The range of form and structure of both generations in the liverworts is so great that no one form can be taken as a satisfactory type. It will, however, be of use to preface the more general description by a brief account of a particular example, and we may take for this purpose a very common and easily recognized thalloid liverwort belonging to the Jungermanniales.Fig. 3. - Pellia epiphylla.
A, Longitudinal section of thallus at the time of fertilization. an, Antheridia; ar, archegonia; in, involucre.
B, Longitudinal section of almost mature sporogonium attached to the thallus. in, Involucre; cal, calyptra; f, foot; s, seta; caps, capsule (semi-diagrammatic).
Fig. 2. - Pellia epiphylla. Group of plants bearing mature sporogonia.
From Cooke, Handbook of British Hepaticae.
Pellia epiphylla (fig. 2) can be found at any season growing in large patches on the damp soil of woods, banks, etc. The broad flat thallus is green and may be a couple of inches long. It is sparingly branched, the branching being apparently dichotomous; the growing point is situated in a depression at the anterior end of each branch. The wing-like lateral portions of the thallus gradually thin out from the midrib; from the projecting lower surface of this numerous rhizoids spring. These are elongated superficial cells, and serve to fix the thallus to the soil and obtain water and salts from it. No leaf-like appendages are borne on the thallus, but short glandular hairs occur behind the apex. The plant is composed throughout of very similar living cells, the more superficial ones containing numerous chlorophyll grains, while starch is stored in the internal cells of the midrib. The cells contain a number of oil-bodies the function of which is imperfectly understood. The growth of the thallus proceeds by the regular segmentation of a single apical cell.
The sexual organs are borne on the upper surface, and both antheridia and archegonia occur on the same branch (fig. 3, A). The antheridia (an) are scattered over the middle region of the thallus, and each is surrounded by a tubular upgrowth from the surface. The archegonia (ar) are developed in a group behind the apex, and the latter continues to grow for a time after their formation, so that they come to be seated in a depression of the upper surface. They are further protected by the growth of the hinder margin of the depression to form a scale-like involucre (in). Fertilization takes place about June, and the sporogonium is fully developed by the winter. The embryo developed from the fertilized ovum consists at first of a number of tiers of cells. Its terminal tier gives rise to the capsule, the first divisions in the four cells of the tier marking off the wall of the capsule from the cells destined to produce the spores. In fig. 4, C, which represents a longitudinal section of a young embryo of Pellia, these archesporial cells are shaded. The tiers below give rise to the seta and foot. The mature sporogonium (fig. 3, B) consists of the foot embedded in the tissue of the thallus, the seta, which remains short until just before the shedding of the spores, and the spherical capsule.
It remains for long enclosed within the calyptra formed by the further development of the archegonial wall and surmounted by the neck of the archegonium. The calyptra is ultimately burst through, and in early spring the seta elongates rapidly, raising the dark-coloured capsule (fig. 2). In the young condition the wall of the capsule, which consists of two layers of cells, encloses a mass of similar cells developed from the archesporium. Some of these become spore-mother-cells and give rise by cell division to four spores, while others remain undivided and become the elaters. The latter are elongated spindle-shaped cells with thick brown spiral bands on the inside of their thin walls. They radiate out from a small plug of sterile cells projecting into the base of the capsule, and some are attached to this, while others lie free among the spores. The latter are large, and at first are unicellular; but in Pellia, which in this respect is exceptional, they commence their further development within the capsule, and thus consist of several cells when shed.
The cells of the capsule wall have incomplete, brown, thickened rings on their walls, and the capsule opens by splitting into four valves, which bend away from one another, allowing the loose spores to be readily dispersed by the wind, assisted by the hygroscopic movements of the elaters. On falling upon damp soil the spores germinate, growing into a thallus, which gradually attains its full size and bears sexual organs.
Fig. 4. - Semi-diagrammatic figures of young embryos of Liverworts in longitudinal section. The cells which will produce the sporogenous tissue are shaded. (After Kienitz-Gerloff and Leitgeb.)
B, Marchantia polymorpha.
C, Pellia epiphylla.
D, Anthoceros laevis.
E, Cephalozia bicuspidata.
F, Radula complanata.
While the general course of the life-history of all liverworts resembles that of Pellia, the three great groups into which they are divided differ from one another in the characters of both generations. Each group exhibits a series leading from more simple to more highly organized forms, and the differentiation has proceeded on distinct and to some extent divergent lines in the three groups. The Marchantiales are a series of thalloid forms, in which the structure of the thallus is specialized to enable them to live in more exposed situations. The lowest members of the series (Riccia) possess the simplest sporogonia known, consisting of a wall of one layer of cells enclosing the spores. In the higher forms a sterile foot and seta is present, and sterile cells or elaters occur with the spores. The lower members of the Jungermanniales are also thalloid, but the thallus never has the complicated structure characteristic of the Marchantiales, and progress is in the direction of the differentiation of the plant into stem and leaf. Indications of how this may have come about are afforded by the lower group of the Anacrogynous Jungermanniaceae, and throughout the Acrogynous Jungermanniacae the plant has well-marked stem and leaves.