RANGE OF VEGETATIVE AND

REPRODUCTIVE STRUCTURES,
MODE OF REPRODUCTION IN
LIVERWORTS - METZGERIALES
BY
M . VA I S H N AV I
M.SC PB&PBT
REG NO.219PB00398
 OBJECTIVE
INTRODUCTION
DISTRIBUTION
CHARACTERISTIC FEATURE
REPRODUCTION
PELLIA – TYPE STUDY
COMPARISON OF METZGERIALES (PELLIA) AND MARCHANTIALES (MARCHANTIA)
 INTRODUCTION
Systematic Position [ According to Schuster (1953,1958) ]
Kingdom : Plantae
Division : Bryophyta
Class : Hepaticopsida
Sub-Class : Jungermanniae
Order : Metzgeriales (23 Genera, 550 Species)

Muller (1954) recognised 6 families. But, Schuster (1966) listed 8 families.

Important families are
a) Pelliaceae : Pellia , Calycularia , Noteroclada
b) Riccardiaceae : Riccardia , Cryptothallus
c) Blasiceae : Blasia
d) Metzgeriaceae : Metzgeria
e) Pallaviciniaceae : Pallavicinia , Moerckia , Symphyogyna
f) Treubiaceae : Treubia
g) Fossombroniaceae : Fossombronia , Sewardiella , Petalophyllum

The Genus name which are Red colored represents the 10 Genera which are
commonly found in India.
PICTURES OF SOME OF THE IMPORTANT GENERA OF ORDER METZGERIALES

Pellia Calycularia crispula Noteroclada

Riccardia Cryptothallus Blasia Metzgeria

Pallavicinia Petalophyllum
Symphyogyna Treubia

Moerckia Fossombronia
 DISTRIBUTION
a) They are Widespread in distribution.
b) Occurs in temperate and tropical regions of the world.
c) 10 Genera in India – Himalayas and South India.
d) Species are hygrophytic and mesophytic.
e) Combination of shade and abundant moisture is
precondition for successful growth.
f) Grow on damp soil, moist rocks, bark of trees.
 CHARACTERISTIC FEATURE
a) Thallus is of Diversified form.
1) Thalloid form – showing wavy margin, dichotomous branching.
eg. Pellia, Riccardia, Pallavicinia.
2) Leafy Form – Gametophyte differentiated into stem and lateral leaves.
eg. Fossombronia
3) Transitional form – Intermediate between thallose and leafy forms.
Thallus with distinct midrib , wings deeply insist to form lobes which
are not distinct enough to be leaves.
eg. Symphyogyne brongniartii
b) Mostly Gametophytes are without internal differentiation of tissues
but certain genera have central strand of thick walled cells.

c) The ventral surface bears smooth walled rhizoids.

Portion of T.S of Thallus
Pellia epiphylla Riccardia
d) The sex organs occur in single or group on upper surface of thallus
(pellia) or on short ventrally / side branches.

e) Archegonia do not develop from apical cells. It arise from the young
segments cut off by the apical cells – Anacrogynous.

f) Thallus continues to grow , as a result archegonia and sporophytes

are dorsal in position.
Section through gametophyte showing sex organs in
Fossombronia
Pellia
g) The cluster of archegonia are protected by tubular or funnel like out
growth of adjacent thallus tissues – Involucre.

h) Capsule wall – 2 to 5 layers of cell thick.

Sporophyte of
Riccardia Pellia
 REPRODUCTION
A) VEGETATIVE REPRODUCTION
1) Gemmae – Bud like structures variable in form and size.
a) Thallose Forms – Dorsal surface Exogenous
Endogenous
Within specialized cell
b) Special gemmiferous branches or flask shaped receptacles.
eg. Blasia pusilla

Two types of Gemma 1) multi cellular star shaped

2) multi cellular globose, stalked

c) Small, 2- celled gemmae eg. Riccardia mutifitida

d) 3-4 celled structure gemma eg. Treubia

e) Flat, plate like multi cellular gemmae on branches of

thallus called gemmiferous branches.
eg. Metzgeria fruticulosa

f) Gemma on leaf margin eg. Radula complanata

2) Fragmentation :
By progressive growth followed by the death of the older regions of the
thallus, the young thallus lobes or branches of the dichotomy become isolated.
The isolated branches are called fragments. Each fragment by apical growth
and branching, grows into new plant. eg. Pellia

Adventitious branches of
3) Adventitious Branches : Calycularia crispula
Some thallose Metzgeriales produce specialised
adventitious branches which arises ventrally get
detached and serve as means of vegetative propagation.
eg. Calycularia crispula
4) Tubers:
 Apical tubers develop at the end of growing season.
 Tip of the plants grows down wards into the soil and thickens to form a
globose or cylindrical mass of cells called tubers.
 It develops corky wall which protect it from desiccation.
 Inner tissues store reserve foods in the form of starch, oil globules.
 The tubers remain dormant over the dry period. With the return of
favourable conditions it resumes its activity.
eg. Fossombronia himalayensis
B) SEXUAL REPRODUCTION
ANTHERIDIUM
1) Arrangement
a) Single
b) Scattered in rows e.g.(Pellia)
c) Groups e.g.(Calycularia cripsula on dorsal surface)
d) Ventral lobes – Metzgeria
e) Side branches – Riccardia pingus
f) Sunken pits – Riccardia, Pellia

2) Structure
a) Antheridium globose , stalked – short multicellular.
b) Sac encloses mass of Androcyte mother cells.
c) Wall of sac – one layer of cells thick.
d) Sperms - Biflagellate with more coils.
Pellia- Antheridium
ARCHEGONIUM

1)Arrangement / Position
a) Single eg. Riccardia
b) Groups eg. Pallavicinia
c) Groups laterally on midrib near growing point protected by young leaves.
eg. Fossombronia.

2) Structure
a) Archegonia have distinct stalk.
b) Venter is slightly broader then neck.
c) It consists of 5 vertical rows of neck cells.
d) Group of Archegonium is covered by envelope called
Involucre or perichaetium formed by upward growth
of adjacent thallus tissue.
Fossombronia- Archegonium
 Sporophyte-Fossombronia
SPOROPHYTE
It is differentiated into 3 parts.
i) Absorbing and anchoring swollen base foot.
ii) A long stalk like seta.
iii) The terminal sac like capsule containing spores & elaters.

a) Foot Pellia
• Usually anchor shaped in longitudinal section absorbs
nutrients from the gametophytes.
b) Seta
• Metzgeriales has long fragile transparent seta is present.
 Pellia
c) Capsule
• Dark mature capsules is variable in form.
• Globose – Pellia, Fossombronia
• Ovoid-cylindrical Blasia , Riccardia.
• Capsule wall 2 or more layers of cell thick.
• No columella.
• Sporogenous tissue forms spore mother cells & sterile cells.
• Sterile cells metamorphosed to elaters.
Fossombronia
• In adition to free elaters some genera have tuft of fixed elaters
at the base of capsules (Pellia) , apex of capsule (Riccardia).
DEHISCENCE
a) Dehiscence is regular manner.
b) Splits by four valve usually to the base.
c) Tuft of fixed elater stands at base of dehisced capsules .eg. Pellia , remain attached
to tip of each valve . eg. Riccardia

Capsule & Dehisced Capsule liberating spores

Pellia
Riccardia
 PELLIA – TYPE STUDY

Division : Bryophyta
Class : Hepaticopsida
Order : Metzgeriales
Genus: Pellia
Species: P.calycina, P. epiphylla, P. neesiana

SEXUAL REPRODUCTION (IN DETAIL)

P.calycina , P. neesiana - Dioecious
P. epiphylla - monoecious ( protandrous)
ANTHERIDIA

1) Arrangement / Position
a) Occur Singly , irregularly scattered in more rows on midrib region.
b) Antheridium is present in sunken condition inside antheridial chamber.
c) Antheridial chamber has opening called ostiole.
d) Sunken condition – efficient protection , discharge of sperms.

2) Structure
a) Antheridium is stalked globular structure.
b) Stalk – short, slender , multicellular.
c) Globular body  outer wall of one cell layer thick.
 inside a central mass of androcyte mother cell.
3) Development of Antheridium
Singly superficial cell (Antheridia initial) (A)
a)Basal Cell
Transverse Division
b) Antheridial mother cell (B)
a) Primary Stalk Cell -Stalk Transverse Division
b) Primary antheridial Cell(C)
Longitudinal Division
a) Daughter Cell b) Daughter Cell (D) Periclinal Division
Smaller jacket or wall initial Larger sister Cell (F) Smaller jacket or wall initial Larger sister Cell
Outer IInd jacket or Inner primary Outer IInd jacket or Inner primary
wall initial (G) Androgonial cell (H) wall initial Androgonial cell

Periclinal Division
Therefore, there is 4 wall initials covering the 2 primary Androgonial cells.
Anticlinal Division
Wall initial wall of antheridium (I)
Re-divide several times
Primary Androgonial cell Spermatocytes
or androcyte mother cell
Divides Diagonally (K)
Spermatids / Androcytes

Antherozoid (L)
• Antherozoid formed mainly from nuclear material.
• Furnished with two long whiplash flagella.
• Flagella are inserted at thin, anterior end of tapering criled body of Antherozoid.
4) Dehiscence of Antheridium
a) On access of moisture, wall cells of mature antheridium imbibe water and
become distended.
b) Distended wall cell abutting against the walls of antheridial pits cause
pressure to the contents within antheridium.
c) Mature antheridium ruptures at apex.
d) Sperms are liberated into the water by dissolution of the walls of androcytes.
ARCHEGONIA
1) Position
a) Present in cluster on the upper surface of thallus lobes.
b) Archegonia in clusters.
c) Archegonia stands on slightly raised transverse ridge of tissue – repectacle.
d) Cluster is surrounded by a complete collar like or incomplete flap like sheath-
Involucre.
e) Involucre is green ,protective in function. It is tubular – P.calycina ,
short, cylindrical – P-neesiana, reduced to pouch – P.epiphylla.
2) Structure
a) Archegonium – flask shaped structure.
b) Archegonium is seated on short, stout stalk.
c) The venter consists of a single layer of cells in thickness prior to
fertilization. At the time of fertilization it is 2 cell layer thick.
d) The venter cavity contains egg, small ventral canal cells.
e) Neck consists of 5 longitudinal rows of cells.
f) Canal enclosed by 6-9 neck canal cells.
g) The rosette of 4 large cover or cap cells at the top of neck.
h) Sometimes lower portion of neck become 2 layers thick.
3) Development : Archegonium develops from younger segments of apical cells –
archegonial initial. It enlarges into papilla like outgrowth.
Archegonial Initial(A)
Basal pedicel cell Upper Archegonial mother cell (B)
(Divides by 3 vertical walls)
Stalk 3 Peripheral Initial (D) Middle primary Axial Cell
(2 Large,1 small) Transverse Division

(vertical wall) outer cover cell lower central cell(F)

5 jacket initial do not divide upper primary Lower primary
upper neck initial lower venter initial neck canal cell ventral cell
Transverse Division
neck canal cell upper ventral Egg Cell
neck venter canal cell
Stages of Development of Archegonium
FERTILISATION
a) It takes place in the presence of water.
b) Axial rows of cells disorganise and mucilage is formed.
c) Mucilage absorb water , forces open the neck of archegonium.
d) Antheridium take up water , swells, bursts liberating antherozoids.
e) Antherozoids swims and enters the neck of archegonium.
f) One of the antherozoid enters the egg , loses flagella.
g) The Union of nucleus of antherozoid takes place with that of egg.
h) Fertilized egg scretes wall –zygote or oospore(2n).
SPOROPHYTE
1)DEVELOPMENT OF SPOROGONIUM
5 or 6 days after fertilization of zygote(A) undergoes segmentation.
Oospore(A) Transverse Division

Upper epibasal cell Lower hypobasal cell haustorium or suspensor (C)

Vertical and Transverse Division
2 tiers of 4 cell each

Lower tier of 4 cell Upper tier of 4 cell Periclinal Division

Foot, Seta Peripheral Amphithecium Central Endothecium

Capsule wall Archesporium

Mass of sporogenous wall

Spores and Elaters

Stages of Development of Sporogonium
 Stages of Sporogenesis
2)SPOROGENESIS

Spore mother cells(2n)(A)

Four lobed structure(D)

Spore tetrad (n)(surrounded by common sheath)(G)

Spores(n)

Calyptra is developed around young sporangium. Seta is quite short.

3) STRUCTURE OF MATURE SPOROGONIUM
Mature Sporogonium
At maturity, it consists of foot, seta, capsule.
a) Foot:
1) Basal absorbing organ.
2) Distinct, conical in forms.
3) forms collar- like structure around base of seta.
b) Seta:
1) Pure white colour.
2) At maturity, it is almost transparent.
3) It terminates in dark green or black capsule.
4) It attains a length upto 5cm.
c) Capsule:
1) Globular with diameter of 1.5mm.
2) Two more cell layers thick. The outer layer consists of
polygonal cells. Radial walls have rod like shrinkage bands.
3) The second layer is flattened, reticulately thickened.
4) The Spores and elaters are intermixed capsule.
5) Capsule wall has vertical strips of thick wall cells on
surface line of dehiscence.

Capsule Showing Spores And Elaterophores

d) Elaters:
1) They are long, slender, spindle shaped cells.
2) Bands of spiral thickening on their wall.
3) They are hygroscopic spore dispersal.
4) There is prominent basal central tuft of 50-100 elaters attached to
center of the base of capsule. This elater cluster is erroneously called
as Elaterophores.

Spores Mixed Up With Elaters

NUTRITION AND PROTECTION
1) Nutrition to sporogonium is provided by foot housed in the thalloid
gametophyte.
2) Protection is given by Calyptra (an enlargement of Archegonia venter
tissue) and involucre.
DEHISCENCE OF CAPSULE
1) Dehiscence of Capsule - by elongation of the seta.
2) Calyptra is a ruptured .
3) Drying effect of surrounding air initiate the hygroscopic movement in
capsule wall.
4) Splitting of capsule along the line of dehiscence into 4 valves.

Capsule Showing Line Of Dehiscence

DISPERSAL OF SPORES
1) The separation of spores from exposed mass is assisted by
hygroscopic movement of elaters.
2) The flicking to and fro of the sessile elaters facilitates gradual
dispersal of loosened spores.
3) When all spores are shed, the elaterophore persists.
Dehisced Capsule with recurved valves Dehisced Capsule liberating spores
 GERMINATION OF SPORE
Spores begin to germinate before they shed, i.e. precocious germination of spores.
By the time, capsule dehisces , several cells of protenema are already formed within exospore.
Exospore (A)
Transverse Division
2 cells(B)
Transverse Division
4 cells (contain apical, basal cells)(C)
2 central cells are produced (D) Vertical Division

6 – 9 celled ovoid mass (E) – (F)

Liberation of spores
Basal cell grows into rhizoid (H)
Central cells undergo longitudinal division (I)
Cyl. Stem like structure (K)
Flattened thallus (L)
PELLIA – GRAPHIC LIFE CYCLE
COMPARISON OF METZGERIALES (PELLIA) AND MARCHANTIALES (MARCHANTIA)

METGZERIALES (PELLIA) MARCHANTIALES (MARCHANTIA)

1. Upper surface of thallus is smooth. There 1. It has areolae marked by dot like central
is no areolae. air pores.

2. No scales on lower surface of thallus. 2. Scales are present.

3. Smooth rhizoids are present whereas 3. Both are present.

tuberculate rhizoids are absent.

4. There is no or little differentiation of 4. Well differentiated thallus. Air pores, air

tissues of thallus. Air pores, air chambers chambers are present.
are absent.
METGZERIALES (PELLIA) MARCHANTIALES (MARCHANTIA)

5. Archegonial neck consists of 5 5. It consists of 6 vertical rows of cells.

vertical rows of cells.

6. Antheridia and archegonia are 6. They are produced on specialized

present on the surface of thallus. branches called antheridiophores and
archegoniophores.

7. Involucre present. 7. Involucre absent.

8. Elaterophores are present in 8. Elaterophores are absent . Instead

addition to in addition to sterile elaters. sterile elaters are present.
 REFERENCES
1) B.R,Vashishta,Botany For Degree Students Bryophyta,8 ed.,S Chand And Company,New
Delhi,1999
2) B.P. Pandey,College Botany,S Chand And Company,New Delhi,1979
3) https://www.sciencephoto.com/media/663133/view/liverwort-pellia-epiphylla-sporophyte
4) https://alchetron.com/Metzgeriales
5) https://www.alamy.com
6) https://en.wikipedia.org
7) http://www.preservearticles.com/botany/3-most-important-classes-of-bryophyta/30331
8) http://www.biologydiscussion.com/bryophyta/classification-bryophyta/classification-of-
bryophytes/21255