vegetative Reproduction
1) Fragmentation
2) Formation of adventitious branches
3) Persistent growing apices
4) Formation of tubers
Sexual Reproduction
• oogamous i.e. union between a motile flagellate male gamate and a resting

non-flagellate female gamete takes place .

• male sex organs are antheridia (borne in antheridial chambers)and female sex
organs are archegonia (borne in archegonial chambers)
• some of the species are monoecious i.e. both the types of sex organs may
develop on the same thallus (R. gangetica, R. glauca) ; and some are
dioecious i.e. the sex organs may develop on different thalli (R. discolor, R.
personii)
• sex organs arise singly in acropetal succession i.e. the first formed (old) sex
organ is behind and the last formed (new) sex organ is near the growing apex.
Structure of mature antheridium
• The mature antheridium remains embedded in the antheridial chamber. This
chamber opens by an ostiole on the dorsal side of the thallus.
• The mature antheridium consists of a few celled stalk and antheridium proper.
• The antheridium proper may be rounded or pointed at its apical end.
• A sterile single layered jacket-layer encircles the antheridium and protects it.
• The mature antheridium contains androcytes within the jacket layer.
• Each androcyte metamorphoses into an antherozoid.
Development of Antheridium
1. The development of antheridium starts from a
superficial antheridial cell, situated on the
dorsal surface of the thallus (Fig. 15 A). It is
a few (2-3) cells away from the apical cell.
2. The antheridial initial enlarges in size,
becomes papillate and divides first by a
transverse division to form an upper outer
cell and a lower basal cell (Fig. 15 B). Basal
cell remains embedded in the tissue of
thallus, undergoes only a little further
development and forms the embedded
portion of the antheridial stalk.
Development of Antheridium
3. Outer cell divides by transverse divisions to
form a filament of 4 cells. Upper two cells of
the 4 celled filament are known as primary
antheridial cells and lower two cells are
known as primary stalk cells (Fig. 15 C-E).
4. Primary stalk cells form the stalk of the
antheridium. Primary antheridial cells
divide by two successive vertical divisions at
right angle to each other to form two tiers of
four cells each (Fig. 15 F, G).

Ref: Vashishta, Page 50-52

Development of Antheridium
5. A periclinal division is laid down in both the
tiers of four cells and there is the formation of
eight outer sterile jacket initials and 8 inner
primary androgonial cells (Fig. 15 H, I).
6. Jacket initials divide by several anticlinal
divisions to form a single layer of atheridial
jacket. Primary androgonial cells divide by
several repeated transverse and vertical
divisions resulting in the formation of large
number of small cubical androgonial cells
(Fig. 15 K-M). The last generation of the
androgonial cells is known as androcyte
mother cells.
Spermatogenesis
• The process of metamorphosis of androcyte mother cells into antherozoids is
called spermatogenesis.
• It completes within a few minutes.

1. Each androcyte mother cell divides by a diagonal

mitotic division to form two triangular cells called
androcytes. Both the androcytes remain enclosed in
the wall of the androcyte mother cell with one
separate wall (Fig. 16 A, B).
2. Each androcyte has a prominent nucleus and a small
extra-nuclear granule called blepharoplast. It lies
near the periphery of the protoplast (Fig. 16 C).
The androcyte soon looses its triangular shape and
becomes somewhat round or oval (Fig. 16 D). Its
blepharoplast elongates into a cord and occupies
about two-thirds of its part (Fig. 16 E).
3. Simultaneously the nucleus also becomes crescent
shaped, homogeneous and ultimately comes in
contact with the blepharoplast (Fig. 16 F). The
cell become coiled.
4. Two large flagella develop from the conspicuously
thickened end of the blepharoplast (Fig. 16 G).
5. A small part of the cytoplasm, which is not utilised
in the formation of flagella may remain attached to
the posterior end of antherozoid as a small vesicle.
Each androcyte thus metamorphosis into an
antherozoid (Fig.16 E-H).
Structure of a mature archegonium
• The archegonium is a flask-shaped body. It is
attached to the thallus by a short stalk. It
consists of an elongated neck and a
bulbous venter.
a) Neck:
• The neck is 6 to 9 cells in height.
• It consists of six vertical rows of the cells.
• The six vertical rows of the cells enclose a neck
canal.
• There are four cover cells at the top of neck
canal.
• Four neck canal cells are found within a neck
canal before maturity.
• They disintegrate into mucilaginous mass on the
maturation of the archegonium.
Structure of a mature archegonium
b) Venter:
• The lower bulbous structure of the archegonium is
venter.
• This has a single layered wall around it.
• This wall is of 12-20 cells in perimeter.
• The venter encloses a venter canal cell and the
large egg.
• The venter canal cell disintegrates on the maturity of
the archegonium .
• Thus only the large egg remains.
❑The mucilaginous mass absorbs water by imbibition
on the maturity. It creates pressure. Thus the cover
cells become separated from each other
and an opening is formed. The mucilaginous mass
extrudes through this opening. It attracts the
antherozoids.
Development of Archegonium
1. Archegonium develops from a superficial cell.
This cell lies two or three cells away from the
apex of the thallus. This cell is called the
archegonial initial. (Fig: A)
2. The archegonial initial divides by a
transverse devision into two cells. The lower
cell is the basal cell and the upper cell is the
outer cell. The lowermost portion of the
archegonium is produced from the basal cell.
The outer cell produces the main body of the
archegonium. (Fig: B)
3. The outer cell divides vertically three times.
Thus it gives rise to three peripheral initials. It
itself becomes primary axial cell. The three
peripheral initials are situated around a
primary axial cell. (Fig: C,D,E)
• Ref: Vashishta, Page 53-54
Development of Archegonium
4. The three peripheral initials divide vertically.
They form six jacket initials. All the six cells
surround the central primary axial cell.
Soon after each jacket initial divides by a
transverse division producing six neck
initials and six venter initials. They are
arranged in two tiers. The tier of six neck cells
is situated above the tier of six venter initials.
(Fig: G,H)
5. Several transverse divisions occur in the
neck initials. It gives rise to a neck of
archegonium 6 to 9 cells in length and six-
celled in perimeter. The venter initials divides
to form jacket of the venter. The jacket of the
venter is :2 to 20 cells in perimeter. (Fig: I,J,K)
Development of Archegonium
6. The primary axial cell divides by an unequal
transverse division. It produces a small primary
cover cell and a central cell. (Fig: F)
7. The primary cover cell divides twice vertically. It
produces four cover cells at the apex of jacket
layer (Fig: I). The central cell divides by a
transverse wall. It produces a primary neck canal
cell and primary venter cell (Fig:G)
8. The primary neck canal cell divides twice. It gives
rise to four neck canal cells. They are situated
within the neck of the archegonium (Fig:K). The
primary venter cell divides by an unequal
transverse division. It gives rise to a venter canal
cell and a large egg. The venter canal cell is
situated above the large egg. The neck canal cells,
venter canal cell and the cover cells are separated
from each other in mature archegonium (Fig:L)
Fertilization or syngamy
• The water is necessary for fertilization.
• The antherozoids reach to the mouth of the neck of
archegonium through water.
• The water is also essential for the separation of the
cover cells.
• Mucilaginous substances along with proteins and
certain inorganic salts ooze out through the mouth of the
archegonium.
• The antherozoids are attracted chemotactically by this
substance.
• The antherozoids enter the mouth of the archegonium.
• They travel through the neck and reach near the egg.
• One of the antherozoids penetrates egg cell and the
fertilization takes place.
Development of embryo
1. Formation of calyptra: The zygote
secretes a wall around it after the
fertilization. It enlarges in size. It nearly
fills the cavity of the venter. The venter
cells divide periclinally and anticlinally
thus the wall of the venter becomes
two-celled in thickness and form two
layered calyptra. The developing
embryo is situated inside the
calyptra (Fig: C,D)

Ref: Vashishta, Page 55

Development of embryo
2. Formation of octant: The first division
of the zygote is transverse. Thus two
equal cells are formed. In most of the
species these two cells are
called epibasal and hypobasal (Fig: C)
They divide vertically and a four celled
embryo of quadrant type is formed (Fig: D).
(In some species, four celled filamentous
type embryo is formed from the two celled
embryo . Sometimes both the types of
embryo are formed in the same genus). This
four celled embryo divides by vertical division.
Thus the eight celled embryo (octant stage) is
formed. It is called octant stage.
Development of embryo
3. Several irregular divisions occur in
octant stage. Thus 20 to 30 celled
embryo is formed (Fig: F). At this stage
periclinal divisions occur. Now the
embryo is differentiated into two
regions. The outer layer is called
amphithecium. The inner mass of the
cells is called endothecium (Fig:F).
4. The amphithecium is protective in
nature and forms the jacket of the
capsule. The endothecium divides
repeatedly. It gives rise to mass of
archesporium or sporogenous cells.
Development of embryo
5. The sporogenous tissue differentiates
into sporocytes than in spore mother
cells and nurse cells. The sporocytes
consist of dense granular cytoplasm.
The nurse cells have this vacuolated
cytoplasm.
6. The nurse cells and the inner jacket
cells disintegrate. They give rise to a
mucilaginous mass. The sporocyte
remain suspended in the mucilaginous
mass (Fig: H). The sporocytes receive
the nourishment from this mucilaginous
fluid during meiosis and form spore
tetrad. Finally, the walls of the spore
mother cells disintegrate. Now they lie
free in the sporogonium.
Sporophyte Characteristics
• Foot and seta are absent.
• The capsule is globose.
• Capsule wall is one layered, disorganized and abortive.
• Amphithecium forms the jacket of the capsule.
• Endothecium forms the archesporium (mass of sporogenous tissue).
• Archesporium give rise to the nurse cells and spore mother cells.
• No special mechanism of dehiscence. Spore are set free after the death
and decay of the thallus.
Germination of spore and formation of
gametophyte
• The spore is the beginning of the gametophytic generation. It has following
process of germination.
1. The two outer layers of spore, exosporium and mesosporium rupture. The
endosporium protrudes out through the opening. It forms a germinal tube. Much of the
cytoplasm is shifted in the terminal end. Now transverse walls are laid down at the apex
of this tube. A rhizoid comes out near the germinal tube, of the spore.
2. The distal end of the germinal tube again divides by a parallel wall. Thus two cells are
formed at the distal end of the germinal tube. Now both the cells divide vertically
twice. They give rise to two tiers of four cells each.
Germination of spore and formation of
gametophyte
3. The four cells divides transversely. They give rise to the cylindrical
elongated posterior portion of a young gametophyte.
4. One cell of the four celled distal tier acts as an apical cell. It possesses two
cutting faces. Thus it gives rise to a multicellular young thallus.
Alternation of Generation
• Riccia shows alternation of generations a haploid gametophyte generation and
a diploid sporophyte generation alternate each other.
• The life cycle of Riccia shows regular alternation of gametophytic and
sporophyte generations. The two generations are morphologically different
hence this type of alternation of generation is known as heteromorphic.
• The haploid generation is called the gametophyte because it undergoes
sexual reproduction to produce male and female gametes. Production of
gametes involves mitosis, so the gametes are also haploid. The male and
female gametes of Riccia fuse to form a diploid zygote which grows into the
next generation the diploid sporophyte generation. Sporophyte produces
spore tetrads. Production of spore involves meiosis. Hence, there is a return to
the haploid condition. The haploid spores give rise to the gametophyte
generation. In Riccia, the gametophyte generation is dominant.
Riccia Life cycle