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was proposed to unite them in one alliance, under the name of Mycetales, in the same manner as the late Dr. Lindley had united allied orders under alliances in his "Vegetable Kingdom;" but, beyond this, there was no predisposition towards the theory since propounded, and which, like all new theories, has collected a small but zealous circle of adherents. It will be necessary briefly to summarize this theory and the arguments by which it is supported and opposed, inasmuch as it is intimately connected with our subject.

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As recently as 1868, Professor Schwendener first propounded his views, and then briefly and vaguely, that all and every individual lichen was but an algal, which had collected about it a parasitic fungal growth, and that those peculiar bodies which, under the name of gonidia, were considered as special organs of lichens, were only imprisoned algae. In language which the Rev. J. M. Crombie † describes as "pictorial," this author gave the general conclusion at which he had arrived, as follows:"As the result of my researches, all these growths are not simple plants, not individuals in the usual sense of the term; they are rather colonies, which consist of hundreds and thousands of individuals, of which, however, only one acts as master, while the others, in perpetual captivity, provide nourishment for themselves and their master. This master is a fungus of the order Ascomycetes, a parasite which is accustomed to live upon the work of others; its slaves are green algae, which it has sought out, or indeed caught hold of, and forced into its service. It surrounds

whereas in the case of lichens the apothecia contain very little, if any, of those substances, but a large amount of the lichenoxanthines so characteristic of the class. Looking upon fungi from this chromatological point of view, they bear something like the same relation to lichens that the petals of a leafless parasitic plant would bear to the foliage of one of normal character-that is to say, they are, as it were, the coloured organs of reproduction of parasitic plants of a type closely approaching that of lichens, which, of course, is in very close, if not in absolute agreement with the conclusions drawn by botanists from entirely different data."

* Schwendener, "Untersuchungen über den Flechtenthallus."

+ Crombie (J. M.) "On the Lichen-Gonidia Question," in "Popular Science Review" for July, 1874.

them, as a spider does its prey, with a fibrous net of narrow meshes, which is gradually converted into an impenetrable covering. While, however, the spider sucks its prey and leaves it lying dead, the fungus incites the algae taken in its net to more rapid activity; nay, to more vigorous increase.' This hypothesis, ushered upon the world with all the prestige of the Professor's name, was not long in meeting with adherents, and the cardinal points insisted upon were-1st. That the generic relationship of the coloured "gonidia" to the colourless filaments which compose the lichen thallus, had only been assumed, and not proved; 2nd. That the membrane of the gonidia was chemically different from the membrane of the other tissues, inasmuch as the first had a reaction corresponding to that of algæ, whilst the second had that of fungi; 3rd. That the different forms and varieties of gonidia corresponded with parallel types of algae; 4th. That as the germination of the spore had not been followed further than the development of a hypothallus, it might be accounted for by the absence of the essential algal on which the new organism should become parasitic; 5th. That there is a striking correspondence between the development of the fruit in lichens and in some of the sporidiiferous fungi (Pyrenomycetes).

These five points have been combated incessantly by lichenologists, who would really be supposed by ordinary minds to be the most practically acquainted with the structure and development of these plants, in opposition to the theorists. It is a fact which should have some weight, that no lichenologist of repute has as yet accepted the theory. In 1873 Dr. E. Bornet* came to the aid of Schwendener, and almost exhausted the subject, but failed to convince either the practised lichenologist or mycologist. The two great points sought to be established are these, that what we call lichens are compound organisms, not simple, independent vegetable entities; and that this compound organism consists of unicellular algæ, with a fungus parasitic upon them. The coloured gonidia which are found in the

* Bornet, (E.), "Recherches sur les Gon dies des Lichens," in "Ann. des Sci. Nat." 1873, 5 sér. vol. xvii.

substance, or thallus of lichens, are the supposed alge; and the cellular structure which surrounds, encloses, and imprisons the gonidia is the parasitic fungus, which is parasitic on something infinitely smaller than itself, and which it entirely and absolutely isolates from all external influences.

Dr. Bornet believed himself to have established that every gonidium of a lichen may be referred to a species of algae, and that the connection between the hypha and gonidia is of such a nature as to exclude all possibility of the one organ being produced by the other. This he thinks is the only way in which it can be accounted for that the gonidia of diverse lichens should be almost identical.

Dr. Nylander, in referring to this hypothesis of an imprisoned algal,* writes: "The absurdity of such an hypothesis is evident from the very consideration that it cannot be the case that an organ (gonidia) should at the same time be a parasite on the body of which it exercises vital functions; for with equal propriety it might be contended that the liver or the spleen constitutes parasites of the mammiferæ. Parasite existence is autonomous, living upon a foreign body, of which nature prohibits it from being at the same time an organ. This is an elementary axiom of general physiology. But observation directly made teaches that the green matter originally arises within the primary chlorophyll- or phycochrom-bearing cellule, and consequently is not intruded from any external quarter, nor arises in any way from any parasitism of any kind. The celiule at first is observed to be empty, and then, by the aid of secretion, green matter is gradually produced in the cavity and assumes a definite form. It can, therefore, be very easily and evidently demonstrated that the origin of green matter in lichens is entirely the same as in other plants." On another occasion, and in another place, the same eminent lichenologist remarks,† as to the supposed algoid nature of gonidia-" that such an unnatural existence as they would thus pass, enclosed in a prison and

Nylander, "On the Algo-Lichen Hypothesis," &c., in "Grevillea," vol. ii. (1874), No. 22, p. 146.

In Regensburg "Flora," 1870, p. 92.

deprived of all autonomous liberty, is not at all consonant with the manner of existence of the other algae, and that it has no parallel in nature, for nothing physiologically analogous occurs anywhere else. Krempelhuber has argued that there are no conclusive reasons against the assumption that the lichen-gonidia may be self-developed organs of the lichen proper rather than algæ, and that these gonidia can continue to vegetate separately, and so be mistaken for unicellular algæ." In this Th. Fries seems substantially to concur. But there is one strong argument, or rather a repetition of an argument already cited, placed in a much stronger light, which is employed by Nylander in the following words:-"So far are what are called algæ, according to the turbid hypothesis of Schwendener, from constituting true algæ, that on the contrary it may be affirmed that they have a lichenose nature, whence it follows that these pseudo-algo are in a systematic arrangement to be referred rather to the lichens, and that the class of algae hitherto so vaguely limited should be circumscribed by new and truer limits.

As to another phase in this question, there are, as Krempelhuber remarks, species of lichens which in many countries do not fructify, and whose propagation can only be carried on by means of the soredia, and the hyphae of such could in themselves alone no more serve for propagation than the hyphæ from the pileus or stalk of an Agaric, while it is highly improbable that they could acquire this faculty by interposition of a foreign algal. On the other hand he argues: "It is much more conformable to nature that the gonidia, as self-developed organs of the lichens, should, like the spores, enable the hyphæ proceeding from them to propagate the individual.*

A case in point has been adduced† in which gonidia were produced by the hypha, and the genus Emericella,‡ which is allied to Husseia in the Trichogastres, shows a structure in the stem exactly resembling Palmella botryoides of Greville, and to what occurs in Synalyssa. Emericella, with one or two other

* Rev. J. M. Crombie, in "Popular Science Review," July, 1874.
+ Berkeley's "Introduction to Cryptogamic Botany," p. 373, fig. 780
‡ Berkeley's "Introduction," p. 341, fig. 76.

genera, must, however, be considered as connecting Trichogastres with lichens, and the question cannot be considered as satisfactorily decided till a series of experiments has been made on the germination of lichen spores and their relation to free algæ considered identical with gonidia. Mr. Thwaites was the first to point out* the relation of the gonidia in the different sections of lichens to different types of supposed algae. The question cannot be settled by mere à priori notions. It is, perhaps, worthy of remark that in Chionyphe Carteri the threads grow over the cysts exactly as the hypha of lichens is represented as growing over the gonidia.

Recently, Dr. Thwaites has communicated his views on one phase of this controversy,† which will serve to illustrate the question as seen from the mycological side. As is well known, this writer has had considerable experience in the study of the anatomy and physiology of all the lower cryptogamia, and any suggestion of his on such a subject will at least commend itselt to a patient consideration.

"According to our experience," he writes, "I think parasitic fungi invariably produce a sad effect upon the tissues they fix themselves upon or in. These tissues become pale in colour, and in every respect sickly in appearance. But who has ever seen the gonidia of lichens the worse for having the 'hypha' growing amongst them? These gonidia are always in the plumpest state, and with the freshest, healthiest colour possible. Cannot it enter into the heads of these most patient and excellent observers, that a cryptogamic plant may have two kinds of tissue growing side by side, without the necessity of one being parasitic upon the other, just as one of the higher plants may have half a dozen kinds of tissue making up its organization? The beautifully symmetrical growth of the same lichens has seemed to me a sufficient argument against one portion being parasitic upon another, but when we see all harmony and robust health, the idea that one portion is subsisting parasitically upon another appears to me to be a perfect absurdity."

"Annals and Magazine of Natural History," April, 1849.
"Gardener's Chronicle" for 1873, p. 1341.

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