Metzgeriales

The order includes about 23 genera and 550 species (in Jungermanniales; about 220 genera and 8500 species).
It includes forms in which the archegonia occurs in group on the dorsal surface.
They originate behind the apical cell, i.e., apical cell never used up to form an archegonium.
Due to this activity, the position of archegonium and sporophyte are always dorsal.

The gametophytic body is usually thallus (Pellia, Riccardia and Pallavicinia lyelii), more rarely a stem with leaves (Fossombronia) and
some genera are intermediate between thallose and leafy form (Symphyogyna and Pallavicinia crispula).

Internally, the gametophytic body has little histological differentiation of tissues (differ from Marchantiales).
The scales are often absent and rhizoids are always smooth walled.

Vegetative reproduction takes place by:

i. Gemmae: in thallose form the gemmae produced on the dorsal surface of the thallus.
-sometimes on flask shaped receptacle (Blasia).
Gemmae are two celled in Riccardia, multicellular and star shaped or globose in Blasia, and circular plate
like in Metzgeria uncigera.

ii. Fragmentation
iii. Ventral adv. branch in Calycularia.
iv. Tubers in Fossombronia spp., Petalophyllum, Sewardiella and Symphyogyna.

The Jacket of the archegonial neck is usually composed of 5 vertical rows of cells, and the neck is often
almost as broad as the venter.
The jacket of the capsule is two or more layered in thickness.

The archesporium give rise to spores and elaters.

The sporogonium usually consists of foot, seta (much elongated at maturity) and capsule.

The mature capsule usually dehisces by four valves.

The order Metzgeriales comprising 23 genera and more than 550 species.

The classification of Metzgeriales is based on two important characteristic features of the gametophyte besides other from the
sporophytic generation. Those are:

a. The presence or absence of mid rib and leaf like appendages

b. And diverse position of the sex organs.

Mueller (1954) recognized 6 families; Pelliaceae, Blasiaceae,

Pallaviciniaceae, Metzgeriaceae, Riccardiaceae, and Monocleaceae but Schuster (1966) listed 8. Additional families are
Treubiaceae and Fossombroniaceae.

Family Pelliaceae

Characteristic features:
The plant body is thallus which is often lobed by irregular incisions.

The antheridia and archegonia are scattered on the dorsal surface of the thallus.

The capsule is either spherical or oval, and has a basal elatophores.

Family Riccardiaceae

Gametophytes are thallose.

The cells of the thallus have distinct, finely segmented oil bodies.

The sex organs are dorsal on short lateral branches.

The involucre is absent but the calyptra is large, thick, fleshy and well developed.

The capsule is ovoid to cylindrical.

There is a distal elatophores, to which some of the elaters are attached.

Family Fossombroniaceae
Plants thalloid or leafy.

Rhizoids purplish or brownish.

Antheridia scattered, archegonia scattered or clustered.

Sporophytes protected by a shoot calyptra and either a caulocalyx or perichaetial pseudoperianth.

Capsules spheroidal, lacking an elaterophore, dehiscence irregular or in 5–7 unequal segments.

Pellia

The mature gametophyte

The gametophytic body is simple, thin, prostrate, dorsivental, dichotomously branched thallus with
a sinuous (wavy) margin.

Dorsal surface with distinct midrib.

Ventral surface with numerous unicellular smooth walled rhizoids.

Tuberculate rhizoids and scales are absent.

Internal structure
The internal structure of the thallus is very simple, and mainly consists of parenchymatous
cells.

The thallus is multilayered at the mid rib region while single layered in wings.

The cells of the wings and the upper layer of the midrib contain abundant chloroplast,
whereas the lower cells of the midrib contain very few or no chloroplasts.

The pores and air chambers are absent.

Development of antheridium

The antheridial initial (develops on the dorsal surface of the thallus) divides by ransverse wall into a basal cell and an outer cell.

The outer cell divides transversely into lower primary stalk cell (forms stalk) and upper primary antheridial cell.

Meanwhile, the dorsal cells of the thallus (that surrounds antheridium) divide repeatedly and produce a ring shaped involucre.

Primary antheridial cell divides vertically into two daughter cells which again divides by nearly periclinal wall into two unequal
cells.

The smaller of these cells is the first jacket initial.

The large cell again divides by another periclinal wall and form outer 2nd jacket initial and an inner primary androgonial cell.
This is the sequence of early stages of development of the antheridium in almost all the Jungermanniales.
Two primary androgonial cells repeatedly divide and form large number of androgonial cells……..androcyte mother cell…….2 diagonal
androcyte…. Antherozoids.

Mature antheridium is similar to the members of Marchantiales.

Rest of the divisions and developmental pattern is similar to the Marchantiales.
The archegonium
The archegonia are formed in groups
of 4-12 at the anterior end of a branch just behind the growing point.

The group of archegonia are protected by an involucre.

Develoment of archegonium
The archegonial initial divides transversely into outer and basal cell.

As in Riccia, three intersecting vertical walls are laid down in the outer
cell, thus forming a central primary axial cell surrounded by three peripheral initials.
Among the three, smaller remains undivided while two others divide vertically, which leads to the formation of 5 jacket initials
(6 in Marchantiales).

Each jacket initial divides transversely into an upper neck initial (forms neck) and the lower ventral initial (forms venter).
The primary axial cell divides transversely And form pri. Cover cell (upper) and central cell (lower).

Rest of the steps are similar to the Marchantiales.

Mature Archegonium
It has short multicellular stalk, a venter and a long neck.
The jacket of the neck consists of five ventral rows of cells and encloses usually
6-8 neck canal cells.

Just before fertilization, the venter and sometimes the lower portion of the neck
become two layered in thickness.

Syngamy- is similar to Riccia.

Development of sporogonium

The zygote divides transversely and form upper epibasal and lower hypobasal cell.
Hypobasal cell does not divide and forms a suspensor, which functions as haustorium.
All the tissues of mature sporogonium i.e. foot, seta and capsule are derived from epibasal cell.
The epibasal cell first divides vertically then transversely and form 4 cells.
The four cells divides by vertical walls and form two tier of 4 cells.
The four cells of the lower tier divides repeatedly and form foot and seta.

Periclinal wall formation in the upper four cells results the formation of peripheral amphithecium and central endothecium.
The endothecium forms the archesporium and after repeated division form a mass of sporogenous cells.
The sporogenous cells differentiate into elatophores, elaters and spores.