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.
The difficulty of tracing the relationships of algae is largely due to the inadequacy of our knowledge of the conditions under which they pass through the critical stages of their life-cycle. Of the thousands of species which have been distinguished, relatively few have been traced from spore to spore, as the flowering plants have been observed Poly-morphism.from seed to seed. The aquatic habit of most of the species and the minute size of many of them are difficulties which do not exist in the case of most seed-plants. From the analogy of the higher plants observers have justly argued that when they have seen and marked the characters of the reproductive organs they have found the plant at the stage when it exhibits its most noteworthy features, and they have named and classified the species in accordance with these observations. While even in such cases it is obvious that interesting stages in the life of the plant may escape notice altogether, in the cases of those plants the reproduction of which is unknown, and which have been named and placed on the analogy of the vegetative parts alone, there is considerable danger that a plant may be named as a distinct species which is only a stage in the life of another distinct and perhaps already known species. To take an example, Lemanea and Batrachospermum are Florideae which bear densely-whorled branches, but which, on the germination of the carpospore, give rise to a laxly-filamentous, somewhat irregularly-branched plant, from which the ordinary sexual plants arise at a later stage. This filamentous structure has been attributed to the genus Chantransia, which it greatly resembles, especially when, as is said to be the case in Batrachospermum, it bears similar monospores. The true Chantransia, however, bears its own sexual organs as well as monospores. To the specific identity of Haplospora globosa and Scaphospora speciosa, and of Cutleria multifida and Aglaozonia reptans, reference has already been made. Again, many Green Algae—some unicellular, like Sphaerella and Chlamydomonas; some colonial forms, like Volvox and Hormotila; some even filamentous forms, like Ulothrix and Stigeoclonium—are known to pass into a condition resembling that of a Palmella, and might escape identification on this account.
It is, on the other hand, a danger in the opposite sense to conclude that all Chantransia species are stages in the life-cycle of other plants, and, similarly, that all irregular colonial forms, like Palmella represent phases in the life of other Green Algae. Long ago Kützing went so far as to express the belief that the lower algae were all capable of transformations into higher forms, even into moss-protonemata. Later writers have also thought that in all four groups of algae transformations of a most far-reaching character occur. Thus Borzi finds that Protoderma viride passes through a series of changes so varied that at different times it presents the characters of twelve different genera. Chodat does not find so general a polymorphism, but nevertheless holds that Raphidium passes through stages represented by Protococcus, Characium, Dactylococcus and Sciadium. Klebs has, however, recently canvassed the conclusions of both these investigators; and as the result of his own observations declares that algae, so far from being as polymorphic as they have been described, vary only within relatively narrow limits, and present on the whole as great fixity as the higher plants. It certainly supports his view to discover, on subjecting to a careful investigation Botrydium granulatum, a siphonaceous alga whose varied forms had been described by J. Rostafinski and M. Woronin, that these authors had included in the life-cycle stages of a second alga described previously by Kütxing, and now described afresh by Klebs as Protosiphon botryoides. In Botrydium the chromatophores are small, without pyrenoids, and oil-drops are present; in Protosiphon the chromatophores form a net-work with pyrenoids, and the contents include starch. Klebs insists that the only solution of such problems is the subjection of the algae in question to a rigorous method of pure culture. It is interesting to learn that G. Senn, pursuing the methods described by Klebs, has confirmed Chodat’s observation of the passage of Raphidium into a Dactylococcus-stage, although he was unable to observe further metamorphosis. He has also seen Pleurococcus viridis dividing so as to form a filament, but has not succeeded in seeing the formation of zoospores as described by Chodat. While, therefore, there is much evidence of a negative character against the existence of an extensive polymorphism among algae, some amount of metamorphosis is known to occur. But until the conditions under which a particular transformation takes place have been ascertained and described, so that the observation may be repeated by other investigators, scant credence is likely to be given to the more extreme polymorphistic views.
In comparison with the higher plants, algae exhibit so much simplicity of structure, while the conditions under which they grow are so much more readily controlled, that they have frequently been the subject of physiological investigation with a view chiefly to the application of the results to the study of the higher plants. (See Physiology.Plants: Physiology of.) In the literature of vegetable physiology there has thus accumulated a great body of facts relating not only to the phenomena of reproduction, but also to the nutrition of algae. With reference to their chemical physiology, the gelatinization of the cell-wall, which is so marked a feature, is doubtless attributable to the occurrence along with cellulose of pectic compounds. There is, however, considerable variation in the nature of the membrane in different species; thus the cell-wall of Oedogonium, treated with sulphuric acid and iodine, turns a bright blue, while the colour is very faint in the case of Spirogyra, the wall of which is said to consist for the most part of pectose. While starch occurs commonly as a cell-content in the majority of the Green Algae no trace of it occurs in Vaucheria and some of
