produces spores and is diploid. Hence all such plants are to this extent polymorphic—that is, the plant assumes these two forms in the course of its life-history. When, as in many Thallophyta, one or other of these forms can reproduce itself by means of gonidia, additional forms may be introduced into the life history, which becomes the more complicated the more pronounced the polymorphism.
The most straightforward life-histories are those presented by the Bryophyta and the Pteridophyta, where there are but the two forms, the sexual and the asexual. In the life-history of a moss, the plant itself bears only sexual organs: it is the sexual form, and is distinguished as the gametophyte. The zygote (oospore) formed in the sexual act develops into an organism, the sporogonium, which is entirely asexual, producing only spores: it is distinguished as the sporophyte. When these spores germinate, they give rise to moss-plants. Thus the two forms, the sexual and the asexual, regularly alternate with each other—that is, the life-history presents that simple form of polymorphism which is known as alternation of generations. Similarly, in the life-history of a fern, there is a regular alternation of a sporophyte, which is the fern-plant itself, with a gametophyte, which is the fern-prothallium.
It is pointed out in the preceding section that, as the result of the sexual act, the nucleus of the zygote contains twice as many chromosomes as those of the fusing sexual cells. This 2x number of chromosomes persists throughout all the cell-generations derived from the zygote, that is, in the cells constituting the sporophyte, up to the time that it begins to produce spores, when meiosis takes place. Again, the cell-generations derived from the spore, that is, the cells constituting the gametophyte, all have the reduced x number of chromosomes in their nuclei up to the sexual act. Hence the sporophyte may also be designated the diplophyte and the gametophyte the haplophyte (Strasburger): in other words, the sporophyte is the pre-meiotic, the gametophyte the post-meiotic generation. Twice in its life-history the plant is represented by a single cell: by the spore and by the zygote. The turning-points in the life-history, the transitions from the one generation to the other, are (1) meiosis, (2) the sexual act.
The course of the life-history in Phanerogams and in those Thallophyta which have been adequately investigated is essentially the same as that of the Bryophyta and of the Pteridophyta as described above, though it is less easy to trace on account of the peculiar relation of the two generations to each other in the Phanerogams and on account of various irregularities that present themselves in the Thallophyta.
In the Phanerogams, as in the Pteridophyta, the preponderating generation is the sporophyte, the plant itself. Inasmuch as they are heterosporous, the gametophyte is represented by a male and a female organism or prothallium, both rudimentary. The male prothallium consists of the few cells' formed by the germinating pollen-grain (microspore); and though it is quite independent, since the microspores are shed, it grows parasitically in the tissues upon which the microspore has been deposited in pollination. The female prothallium may consist of many cells with well-developed archegonia, as in the Gymnosperms, or of only a few cells with the female organ reduced to the oosphere, as in the Angiosperms. In either case it is the product of the germination of a megaspore (embryo-sac) which is not shed from its sporangium (ovule): hence it never becomes an independent plant, and was long regarded as merely a part of the sporophyte until its true nature was ascertained, chiefly by the researches of Hofmeister, who first explained the alternation of generations in plants. This intimate and persistent connexion between the two generations affords the explanation of the characteristic features of the Phanerogams, the seed and the flower. The ovule containing the embryo-sac, which eventually contains the embryo, persists as the seed—a structure that is distinctive of Phanerogams, which have, in fact, on this account been also termed Spermatophyta. With regard to the flower, it has been already mentioned that it is, like the cone of an Equisetum or a Lycopodium, a shoot adapted to the production of spores. But it is something more than this: for whereas in Equisetum or Lycopodium the function of the cone comes to an end when the spores are shed, the flower of the Phanerogam has still various functions to perform after the maturation of the spores. It is the seat of the process of pollination—that is, the bringing of the pollen-grain by one of various agencies into such a position that a part (the pollen-tube) of the male prothallium developed from it may reach and fertilize the oosphere in the embryo-sac. Thus the flower of Phanerogams is a reproductive shoot adapted not only for spore-production, but also for pollination, for fertilization, and for the consequences of fertilization, the production of seed and fruit. However, in spite of these complications, it is possible to determine accurately the limits of the two generations by the observation of the nuclei. The meiosis preceding the formation of the spores marks the beginning of the (haploid) gametophyte, male and female; and the sexual act marks that of the (diploid) sporophyte.
The difficult task of elucidating the life-histories of the Thallophyta has been successfully performed in certain cases by the application of the method of chromosome-counting, with the result that alternation of generations has been found to be of general occurrence. To begin with the Algae. In the Dictyotaceae (Brown Algae) there are two very similar forms in the life-history, the one bearing asexual reproductive organs (tetrasporangia), the other bearing sexual organs (oogonia and antheridia). It has been shown (Lloyd Williams) that the former is undoubtedly the sporophyte and the latter the gametophyte, since the nuclei of the former contain 32 chromosomes, and those of the latter 16. Meiosis takes place in the mother-cell of the tetraspores, which, on germination, give rise to the sexual form. Quite a different life-history has been traced in Fucus, another Brown Alga. Here no spores are produced: there is but one form in the life-history, the Fucus-plant, which bears sexual organs and has, on that account, been regarded as a gametophyte. The investigation of the nuclei has, however, shown (Farmer) that the Fucus-plant is actually diploid, that it is, in fact, a sporophyte; but since there is no spore-formation, meiosis immediately precedes the development of the sexual cells, which alone represent the gametophyte (see below, Apospory).
Similarly, two types of life-history have been discovered in the Red Algae. In Polysiphonia violacea, a species in which the tetraspores and the sexual organs are borne by similar but distinct individuals, it has been ascertained (Yamanouchi) that, as in Dictyota, meiosis takes place in the mother-cell of the tetraspores, so that the nuclei of these spores, as also those of the sexual plants to which they give rise, contain 20 chromosomes: and further, that the nuclei of the carpospores (diplospores) produced in the cystocarp as the result of fertilization, contain 40 chromosomes, as do also those of the asexual plant to which the carpospores give rise. Hence the sporophyte is represented by the cystocarp and the resulting tetrasporangiate plants: the gametophyte, by the sexual plants. Though it is the rule in the Red Algae that the tetrasporangia and the sexual organs are borne on distinct individuals, yet cases are known in which both kinds of reproductive organs are borne upon the same plant; and to those the above conclusions obviously cannot apply. They have yet to be investigated.
The second type of life-history has been traced in Nemalion. Here there is no tetrasporangiate form, consequently meiosis takes place at a different. stage in the life-history. It has been observed (Wolfe) that the nuclei of the sexual plant contain 8 chromosomes; those of the gonimoblast-filaments of the developing cystocarp contain 16, whilst those of the carpospores contain 8: hence meiosis takes place in the carposporangia. Here the plant is the gametophyte; the sporophyte is only represented by the cystocarp. The carpospores here are true spores (haplospores).
Among the Green Algae, Coleochaete is the only form that has been fully investigated (Allen). Here meiosis takes place in the germinating oospore: consequently the plant is the
