Life Cycle of Funaria

Gametophytic Phase of Funaria:

(i) External Features:
Plant body is gametophytic and consists of two different
stages namely:
(i) Juvenile stage represented by primary protonema and

(ii) The leafy gametophore which represents the adult form.

The adult gametophyte (gametophore) is differentiated into rhizoids,

axis or ‘stem’ and ‘leaves’ (Fig. 1 A). Rhizoids arise from the base of
the axis. They are slender, branched, and multicellular and have
oblique septa.
Axis is 1—3 cm. high, upright, slender and branched. Each branch is
extra axillary i.e., arise below a leaf. Leaves are sessile, oblong-ovate
with entire margin, pointed apex and are arranged spirally on the
branches and ‘stem’. Each ‘leaf’ is traversed by a single mid rib (Fig.
1B). ‘Leaves’ are borne in 1/3 phyllotaxy which becomes 3/8 at
maturity.

(ii) Internal Structure:

1. Axis or ‘stem’
The transverse section (T. S.) of axis can be differentiated
into three distinct regions:
(i) Epidermis

(ii) Cortex

(iii) Central conducting strand or central cylinder.

(i) Epidermis:
It is the outer most single layered protective covering consisting of
small tangentially elongated chlorophyll bearing cells. Cuticle and
stomata are absent (Fig. 2).

(ii) Cortex:
It is present between the epidermis and conducting tissue. It is made
up to parenchymatous cells. Younger part of the cortex contains
chloroplasts but in the older part they are lacking. At maturity few
outer layers of cortex become thick walled and are reddish brown in
colour but those of the inner layers become thin walled.

(iii) Central Conducting Strand:

It is made up of long, narrow thin walled dead cells which lack
protoplasm. These cells are now commonly called as hydroids.
Conducting strand besides providing a certain amount of mechanical
support, functions in the upward conduction of water and solutes.
2. Leaf:
Transverse section (T. S.) of ‘leaf’ shows a well-defined midrib with
two lateral wings. Except the midrib region, the ‘leaf’ is composed of
single layer of parenchymatous polygonal cells. The cells contain many
large and prominent chloroplasts (Fig. 3). The central part of the mid
rib has narrow conducting strand of thick walled cells which help in
conduction.

Reproduction in Funaria:
Funaria reproduces by vegetative and sexual methods.

(i) Vegetative Reproduction:

It takes place by the following methods:
1. By multiplication of primary protonema:
In Funaria, spores on germination form a branched, filamentous,
multicellular structure. It’s called primary protonema. In it certain
colourless separation cells are formed by intercalary divisions. These
cells die out and break up the protonema into single cell or many
celled fragments. These fragments grow into new protonemata which
bear buds. Each bud develops into a leafy gametophore.

2. By secondary protonema:
When protonema is developed by other than the germination of spore,
it is called secondary protonema. It may be developed from any
detached living part of the gametophyte such as ‘stem’, ‘leaves’,
antheridium, archegonium paraphysis, sterile cells of capsule, seta or
when the rhizoids are exposed to sun light in moist atmosphere (Fig.
16 G). It is similar to primary protonema and develops into leafy
gametophore.

3. By Gemmae:
During unfavorable conditions, the terminal cells of the protonemal
branches divide by transverse, longitudinal divisions and form green
multicellular bodies of 10-30 cells (Fig. 4). These are called gemmae.
At maturity gemmae become slightly reddish brown in colour. On the
return of favourable conditions gemmae germinate and form new
plants.

4. By Bulbils:
When such gemmae like structures are produced on rhizoids inside
the substratum, these are called bulbils. These are devoid of
chloroplasts but capable of developing into leafy individuals under
favourable conditions.

5. Apospory:
Development of gametophyte from sporophyte without the formation
of spores is known as apospory. Any vegetative cell of the sporophyte
may form green protonemal filaments which bear lateral buds. These
buds later develop into leafy gametophores.

The gametophores thus formed are diploid. Sexual reproduction in

such gametophores results in the formation of tetraploid (4n) zygote.
The sporophytes from tetraploid are sterile because they are not
capable of bearing spores.

(ii) Sexual Reproduction in Funaria:

Sexual reproduction is oogamous. Male reproductive structure is
known as antheridium and female as archegonium. Funaria is
monoecious (having male and female sex organs on the same thallus)
and autoicous (antheridia and archegonia develop on separate
branches of the same thallus). Sex organs are borne on leafy
gametophores in terminal clusters.

The main shoot of the leafy gametophore bears antheridia and act as
male branch. Female branch develops as a lateral outgrowth from the
base of the male branch and bears archegonia. It grows higher than
the male branch. Funaria is protandrous (antheridia mature before the
archegonia). It ensures the cross fertilization.

Male Branch or Antheridiophore:

Longitudinal section (L. S.) of male branch shows that its apex is
expanded and convex shaped. It bears large number of reddish brown
or orange antheridia in different stages of development. Projected
antheridia are surrounded by a rosette of spreading leaves called
perigonial leaves.

The antheridial cluster with surrounding perigonial leaves is called

perigonium. The antheridia are intermingled with large number of
sterile hair like club shaped structures called paraphyses (Sing,
paraphysis) (Fig. 5). Paraphyses store water, protect developing
antheridia, help in photosynthesis and dehiscence of antheridia.
Structure of an Antheridium:
The antheridium is club shaped. It can be differentiated into
two parts:
(a) Short multicellular stalk

(b) Body of antheridium (Fig. 6K).

Body of antheridium has sterile, single layered jacket of polyhedral

flattened cells. When young the cells of the jacket contain chloroplasts
which turn orange or reddish brown at maturity. Jacket encloses a
large number of androcytes (antherozoid mother cells).

At maturity the distal end of the antheridium bears one or two thick
walled, colourless cells called operculum. The opercular cells become
mucilaginous, absorb water and swell, break connections with the
neighbouring cells and form a narrow pore. Androcytes ooze out in the
form of a viscous fluid through this pore.

Development of Antheridium:
Antheridium develops from a single superficial slightly projected cell.
It is called antheridial initial. It develops at the apex of the male
branch. It divides by a transverse division to form a basal cell and an
outer cell (Fig. 6 A, B). The basal cell form the embedded portion of
the stalk. The outer cell forms the entire antheridium and is known as
antheridial mother cell. It divides by transverse divisions to form a
short filament of 2-3 cells (Fig. 6 C).

The lower cells form the lower part of the stalk of the antheridium.
The terminal cell of the filament divides by two vertical intersecting
walls, thus an apical cell with two cutting faces is differentiated (Fig.
6D).

The apical cell cuts segments in two rows in regular alternate

sequence. In this way 5-7 segments cut off (Fig. 6E). Simultaneously
when the apical cell is dividing, the third or fourth segments below the
apical cell, starts dividing from base, upwards by diagonal vertical
walls.

The first wall divides the segment into two cells of unequal size. Small
cell is called jacket initial (Fig. 6 F). Larger cell further divides
periclinally into an inner large primary androgonial cell (Fig. 6 G) and
outer jacket initial. In a transverse section (T. S.) primary androgonial
cell appears as a triangular cell (Fig. 6G). Such type of divisions takes
place in all the upper segments except the apical cell which develops
into operculum.

All the jacket initials divide only by anticlinal divisions to form a single
layered wall of antheridium (Fig. 6 H-J). Primary androgonial cells
divide and re-divide to form the androcyte mother cells. The cells of
the last cell generation are called androcyte mother cells.
Each androcyte mother cell divides further and form two androcytes.
Each androcyte produces a single biflagellate sperm or antherozoid or
spermatozoid. Each antherozoid is elongated, spirally coiled, bi-
flagellated structure (Fig. 6L, M).

Female Branch or Archegoniphore:

The female branch arises from the base of the male branch.
Longitudinal section (L. S.) of female branch shows that many
archegonia intermingled with paraphyses occurs at its apex (Fig. 7A).
The terminal cell of paraphyses is not swollen. The cluster of
archegonia is enclosed by a group of green foliage ‘leaves’ called
perichaetial leaves. The archegonial cluster with the surrounding
perichaetial leaves is called perichaetium.

Structure of an Archegonium:
A mature archegonium is flask shaped structure. It remains attached
to the female branch by a massive stalk. It consists upper elongated
slender neck and basal globular portion called venter (Fig. 7B).

The neck is slightly tubular, twisted, single layered and consists of six
vertical rows of neck cells, which enclose an axial row of ten or more
neck canal cells. The venter wall is two layered and encloses venter
canal cell and egg cell. Venter canal cell is situated just below the neck
canal cells.

Development of Archegonium:
Archegonium develops from a single superficial cell called the
archegonial initial (Fig. 8 A). It differentiates at the apex of the female
branch. Archegonial initial divides by transverse division to form the
basal cell or stalk cell and a terminal cell.

The basal cell divides and re-divides to form the stalk of the
archegonium. The terminal cell functions as archegonial mother cell
(Fig. 8 B). It divides by three intersecting walls forming three
peripheral cells enclosing a tetrahedral axial cell (Fig. 8 C, D). The
peripheral cells divide anticlinally to form a single layered wall of
venter which later becomes two layered.

The axial cell divides, transversely to form an outer primary cover cell
and inner central cell (Fig. 8F). The outer primary cover cell functions
as aplical cell with four cutting faces (three lateral and one basal). It
cuts off three lateral segments and one basal segment. Each lateral
segment divides by a vertical wall so that the six rows of cells form the
neck of the archegonium (Fig. G, H). Each basal cell adds to neck canal
cell.

The inner central cell divides by transverse division into an outer

primary neck canal cell and an inner primary venter cell (Fig. 6H).
Primary neck canal cell undergoes transverse divisions to form a row
of neck canal cells.

Thus in Funaria the neck canal cells have double origin (lower and
middle neck canal cells in the neck canal are derived from the primary
neck canal cell while those in the upper portion of neck are derived
from the primary cover cell). The primary venter cell divides by
transverse division to form the venter canal cell and egg cell (Fig. 8 I,
J).
Fertilization in Funaria:
Water is essential for fertilization. The opercular cells of the
antheridium rupture and releases mass of antherozoids. When
archegonium reaches at maturity, the neck canal cells and venter canal
cell disintegrate to form a mucilaginous mass. It absorbs water, swells
up and comes out of the archegonial mouth by pushing the cover cells
apart. This mucilaginous mass consists chemical substances (mainly
sugars).

The cover cells of the neck separate widely from each other and form a
passage leading to the egg. Rosette like perigonial leaves serve as
splash cup from which rain drops disperse antheroziods to some
distance (rain drops falling on the archegonial cluster situated at lower
level).

Many antherozoids enter the archegonial neck because of chemical

response but only one of them fuses with the egg to form the zygote.
Union of male and female nuclei is complete within 10 hours.
Fertilization ends the gametophytic phase.
Sporophytic Phase:
Zygote is the first cell of the sporophytic phase. Development of
sporophyte takes place within the venter of the archegonium.

Structure of Sporophyte:
The sporophyte is semi-parasitic in nature, the mature sporophyte can
be differentiated into three distinct parts—foot, seta and capsule.

(i) Foot:
It is the basal portion of the sporogonium. It is small dagger like
conical structure embedded in the apex of female branch. It functions
as anchoring and absorbing organ.

(ii) Seta:
It is long, slender, stalk like hygroscopic structure. It bears the capsule
at its tip. It raises the capsule above the apex of leafy gametophore. Its
internal structure is more or less similar to axis. The epidermis is
followed by thick walled cortex which surrounds the axial cylinder. It
is mechanical in function and also conducts the water and nutrients to
the developing capsule (Fig. 10).

(iii) Capsule:
It is the terminal part of the sporophyte and is developed at the apex of
the seta. It is green in colour when young but on maturity it becomes
bright orange coloured. It is covered by a cap like structure called
calyptra. (gametophytic tissue develops from the upper part of the
archegonium).

Internal Structure of the Capsule:

Longitudinal Section (L.S.) of the capsule shows that it can be
differentiated into three distinct regions-apophasis, theca and
operculum (Fig. 11A).
(a) Apophysis:
It is the basal sterile part of the capsule. It is bounded by the single
layered epidermis which is interrupted by stomata. The sotmata have
single ring like guard cells (Fig. 11 C, D). Below the epidermis is
spongy parenchyma. The central part of the apophysis is made up of
elongated thin walled cells forming a conducting strand. It is called
neck of the capsule. It is the photosynthetic region and connects seta
with capsule.

(b) Theca:
It is the middle, slightly bent spore bearing region of the capsule. It
lies between the apophysis and operculum.

Longitudinal section (L. S.) passing through the theca shows

the following regions:
(i) Epidermis:
It is the outer most layer. It is single layered with or without stomata.
(ii) Hypodermis:
It is present below the epidermis. It consists two to three layers of
compactly arranged colourless cells.

(iii) Spongy parenchyma:

It consists two to three layers of loosely arranged chlorophyllous cells.
It is present inner to hypodermis. These cells are capable to
manufacture their own food but dependent on gametophyte for water
and mineral nutrients. Therefore, the sporophyte of Funaria is
partially dependent on gametophyte.

(iv) Air spaces:

These are present just below the spongy parenchyma and outside the
spore sacs. Air spaces are traversed by green cells (chlorenchymatous
cells) called trabecular (elongated parenchymatous cells).

(v) Spore sac:

These are present below the air spaces on either side of the columella.
It is ‘U’ shaped and broken at the base. (It separates its both arms).

It has an outer wall (3-4 cells thick) and an inner wall (single cell in
thickness). Between the outer wall and inner wall is the cavity of the
spore sac. When young, the cavity of the spore sac is filled with many
spore mother cells. At maturity the spore mother cells divide by
meiotic divisions and form many haploid spores.

(vi) Columella:
It is the central part of the theca region. It is made up of compactly
arranged colourless parenchymatous cells. It is wide above and narrow
below, connecting the central strand of apophysis. It helps in
conduction of water and mineral nutrients.
(c) Operculum:
It is the upper region of the capsule. It is dome shaped and consists
four to five layers of cells. The outermost layer is thick walled and
called epidermis. Operculum is differentiated from theca by a well-
marked constriction. Just below the constriction there is a diaphragm
(rim). It is composed of two to three layers of radially elongated pitted
cells.

Immediately above the rim is annulus which consists of 5-6

superimposed layers of cells. Its upper cells are thick but two
lowermost layers of cells are thin. Annulus separates the theca from
the operculum. Below the operculum lies the peristome (Fig. 11 B). It
is attached below to the edge of the diaphragm. The peristome consists
of two rings of radially arranged peristomial teeth. In each ring there
are sixteen teeth.

The teeth are not cellular but they are simply the strips of the cuticle.
The teeth of the outer ring are conspicuous, red with thick transverse
bands while the teeth of the inner ring are small, delicate, colourless
and without transverse bands. Inner to peristome teeth lies a mass of
thin walled parenchymatous cells. (Fig. 1 IB).

Development of Sporophyte:
Soon after fertilization, the zygote secretes a wall around it and
enlarges in size. It divides by a transverse wall forming an upper
epibasal cell and lower hypo basal cell (Fig. 12 A, B). Epibasal cell
divides by two intersecting oblique walls. It differentiates an apical cell
with two cutting faces in the epibasal cell (Fig. 12C). Similarly, the
hypo basal cell differentiates an apical cell (Fig. 12 D).

The entire sporophyte is differentiated by the activity of these two

apical cells. So, the development of embryo sporophyte is bi-apical.
Epibasal apical cell develops into capsule and upper portion of the seta
while the hypo basal apical cell develops into foot and remaining part
of the seta. Both apical cells cut out alternate segments and form the
elongated filamentous structure of young sporogonium (Fig. 12 E, F).
Development of Capsule:
A cross section through the upper portion of the young sporogonium
shows a two identical segments (Fig. 13A) which divide by a vertical
division at right angle to the previous one to form a quadrant (4 celled
stage) (Fig. 13B).

Each cell of the quadrant divides by anticlinal wall (Fig. 13C) in such a
way that a smaller almost triangular cell and a larger more or less
rectangular cell is formed. Each rectangular cell now divides by a
periclinal division (Fig. 13D).

It results in formation of a group of four central cells surrounded by 8

peripheral cells. The central tissue is known as endothecium and the
peripheral cells from the amphithecium (Fig. 13D). From these two
group of cells the further development takes place. There is formation
of different rings by anticlinal and periclinal divisions.

The amphithecium divides by periclinal division to form two

concentric layers. The inner layer of 8 cells is called first ring (Fig.
13E). The cells of the outer layer divide by anticlinal divisions to form
16 cells (Fig. 13F). This is followed by the periclinal division in this
layer.
The inner part of this layers is called the second ring (Fig. 13G). Again
the outer layer of these two layers divides anticlinally to form 32 cells.
This layer divides periclinally to form two layers of 32 cells. The inner
layer is called third ring. Similarly by periclinal divisions fourth and
fifth ring of 32 cells are formed (Fig. 13G-I).

The four cells of the endothecium also divides similarly to

amphithecium. The first division is curved and anticlinal (Fig. 13F).
The second division is periclinal (Fig. 13G). It results in the formation
of a central group of 4 endothecial cells, surrounded by 8 peripheral
endothecial cells. Further development of the tissues in the capsule
region takes place by these amphithecial rings and endothecial cells.
It can be studied under the following three headings (Fig.
13H-I). Development of fertile (theca) region (middle
portion of young capsule):

The fertile region in capsule comprises archesporium lined by outer

and inner spore sac. Archesporium is endothecial in origin. Its cells
may undergo sub-divisions to form two cell layers thick spore mother
cells which by meiosis form tetrad of spores. Elaters are absent.

Dehiscence of the Capsule:

Funaria is a stegocarpous moss (dehisce along a pre-determined line)
Dehiscence of the capsule is achieved by ‘breaking off’ of annulus. As
the capsule matures it becomes inverted due to epinasty. The thin
walled cells of the annulus break away, the operculum is thrown off
and the peristome teeth are exposed.

The outer peristomial teeth (exostome) are hygroscopic. The inner

peristomial teeth (endostome) do not show any hygroscopic
movements but act as a sieve allowing only a few spores to disperse at
a time. The lengthening and shortening of the outer peristomial teeth
help in the dispersal of spores.

In high humidity the exostome absorb water, increase in length and

curve inwards. In dry weather, the exostome teeth lose water, bend
outwards with jerky movements. It allows the dispersal of spores from
the capsule in instalments. At maturity the seta also shows jerky
movements. Twisting and swinging of seta in dry weather further aids
in the dispersal of spores.

Structure and Germination of Spore:

Spore is the first cell of the gametophytic phase. Each spore is
spherical (Fig. 15A), 12-20 µ in diameter and surrounded by two wall
layers (Fig. 16 A). The outer wall is thick, smooth, brown and known as
exosporium, while the inner wall is thin, hyaline and called
endosporium. Spore wall encloses single nucleus, chloroplasts and
many oil globules.
Under favourable conditions (sufficient moisture) spores germinate.
Exosporium ruptures and endosporium comes out in the form of one
or two germ tubes (Fig. 16B, C). Each germ tube is multicellular, green
with oblique septa. The germ tube grows in length, divides by septa to
form green algal filament like structure called primary protonema
(Fig. 16 D, E).

Primary Protonema:
It is the juvenile (young) stage of the gametophyte formed by the
germination of spore. It forms two different types of branches.

Most of the branches grow horizontally on the moist surface of the soil
and are known as chloronemal branches (positive phototrophic, thick
and rich in chloroplast) while some branches grow down in the soil
and are called rhizoidal branches (non-green, thin and possess oblique
septa) (Fig. 16F). These branches can develop chlorophyll if expose to
light.

Rhizoidal branches function as anchoring and absorbing organs while

chloronemal branches develop minute green buds behind the cross
walls which develop into leafy gametophores. From one primary
protonema many moss plants develop, so the moss is gregarious in
habit. Primary protonema is short lived.
According to Sirnoval (1947) development of protonema under
laboratory conditions can be differentiated into two stages—
chloronemal stage and caulonemal stage. Chloronemal stage is
characterised by irregular branching, right angle colourless cross
walls, and many evenly distributed discoid chloroplast.

It is positive phototropic but never produce buds. Nearly after 20 days

chloronemal stage matures into caulonemal stage. This stage is
characterised by regular branching brown cell walls, oblique cross
walls and fewer chloroplasts. It is negative phototropic and produce
buds which later develop into leafy gametophores. Rhizoids arise from
the base of a bud (Fig. 16G. 17, 18).