Bryopsida / Musci / Mosses

General characteristics

Bryopsida is the third largest class of the Bryophyta and consists of about 660 genera and 15000 species.

The gametophyte is differentiated into two stages:

-a branched, prostrate, juvenile filamentous or thalloid stage (protonema) which is usually transitory and
-an erect radial leafy shoot, the gametophore.

The gametophore consists of stem and spirally arranged leaves and bears sex organs.

Stem may be branched or unbranched. The branch always arise from below a leaf and is never axillary. Dichotomy is
unknown.

In majority of mosses, leaf has a mid rib (exception Sphagnum and Hypnum sp.).

• The leaves are arranged spirally in three ranks on the stem (1/3 divergence).

• In many cases 1/3 divergence is lost due to growth of the stem resulting divergence of 2/5 in Sphagnum,
3/8 in Funaria, 5/13 in Polytrichum and 4/11 in Dicranium.

• Rhizoids are branched, multicellular with oblique septa between the cells.

• Sex organs are stalked and the stalks are longer than those of other bryophytes.

• Early growth of the sex organs is by means of an apical cell.

• The sporophyte is more elaborate and complex and shows high degree of specialization and sterilization.
Except Sphagnum, the archesporium is differentiated from the outer layer of the endothecium and differentiated
into spores.

Note: In Anthoceros and Sphagnum; periclinal div. appears in the amphithecium dividing it into two regions.

Outer forms wall of the capsule and inner fertile region acts as an archesporium.

In other mosses: outermost layer of endothecium form spores while the central region forms columella.

Except Andraea, the capsule usually dehisces by the separation of a lid.

Except Sphagnum, Andreaea and some reduced forms of true mosses, peristome is present.
Classification
Bower (1935), Wettstein (1933-35) and Campbell (1940) divided the class Bryopsida into three orders
1. Sphagnales 2. Andreales 3. Bryales
Smith (1938, 1955) divided class Bryopsida into three sub-classes
1. Sphagnobrya 2. Andreaeobrya 3. Eubrya
Reimers (1954) recognized 5 sub-classes as
1. Sphagnidae: with single order Sphagnales
2. Andreaeidae: includes single order Andreales
3. Bryidae: comprises 12 orders
4. Buxbaumidae: single order Buxbaumiales
5.Polytrichidae: with two orders Polytrichales and Dawsoniales
Parihar (1955) divided the class Bryopsida into three sub classes
1. Sphagnidae
2. Andreaeidae
3. Bryidae
Latest classification was given by Buck and Goffinet (2000)
They have excluded liverworts and hornworts under division Marchantiophyta and Anthocerotophyta respectively.

Thus, for remaining mosses, the have assigned to division Bryophyta

Division Bryophyta
Class 1. Takakiopsida
2. Sphagnopsida 1. Diphysciidae
Sub-class
3. Andreaeopsida 2. Funariidae
4. Andreaeobryopsida 3. Dicranidae
5. Polytrichopsida 4. Bryidae

Order Takakiales

• Based on the characteristics of female gametophyte, Udar (1976) believed Takakia as a nearest
ancestors of Hepaticopsida.

• After the discovery of antheridia and sporophyte, Takakia is held to be the ancestor of the class
Bryopsida and occupies a separate rank sub-class Takakiidae, having single order Takakiales and
single family Takakiaceae which is monogeneric.
• General Characteristics:
• The gametophytic body is small bright, green and radially symmetrical.

• Body is differentiated into underground creeping rhizome and erect

leafy gametophore.
• Both the rhizome and leafy axes are completely devoid of rhizoids, instead,
from the base of erect gametophore one or more vertically descending leafless
branches arise and grows downwards.
The leaf like appendages born on the erect gametophore axis are termed as
‘phyllids’.

Phyllids are isophyllous and spirally arranged.

The gametophyte bears slime papillae of two types:

-Non beaked and axillary in position, 1-2 per leaf.
-Beaked and occurs in stolons only.
Note: mucilage is secreted through these papillae.
The gametophytes are heterothallic.

The antheridia are found in the axil of 3

to 4 lobed leaves.

The archegonia on the female plants

occurs singly or occasionally in a group of
2-3.

The archegonia has a short neck which

consists of 6 rows of neck cells.

The sporophyte is differentiated into foot, seta and capsule.

The developing sporophyte is protected by vaginula at the base and remnant

part of calyptra at apex.

The capsule lack operculum, apophysis, theca and peristome teeth.

The presence of columella, lack of elaters and pseudoelaters are the features
showing affinity of this group with the mosses.
Sub class- Sphagnidae
Members of this group are commonly called peat or bog moss.

The sub-class Sphagnidae includes a single order Sphagnales with a single family
Sphagnaceae represented by a single genus Sphagnum (more than 336 species).

The gametophytic body has two distinct stages:

i. Juvenile stage: It is formed by germination of spore (protonema).

Plate like thallose protonema is fixed to the substratum by multicellular
rhizoids (obliquely septate).
Leafy gametophyte or gametophore arise from the protonema.
As the gametophore grows, the rhizoids soon disappear.
II. Leafy gametophyte:
It consists of stem and leaves.
Plants growing on land possesses two
types of branches:
-Divergence: grow laterally and
-Drooping
Internally, the stem is differentiated into 3 zones:
-Cortex or hyalodermis: at maturity, they becomes dead and
helps to store water.
-The Hadrome (prosenchymatous region): supporting tissue.
-Medulla (axial cylinder): storage region.

The leaf consists of two kind of cells, the narrow, living,

green assimilatory cells and the large colourless dead
capillary cells.
Vegetative reproduction

a. Innovations: occasionally one of the branches in the

axillary cluster may turn upward and grows more
vigorously than others.
This branch takes on all the characteristics of the main
stem.

b. Multiplication of protonemal stage:

Any marginal cell of the thallose primary protonema
may become meristematic and give rise to secondary
protonema. Leafy gametophore arises from it.

c. Gemmae: These are subglobose and double walled.

Sexual reproduction
Sphagnum is monoecious or dioecious.

Antheridia
Antheridial branches are usually shorter but stouter than the
vegetative branches.

The antheridia occurs singly in the axil of

leaves towards the tip of antheridial
branches.
The mature antheridium consists of a globular body and
a long stalk (2 cells broad).

Body of antheridium has a jacket layer which encloses a

mass of androcyte mother cells.

Each androcyte mother cell divides into two androcytes

(division is not diagonal).
The androcyte metamorphoses into sperms.
Archegonia

The female branches are very short and more or less globular in
form.

They are green and bud-like aggregates of bracts and archegonia.

Some leaves of the terminal region forms perichaetium (protect

archegonia).

Archegonia occurs in a group of 2-5 (usually 3), of them the central

or the middle archegonium in the cluster grows directly from the
apical cells of the branch.

The body of archegonium consists of long twisted neck and a massive venter.
The neck consists of six vertical rows of neck cells.
The neck canal contains 8-9 neck canal cells.
The venter and the lower portion of the neck is 2-3 layers of cells in thickness.
The venter cavity contains venter canal cell and an egg.
Development of archegonium is similar to Jungermanniales.
Fertilization
Sporophyte phase
The mature sporophyte consists of a foot, suppressed seta and capsule.

Foot is bulbous structure and embedded in the tissue of the dilated apex
of the pseudopodium.
Capsule is a small globose structure and contains:
Columella:
Spore sac: dome shaped.
Capsule wall: 4-6 layered with chloroplast.
Epidermis: with rudimentary stomata.
Operculum / lid: it is marked off from rest of the capsule region by a
ring like groove and thin walled cells, the annulus.

Development of sporophyte
• The zygote divides by transverse wall and form 3 to 4 celled filamentous embryo.
• The lower portion of the embryo forms foot and the constriction like seta.
• Each segment of the upper part of the embryo undergoes two successive vertical division.
• Then, periclinal division differentiate the region into
-amphithecium and
-endothecium (forms columella)

• Amphithecium divides periclinally into:

-outer (forms wall of the capsule) and
-inner region ….. archesporium ……spore mother
cells……meiosis……spore tetrad.
Dehiscence of capsule
By explosive mechanism.
Germination of spore…..new plant through protonemal stage.