Funaria

Latin word funis =rope, denote characteristic twisting of the seta

SYSTEMATIC POSITION
Division : Bryophyta

Class : Bryopsida

Order : Funariales

Family : Funariaceae

Genus : Funaria

OCCURRENCE
➢ Funaria includes 117 species which are
cosmopolitan in distribution.
➢ It is terrestrial moss, which grows in dense
patches of bright green colour.
➢ It usually grows in moist shady banks and prefer
alkaline soil.
➢ Funaria hygrometrica, is the best-known species
of the genus.
➢ Shows heteromorphic alternation of generations
Gametophyte Sporophyte
• Haploid • Diploid
• Autotrophic & main plant body • Dependent & attached to gametophyte
• Thalloid and leafy form. • Foot, Seta, Capsule
• Produce archegonia and antheridia • Sporogenous tissue form spores by
(gametes) meiosis.
 GAMETOPHYTE
MORPHOLOGY
➢ Funaria main plant body is free living
gametophyte. Body consists of leafy
axis which is attached to substratum by
rhizoids.
➢ The axis is upright, branched and
covered by spirally arranged simple
sessile and ovate leaves.
➢ Rhizoids are branched, slender and
multicellular and characterized by the
presence of oblique septa (Fig. 1).

Fig 1. Funaria hygrometica, Plant body with sporophyte (A), leaf (B),
➢ The sporophytic plant body is photosynthetic, rhizoids showing oblique septa (C).

hence semi-parasitic on gametophore.

 ANATOMY
Stem
A transverse section of mature axis
shows three distinct regions (Fig 2A)
Epidermis: It is outermost single
layered thin-walled chlorophyllous
cells. It is devoid of cuticle and stomata.
Cortex: Present below the epidermis
and is made up of many layers of thin-
walled parenchymatous cells. The
younger cells contain chloroplasts.
Central cylinder: Consists of
vertically elongated thin walled narrow,
compactly arranged cells without
protoplasm. They help in the
conduction of water and nutrients. The
non-lignified central strands cells are
known as hydroids which considered
as tracheids in being dead at maturity. Fig 2. Funaria hygrometica, T.S of stem and leaf cut at a slightly higher level.

Leaf
➢ Leaf is composed of midrib and wings or lamina. Midrib is several celled thick.
It is composed of elongate, thick-walled cells, which help in conduction (Fig 1B,
2B).
➢ The leaf lamina consists of a single layer of parenchyma, which have
chloroplasts. The leaf does not have stomata or hairs.
 REPRODUCTION
Funaria gametophyte reproduces by vegetative and sexual reproduction.
Vegetative reproduction
➢ Fragmentation of Primary Protonema: Sometimes, primary protonema break
into many fragments and the fragments bearing buds develop into a new plant.
➢ Secondary Protonema: The protonema developing from any part of the plant
other than spores are called secondary protonema. Secondary protonema are
formed on injured rhizoids, stems, leaves or reproductive structures. These
injured structure bear buds which grow into a new plant.

Protonema

Bulbil

Gemmae

➢ Gemmae: During unfavourable condition gemmae are formed. Gemmae is

multicellular green bodies formed from the terminal cells of the protonema. In
favourable condition it germinates and from a new plant.
➢ Bulbil: Bulbil are similar to gemmae but develop on the rhizoidal branches
and devoid of chloroplast. Bulbil on detachment grow into a new plant.
➢ Apospory: Development of gametophyte form the sporophyte without forming
spores is known as apospory. Gametophytic protonema of Funaria may develop
from any unspecialised cells of the sporophyte.
 Sexual Reproduction
➢ Funaria is monoecious. Male and female sex organs
develop on different branches of the same plant.
➢ The sex organs i.e. antheridia and archegonia
develop at the apices of separate erect branches.
These branches are called gametophores.
➢ The female branch grows more vigorously and
becomes longer as compared to male branch.

Antheridial / Male Branch

➢ The leafy gametophore bearing antheridia is called male branch. Antheridia
develop in group at the convex shaped apex of a male branch.
➢ The antheridia are
intermingled with
paraphyses which are 4-6
cells high and one cell wide
sterile hairs. Paraphyses
perform following functions.
1. Photosynthesis (contain
chloroplast)
2. Protect the young antheridia
against desiccation
3. Help in the liberation of
antherozoids.
➢ Antheridia are also covered by
a number of bract-like leaves
called perichaetial leaves and
these form an envelope known Fig 3. Funaria. L. S of an antheridial branch (A), a single antheridium (B).

as perichaetium (Fig 3A).

Antheridium
➢ A mature antheridium is elongated and club-shaped raised on multicellular stalk.
➢ The main body of antheridium contains a mass of spermatogenesis cells. These
cells are surrounded by a layer of jacket cells.
➢ At the tip of jacket layer there is large hyaline cell, which forms operculum.
 Development of Antheridium
➢ The antheridium develops from a superficial antheridial initial which divides
transversely and produced an outer cell and a basal cell. The outer cell divides
further by successive transverse divisions to form a linear filament of 2 to 4 cells.
Basal cells form the stalk of the antheridium (Fig. 4 A-C).
➢ The terminal apical cell of the filament divides by two vertical intersecting walls
so the apical cell with two cutting faces formed (Fig. 4D).
➢ Apical cell cuts off cells in two rows. Each young segment of the upper 3 to 4
cells now divides by a vertically diagonal wall to form two unequal cells (Fig.
4E-F).

Fig 4. Funaria hygrometrica, development of antheridium (A-K)

➢ The smaller peripheral cells act as a first jacket initial. Whereas, the larger
daughter cell, through the similar division, forms the outer second jacket
initials and the inner primary androgonial cell. Jacket initial divide
anticlinally and forms antheridial jacket (Fig. 4G-H).
➢ Apical cell of the filament forms operculum or cap region of the antheridial
wall (Fig. 4I).
➢ The primary androgonial cell divides and re-divides to form androcyte
mother cells. Each androcyte mother cell divides to form two androcytes.
The androcytes transform into biflagellate antherozoids or sperms (Fig. 4
J-K).
➢ Antheridia in an antheridial head mature at different times.
 Archegonial / Female Branch
➢ Archegonial branch arises from the base of male branch.
➢ Archegonia develops in clusters and surrounded by perichaetial leaves.
➢ They are intermingled with paraphyses (Fig 5).
Archegonium
➢ A mature archegonium is flask-
shaped (Fig 5). It is borne on
short stalk consisting two parts a
basal swollen part called
venter, and elongated neck.
➢ The venter is surrounded by a
double layered sterile jacket,
whereas the jacket around the
neck is single layered.
➢ The venter contain venter canal
cell and an egg

Fig 5. Funaria LS of an archegonial branch bearing archegonia.

 Development of archegonia
➢ Archegonia develop from the archegonial initial, present at the apex of archegonial
branch.
➢ It divides transversely to form an upper cell and a lower cell.
➢ The upper cell divides by two intersecting oblique walls forming an apical cell with
two cutting faces (Fig. 6 A-C). archegonia mother cell
➢ The apical cell cuts off 4- 8 segments. They develop into stalk. Afterwards the apical
cell divides in such a way that it gives rise to central axial cell and peripheral cells.
(Fig. 6 D-E).

Fig 6. Funaria showing successive stages of the development of archegonium (A-I).

➢ The peripheral cells divide to form jacket initials. These initials divide to form jacket
layers around the venter and neck.
➢ The axial cell divides to form primary cover cell and central cell. (Fig. 6 F).
➢ Primary cover cell later forms the cover cell and the central cell divides to
form primary neck canal cell and primary venter cell
➢ The primary neck canal cell gives rise to neck canal cells (Fig.6 G) and the primary
venter cell divides to produce venter canal cell and an egg (Fig.6 H-I).
 Fertilization
➢ The dehiscence of the mature antheridium only takes place in presence of water. The
opercular cell absorbs water and the inner wall rupture and eventually a pore is formed
at the distal end of the antheridium.
➢ The androcytes release through the pore in the form of a viscous fluid due to the
hygroscopic pressure developed within the antheridial cavity.
➢ In archegonium the ventral canal cell and the neck canal cells dissolve forming a
mucilaginous substance. This mucilaginous substance absorbs water accumulated as
rain or dew water, then swells up and the resultant pressure breaks apart the terminal
cover cell and clear passage is created.
➢ Now antherozoids are chemotactically move towards the archegonia. A large number
of antherozoids enter the neck, but only one of them fertilizes with the egg nucleus and
produces diploid zygote.
➢ The process of fertilization stimulates cells of venter wall and they divide to form
calyptra. Calyptra later form the protective layer of capsule.
 SPOROPHYTE
➢ The fertilised egg or zygote is the first cell of the sporophytic generation (Fig.
7A). The zygote swells up, increases in size and forms a wall before further
divisions.
Development of Sporogonium
➢ The zygote enlarges and divides
transversely into an
upper epibasal and a
lower hypobasal cell (Fig. 7B).
➢ Both epibasal and hypobasal
cells obliquely divide to form
two apical cells (Fig. 7 C).
➢ The segments cut off from
epibasal apical cells
form capsule and upper part of
the seta.
Fig 7. Funaria, mature zygote inside the archegonium (A), early stages of the
➢ The segments cut off from development of sporophyte (B-E).
hypobasal apical cell develop
into lower part of seta and foot
(Fig. 7D-E).
➢ In a transverse section, just below the apical cell,
the two derivatives, one each from the two faces
of the epibasal apical cell, form an almost
spherical segment (Fig.A). Both these derivatives
divide by a vertical wall forming a quadrant
embryo (Fig.B).
➢ Each cell of the quadrant divides and form 8-
celled embryo; the four triangular and the four
rectangular cells are arranged alternately
(Fig.C).
➢ Each rectangular cell now divides periclinally
and at this stage eight peripheral cells, which
form the amphithecium, encircle a central core
of four cells, the endothecium.
➢ The amphithecium and the endothecium differentiate further and contribute to the three
regions of capsule; operculum, theca and apophysis.
➢ The cells of amphithecium divide periclinally. They form outer and inner layers of
eight cells each. The inner layer is called first ring. Its cells divide and mature into
outer wall of spore sac (Fig.E).
➢ The cells of outer layer divide anticlinally first and then periclinally. They produce an
inner 16-celled second ring. These cells give rise to trabeculae. (Fig.8F).
➢ The cells outer to second ring divides again anticlinally and periclinally (Fig.8G). They
produce a 32-celled inner third ring (Fig.8H). The cells of this layer differentiate
into spongy layer.
➢ The cells external to third ring divide periclinally to separate fourth and fifth rings
(Fig. I). Each ring is 32-celled. The cells of these rings differentiate
into hypodermis and epidermis.
➢ The endothecium cell divides first by a curved vertical wall. It separates a triangular
and a rectangular cell. Then periclinal wall produce four central cells and a ring of eight
peripheral cells.
➢ The central cells divide and differentiate into central columella. The peripheral cells
divide periclinally into an outer layer and an inner layer. The cells of outer layer divide
repeatedly to form sporogenous tissue (archesporium). The cells of inner layer give
rise to inner wall of the spore sac. In Funaria the outer wall of spore sac develops
from the endothecium.
➢ Sporogenous tissue later differentiate into spore mother cells. Each spore mother cell
now divided by meiosis and forms four haploid spores.

Structure of Sporogonium
Morphology
➢ The sporophyte stage of bryophytes is
also called sporogonium.
➢ A mature sporogonium is differentiated
into a massive foot, a long seta, and a
pear-shaped capsule (Fig9).
➢ The foot is embedded in the apical
tissue of female branch. It absorbs
water and nutrients from the
gametophyte.
➢ The seta is a long stalk. It carries
capsule at its apex.
➢ The capsule is pear-shaped highly
organized spore-producing structure.
The young capsule is green but later it
becomes dark. The apical part of the
capsule is covered with remains of
Fig 9. Funaria, a gametophyte with sporophyte (A), L.S of capsule (B).
 ruptured calyptra. The capsule has
considerable differentiation of tissues.

Anatomy
➢ The foot is bulbous mass of tissue and
it gives rise to seta.
➢ The seta consists of central conducting
strand and are composed of thin-walled
cells. These cells are surrounded by
cortex and epidermis. The epidermis is
covered with cuticle.

➢ The capsule is differentiated into three regions, the apophysis, theca and apical
region.
➢ Apophysis
➢ Apophysis is slightly swollen basal
sterile region of the capsule.
➢ Beneath the epidermis is
photosynthetic spongy layer. Thus
partially depend upon the
gametophyte.
➢ Strand of thin-walled, vertically
elongated cells is present in the
centre.
Theca
➢ It is central part of the capsule. It has
sterile central column of tissue
called columella.
➢ The columella is surrounded by
barrel-shaped spore sac. The spore
sac contains spore mother cells.
➢ A wide air space is present outer to
spore sac. This air space contains
transverse strands called
trabeculae. These trabeculae
connect the capsule wall with the
wall of the spore sac.
 ➢ The wall of the theca consists of epidermis, hypodermis and two cells of thick
photosynthetic spongy layer.
Apical Region:
➢ Consist of two important part, the
operculum and peristome.
➢ The operculum is conical cap on
terminal region of the capsule. It consists
of 3-4 layers of thin-walled cells.
Immediately below the operculum is
peristome.
➢ The peristome are simply strips of
cuticle and they are not cellular.
Peristome consists of two rings. In each
ring there are outer 16 long, triangular
teeth and inner 16 thin-walled teeth (Fig.
11). The peristome teeth are
hygroscopic. They respond to slight
changes in humidity.
➢ The peristome is attached to a ring of thin-walled cells which form rim of the capsule.

Dehiscence of capsule
➢ When sporophytes reach at maturity, the water supply to the capsule is cut off. As a
result, all tissues of the capsule shrinks, expect spores. It loosens the connection
between operculum and underlying tissue. Finally, the operculum is shed. It exposes
the peristome and teeth of the peristome regulates dispersal of spores.
 Germination of Spores and Development of Protonema
➢ The spore shed from the capsule
and it start germination. The
spore increases in size and the
outer spore wall, exospore
ruptures.
➢ The inner spore wall,
endospore grows out. It forms
one or two germ tubes (Fig 12 A-
B).
➢ A cross wall is formed near its
point of emergence. It separates
the germ tube from the spore.
➢ The cell cut off soon develops
Fig 12. Funaria, spore (A), germination of spore (B), successive stages of the
into a branched multicellular formation of primary protonema, secondary protonema and young
gametophore (C-E).
filament, the protonema (Fig. 12 C).
➢ A protonema is differentiated into two kinds of branches:
o Chloronema: It grows along the surface of the substratum or into the air.
o Rhizoids: They penetrate into the substratum.
➢ The chloronemal stage grows extensively. After 20 days, most of the cells of
chloronema degenerate. Only a few apical cells are left. These cells give rise to
another type of filaments, the caulonema.
➢ Buds develop on the caulonema filament and this bud gives rise to gametophores
(Fig. 12 D-E).
 LIFE CYCLE
The life cycle of Funaria hygrometrica is presented in below figure.
Funaria