LICHENS

This article appears in Volume V16, Page 578 of the Encyclopedia Britannica.

Encyclopedia Britannica - Main :: LEO-LOB

LICHENS , in botany, compound or dual organisms each consisting of an association of a higher fungus, with a usually unicellular, sometimes filamentous, alga. The fungal part of the organism nearly always consists of a number of the Discomycetes or Pyrenomycetes, while the algal portion is a member of the Schizophyceae (Cyanophyceae or Blue-green Algae) or of the Green Algae; only in a very few cases is the fungus a member of the Basidiomycetes. The

special

SPECIAL

fungi which take part in the association are, with rare exceptions, not found growing separately, while the algal forms are constantly found free. The reproductive organs of the lichen are of a typically fungal character, i.e. are apothecia or perithecia (see FUNGI) and spermogonia. The algal cells are never known to form spores while part of the lichen-thallus, but they may do so when separated from it and growing free: The fungus thus clearly takes the upper hand in the association.
Owing to their peculiar dual nature, lichens are able to livein situations where neither the alga nor fungus could exist alone. The enclosed alga is protected by the threads (hyphae) of the fungus, and supplied with water and salts and, possibly, organic nitrogenous substances; in its turn the alga by means of its green or blue-green colouring matter and the sun's energy manufactures carbohydrates which are used in part by the fungus. An association of two organisms to their mutual advantage is known as symbiosis, and the lichen in botanical language is described as a symbiotic union of an alga and a fungus. This form of relationship is now known in other groups of plants (see BACTERIOLOGY and FUNGI), but it was first discovered in the lichens. The lichens are characterized by their excessively slow growth and their

great

GREAT

length of life.
Until comparatively

recent

RECENT

times the lichens were considered as a group of simple organisms on a level with algae and fungi. The green (or blue-green) cells were termed gonidia by Wallroth, who looked upon them as asexual reproductive cells, but when it was later realized that they were not reproductive elements they were considered as mere outgrowths of the hyphae of the thallus which had developed chlorophyll. In 1865 De Bary suggested the possibility that such lichens as Collema, Ephebe, &c., arose as a result of the attack of parasitic Ascomycetes upon the algae, Nostoc, Chroococcus, &c. In 1867 the observations of Famintzin and Baranetzky showed that the gonidia, in certain cases,,,were able to live outside the lichen-thallus, and in the case of Physcia, Evernia and Cladonia were able to form zoospores. Baranetzky therefore concluded that a certain number, if not all of the so-called algae were nothing more than free living lichen-gonidia. In 1869 Schwendener put forward the really

illuminating

ILLUMINATING

viewexactly opposite to that of Baranetzkythat the gonidia in all cases were algae which had been attacked by parasitic fungi. Although Schwendener supported this view of the " dual " nature of lichens by very strong evidence and identified the more common lichen-gonidia with known free-living algae, yet the theory was received with a

storm

of opposition by nearly all lichenologists. These workers were unable to consider with equanimity the loss of the autonomy of their group and its reduction to the level of a

special

SPECIAL

division of the fungi. The, observations of Schwendener, however, received ample support from Bornet's (1873) examination of 6o genera. He investigated the exact relation of fungus and alga and showed that the same alga is able to combine with a number of different fungi to form lichens; thus Chroolepns umbrinus is found as the gonidia of 13 different lichen genera.
The view of the dual nature of lichens had hitherto been based on

analysis

ANALYSIS (Gr. avci and Meta, to break up into parts)

; the final proof of this view was now supplied by the actual synthesis of a lichen from fungal and algal constituents. Rees in 1871 produced the sterile thallus of a Collema from its constituents; later Stahl did the same for three species. Later

Bonnier

BONNIER, ANGE ELISABETH LOUIS ANTOINE (1749-1799)

(1886) succeeded in producing fertile thalli by sowing lichen spores and the appropriate algae upon sterile glass plates or portions of bark, and growing them in sterilized air (fig. i). Moller also in 1887 succeeded in growing small lichen-thalli without their algal constituent (gonidia) on nutritive solutions; in the case of Calicium pyenidia were actually produced under these conditions.
The thallus or

body

BODY

of the lichen is of very different form in different genera. In the simplest filamentous lichens (e.g. Ephebe pubescens) the form of thallus is the form of the filamentous alga which is merely surrounded by the fungal hyphae (fig. 2). The next simplest forms are gelatinous lichens (e.g. Collemaceae); in these the algae are Chroococcaceae and Nostocaceae, and the fungus makes its way into the gelatinous membranes of the algal cells and ramifies there (fig. 3). We can distinguish this class of forms as lichens with a homoioinerous thallus, i.e. One in which the alga and fungus are equally distributed. The

majority

MAJORITY (Fr. majorite; Med. Lat. majoritas; Lat. major, greater)

of the lichens, however, possess a stratified thallus in which the gonidia are found as a definite layer or layers embedded in a pseudo-parenchymatous mass of fungal hyphae, i.e. they are heteromerous (

figs

FIGS

. 8 and 9). Obviously these two conditions may merge

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