The sexual generation, the gametophyte. The protonema in the Liverworts is very insignificant, and not always very sharply demarcated from the more highly developed parts of the nutritive system. In the true Mosses the protonema is well-developed, and consists of a branched, alga-like filament of cells, the dividing cell-walls being always placed obliquely. In the parts exposed to the light it is green, but colourless or brownish in those parts which are underground (Fig. [186]). The protonema is considered to be a lower form of the stem, and grows in the same manner by means of an apical cell; at its apex it may directly develope into a leaf-bearing stem, or these arise from it as lateral branches (Fig. [186] k).
The more highly differentiated part of the vegetative system, the “Moss-plant,” which is thus developed from the protonema, is in the “thalloid” Liverworts generally a dichotomously-branched thallus without any trace of leaf-structures (Fig. [194]); in Marchantia (Fig. [197]) and others, scale-like leaves (amphigastria) are found on the under surface. The higher Liverworts and the Leafy-Mosses are differentiated into a filamentous, ramified stem with distinct leaves arranged in a definite manner, resembling the stem and leaves of the higher plants (Figs. [186], [195], [200]).
True roots are wanting, but are biologically replaced by rhizoids. These are developed on the stems or thallus: in the Liverworts they are unicellular, but in the Leafy-Mosses generally multicellular and branched. In the latter group they are considered identical with the protonema, and may become true protonema, and new plants may be developed from them (Fig. [186] b).
Fig. 186.—A Lower portion of a Moss-plant with rhizoids (r), one of which bears a reproductive bud (b). The dotted line indicates the surface of the ground; the portions projecting above this become green protonema (p); k is a young Moss-plant formed on one of these. B Germinating spore of Funaria hygrometrica, with exospore still attached. C, D Older stages of the protonema.
The internal structure of the sexual generation is very simple. The leaves in nearly all cases are formed of a single-layered plate of cells; in the Leafy-Mosses, however, a midrib is very often formed, and sometimes, also, marginal veins; and along these lines the leaves are several layers of cells in thickness. The stem is constructed of cells longitudinally elongated, the external ones of which are narrower and sometimes have thicker walls than the more central ones. Vessels are not found, but in several Mosses there is in the centre of the stem a conducting strand of narrow, longitudinal cells, which represents the vascular bundle in its first stage of development. This strand contains elements for conveying water as well as sieve-tubes. Stomata are entirely wanting in the sexual generation of the Leafy-Mosses; they are found in a few Liverworts (Marchantia), but their structure is not the same as in the higher plants.
Vegetative reproduction takes place by gemmæ or buds which arise on the protenema, the rhizoids, the thallus, or the shoots, and become detached from the mother-plant; or else the protonema and the older parts of the plant simply die off, and their branches thus become independent plants. This well-developed vegetative reproduction explains why so many Mosses grow gregariously. In certain Marchantiaceæ special cupules, in which gemmæ are developed, are found on the surface of the thallus (Fig. [197] A, s-s). Again, protonema may also arise from the leaves, and thus the leaves may act as reproductive bodies. Certain Mosses nearly always reproduce vegetatively, and in these species the oospheres are seldom fertilised.
Fig. 187.—Marchantia polymorpha: a mature antheridium.