1911 Encyclopædia Britannica/Algae/Alternation of generations
In comparing algae with the great archegoniate series which has doubtless sprung from them, it is natural to inquire to what Alternation of generations. extent, if any, they present evidence of the existence of the marked alternation of generations which dominates the life-history of the higher plants. Turning first to the Rhodophyceae, both on account of the high place which they occupy among algae and also the remarkable uniformity in their reproductive processes, it is clear that, as is the case among Archegoniatae, the product of the sexual act never germinates directly into a plant which gives rise to the sexual organs. Even among Bangiaceae the carpospores arise from the fertilized cell by division, while in all other Rhodophyceae the oospore, as it may be called, gives rise to a filamentous structure, varying greatly in its dimensions, epiphytic, and to a large extent parasitic upon the egg-bearing parent plant, and in the end giving rise to carpospores in the terminal cells of certain branches. There is here obviously a certain parallelism with the case of Bryophyta, where the sporogonium arising from the oospore is epiphytic and partially parasitic upon the female plant, and always culminates in the production of spores. Not even Riccia, with its rudimentary sporogonium, has so simple a corresponding stage as Bangia, for, while there is some amount of sterile tissue in Riccia, in Bangia the oospore completely divides to form carpospores. Excluding Bangiaceae, however, from consideration, the Euflorideae present in the product of the development of the oospore like Byrophyta a structure partly sterile and partly fertile. There is, nevertheless, this important difference between the two cases. While the spore of Bryophyta on germination gives rise to the sexual plant, the carpospore of the alga may give rise on germination to a plant bearing a second sort of asexual cells, viz. the tetraspores, and the sexual plant may only be reached after a series of such plants have been successively generated. It is possible, however, that the tetraspore formation should be regarded as comparable with the prolific vegetative reproduction of Bryophyta, and in favour of this view there is the fact that the tetraspores originate on the thallus in a different way from carpospores with which the spores of Bryophyta are in the first place to be compared; moreover, in certain Nemalionales the production of tetraspores does not occur, and the difficulty referred to does not arise in such cases. Altogether it is difficult on morphological grounds to resist the conclusion that Florideae present the same fundamental phenomenon of alternation of generations as prevails in the higher plants. It is by means of the cytological evidence, however, that this problem will finally be solved. As is well known, the dividing nuclei of the cells of the sporophyte generation of the higher plants exhibit a double number of chromosomes, while the dividing nuclei of the cells of the gametophyte generation exhibit the single number. In a fern-plant, for example, which is a sporophyte, every karyokinesis divulges the double number, while in the prothallium, which is the gametophyte generation, the single number appears. The doubling process is provided by the act of fertilization, where an antherozoid with the single number of chromosomes fuses with an oosphere also with the single number to provide a fertilized egg with the double number. The reduction stage, on the other hand, is the first division of the mother-cell of the spore. From egg to spore-mother-cell is sporophyte; from spore-mother-cell to egg is gametophyte. And since this rule has been found to hold good for all the arghegoniate series and also for the flowering plants where, however, the gametophyte generation has become so extremely reduced as to be only with difficulty discerned, it is natural that when alternation of generation is stated to occur in any froup of Thallophyta it sshould be required that the cytological evidence should support the view. The genus Nemalion has been recently investigated by Wolfe with the object of examining the cytological evidence. He finds that eight chromosomes appear in karyokinesis in the ordinary thallus cells, but sixteen in the gonimoblast filaments derived from the fertilized carpogonium. Eight chromosomes appear again in the ultimate divisions which give rise to the carpospores. Upon the evidence it would seem therefore that so far as Nemalion is concerned an alternation occurs comparable with that existing in the lower Bryophyta where the sporophyte is relatively small, being attached to and to some extent parasitic upon the gametophyte. Nemalion is, however, one of those Florideae in which tetraspores do not occur. What is the case with those Florideae which have been described as trioecious? If the sporophyte generation is confined to the cystocarp, is the tetrasporiferous plant, as has been suggested, merely a potential gametophyte reproducing by a process analogous to the bud-formation of the Bryophyta? In answer to this question a recent writer, Yamanouchi, states in a preliminary communication that he has found that in Polysiphonia violacea the germinating carpospores exhibit forty chromosomes, and the germinating tetraspores twenty chromosomes. From this it would seem that in this plant reduction takes place in the tetraspore mother-cell, and that the tetrasporiferous plants are sporophytes which alternate with sexual plante. Novel as this result may seem, the tetraspores of Florideae become hereby comparable with the tetraspores of Dictyota, to which reference will be made hereafter. But it is clear that it becomes on this view increasingly difficult to explain the occasional occurrence of tetraspores on male, female and monoecious plants or the rôle of the carpospores in the life-cycle of Florideae. The results of future research on the cytology of the group will be awaited with interest.
Among Phaeophyceae it is well known that the oospore of Fucaceae germinates directly into the sexual plant, and there is thus only one generation. Moreover, it is known that the reduction in the number of chromosomes which occurs at the initiation of the gametophyte generation in Pteridophyta occurs in the culminating stage of Fucus, where the oogonium is separated from the stalk-cell, so that unless it be contended that the Fucus is really a sporophyte which does not produce spores, and that the gametophyte is represented merely by the oogonium and antheridium, there is no semblance of alternation of generation in this case. The only case among Phaeophyceae which has been considered to point to the existence of such a phenomenon is Cutleria. Here the asexual cells are borne upon the so-called Aglaozonia reptans and the sexual cells upon the plants known as Cutleria. The spores of the Aglaozonia form are known to give rise to sexual plants, and the oospore of Cutleria has been observed to grow into rudimentary Aglaozonia. Latterly, however, as the result of the cytological investigations of Mottier and Lloyd Williams, great advance has been made in our knowledge of the conditions existing in Dictyota. Mottier first observed that a reduction in the number takes place in the mother-cells of the tetraspore. It will be remembered that, as in most Florideae, the male, female and asexual plants are distinct in this genus. Mottier’s observation has been confirmed by Lloyd Williams, who has shown, moreover, that the single number occurs in germlings from the tetraspore, and also in the adult stages of all sexual plants, while the double number occurs in germlings from the oospore, and in adult stages of all asexual plants. It is probable, therefore, that we have here a sharp alternation of generations, both generations being, however, precisely similar to the eye up to point of reproduction. Among Chlorophyceae it is often the case that the oospore on germination divides up directly to form a brood of zoospores. In Coleochaete this seems to be preceded by the formation of a minute parenchymatous mass, in each cell of which a zoospore is produced. In Sphaeroplea it is only at this stage that zoospores are formed at all; but in most cases, such as Oedogonium, Ulothrix, Coleochaete, similar zoospores are produced again and again upon the thallus, and the product of the oospore may be regarded as merely a first brood of a series. It has been held by some, however, that the first brood corresponds to the sporophyte generation of the higher plants, and that the rest of the cycle is the gametophyte generation. Were the case of Sphaeroplea to stand alone, the phenomenon might perhaps be regarded as an alternation of generations, but still only comparable with the case of Bangia, and not the case of the Florideae. But it is difficult to apply such a term at all to those cases in which there intervene between the oospore and the next sexual stage a series of generations, the zoospores of which are all precisely similar.