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Schwendener[345] was the first who, with the improved microscope, made a systematic study of the minute structure of lichens. He examined typical species in genera of widely different groups and described their anatomy in detail. The most variable and perhaps the most important of the tissues of lichens is the cortex, which is most fully developed in the larger thalli, and as the same type of cortical structures recurs in lichens widely different in affinity as well as in form, it seems well to group together here the ascertained facts about these covering layers.

a. Types of Cortical Structure. Zukal[346], and more recently Hue[347], have made independent studies in the comparative morphology of the thallus and have given particular attention to the different varieties of cortex. They each find that the variations come under a definite series of types. Zukal recognized five of these:

1. Pseudoparenchymatous (plectenchyma): by frequent septation of regularly arranged hyphae and by coalescence a kind of continuous cell-structure is formed.

2. Palisade cells: the outer elongate ends of the hyphae lie close together in a direction at right angles to the surface of the thallus and form a coherent row of parallel cells.

3. Fibrous: the cortical hyphae lie in strands of fine filaments parallel with the surface of the thallus.

4. Intricate: hyphae confusedly interwoven and becoming dark in colour form the lower cortex of some foliose lichens.

These four types, Zukal finds, are practically without interstices in the tissue and form a perfect protection against excessive transpiration. He adds yet another form:

5. A cortex formed of hyphae with dark-coloured swollen cells, which is not a protection against transpiration. It occurs among lower crustaceous forms.

Hue has summed up the different varieties under four types, but as he has omitted the “fibrous” cortex, we arrive again at five different kinds of cortical formation, though they do not exactly correspond to those of Zukal. A definite name is given to each type:

1. Intricate: an intricate dense layer of gelatinous-walled hyphae, branching in all directions, but not coalescent (Fig. 44). This rather unusual type of cortex occurs in Sphaerophorus and Stereocaulon, both of which have an upright rigid thallus (fruticose).

Fig. 44. Sphaerophorus coralloides Pers. Transverse section of cortex and gonidial layer near the growing point of a frond × 600.

Fig. 45. Roccella fuciformis DC. Transverse section of cortex near the growing point of a frond × 600.

2. Fastigiate: the hyphae bend outwards or upwards to form the cortex. A primary filament can be distinguished with abundant branches, all tending in the same direction; anastomosis may take place between the hyphae. The end branches are densely packed, though there are occasional interstices (Fig. 45). Such a cortex occurs in Thamnolia; in several genera of Roccellaceae—Roccellographa, Roccellina, Reinkella, Pentagenella, Combea, Schizopelte and Roccella—and also in the crustaceous genus Dirina. The fastigiate cortex corresponds with Zukal’s palisade cells.

3. Decomposed: in this, the most frequent type of cortex, the hyphae that travel up from the gonidial layer become irregularly branched and frequently septate. The cell-walls of the terminal branches become swollen into a gelatinous mass, the transformation being brought about by a change in the molecular constituents of the cell-walls which permits the imbibition and storage of water. The tissue, owing to the enormous increase of the wall, is so closely pressed together that the individual hyphae become indistinct; the cell-lumen finally disappears altogether, or, at most, is only to be detected in section as a narrow disconnected dark streak. The decomposed cortex is characteristic of many lichens, crustaceous (Fig. 46) and squamulose, as well as of such highly developed genera as Usnea, Letharia, Ramalina, Cetraria, Evernia and certain Parmeliae.

Fig. 46. Lecanora glaucoma var. corrugata Nyl. Vertical section of cortex × 500 (after Hue).

Zukal took no note of the decomposed cortex but the omission is intentional and is due to his regarding the structure of the youngest stages of the thallus near the growing point as the most typical and as giving the best indication as to the true arrangement of hyphae in the cortex. He thus describes palisade tissue as the characteristic cortex of Evernia, since the formation near the growing point of the fronds is somewhat palisade-like and he finds fibrous cortex at the tips of Usnea filaments. In both these instances Hue has described the cortex as decomposed because he takes account only of the fully formed thallus in which the tissues have reached a permanent condition.

Fig. 47. Peltigera canina DC. Vertical section of cortex and gonidial zone × 600.

4. Plectenchymatous: the last of Hue’s types corresponds with the first described by Zukal. It is the result of the lateral coherence and frequent septation of the hyphae into short almost square or rounded cells (Fig. 47). The simplest type of such a cortex can be studied in Leptogium, a genus of gelatinous lichens in which the tips of the hyphae are cut off at the surface by one or more septa. The resulting cells are wider than the hyphae and they cohere together to form, in some species, disconnected patches of cells; in others, a continuous cortical covering one or more cells thick, while in the margin of the apothecium they form a deep cellular layer. The cellular type of cortex is found also, as already stated, in some crustaceous Pertusariae, and in a few squamulose genera or species. It forms the uppermost layer of the Peltigera thallus and both cortices of many of the larger foliose lichens such as Sticta, Parmelia, etc.

5. The “fibrous” cortex must be added to this series, as was pointed out by Heber Howe[348] who gave the less appropriate designation of “simple” to the type. It consists of long rather sparingly branched slender hyphae that grow in a direction parallel with the surface of the thallus (Fig. 48). It is characteristic of several fruticose and foliose lichens with more or less upright growth, such as we find in several of the Physciae, and in the allied genus Teloschistes, in Alectoria, several genera of Roccellaceae, in Usnea longissima and in Parmelia pubescens, etc. Zukal would have included all the Usneae as the tips are fibrous.

Fig. 48. Physcia ciliaris DC. Vertical section of thallus. a, cortex; b, gonidial zone; c, medulla. × 100.

More than one type of cortex, as already stated, may appear in a genus: a striking instance of variability occurs in Solorina where, as Hue[349] has pointed out, the cortex of S. octospora is fastigiate, that of all the other species being plectenchymatous. Cortical development is a specific rather than a generic characteristic.

b. Origin of Variation in Cortical Structure. The immediate causes making for differentiation in cortical development are: the prevailing direction of growth of the hyphae as they rise from the gonidial zone; the amount of branching and the crowding of the filaments; the frequency of septation; and the thickening or degeneration of the cell-walls which may become almost or entirely mucilaginous. In the plectenchymatous cortex, the walls may remain quite thin and the cells small as in Xanthoria parietina, or the walls may be much thickened as in both cortices of Sticta. As a result of stretching the cell may increase enormously in size: in some instances where the internal hyphae are about 3 µ to 4 µ in width, the cortical cells formed from these hyphae may have a cell cavity 15 µ to 16 µ in diameter.

c. Loss and Renewal of Cortex. Very frequently the cortex is covered over by a layer of homogeneous mucilage which forms an outer cuticle. It arises from the continual degeneration of the outer cell-walls and it is liable to friction and removal by atmospheric agency as was first described by Schwendener[350] in the weather-beaten cortex of Umbilicaria pustulata. He had noted the irregular jagged outline of the cross section of the thallus, and he then suggested, as the probable reason, the decay of the outer rind with the constant renewal of it by the hyphae from the underlying gonidial zone, though he was unable definitely to prove his theory. The peeling of the dead outer layer (with its replacement by new tissue) has however been observed many times since his day. It has been described by Darbishire[351] in Pertusaria: in that genus there is at first a primary cortex formed of hyphae that grow in a radial direction, parallel to the surface of the thallus. The walls of these hyphae become gradually more and more mucilaginous till the cells are obliterated. Meanwhile short-celled filaments grow up in serried ranks from the gonidial layer and finally push off the dead “fibrous” cortex. The new tissue takes on a plectenchymatous character, and the outer cells in time become decomposed and provide a mucilaginous cuticle which in turn is also subject to wasting.

The same process of peeling was noted by Rosendahl[352] in some species of brown Parmeliae, where the dead tissues were thrown off in shreds, though only in isolated patches. But whether in patches or as a continuous sheath, there is constant degeneration, with continual renewal of the dead material from the internal tissues.

The cortex is the most highly developed of all the lichen structures and is of immense importance to the plant as may be judged from the various adaptations to different needs[353]. The cortical cell-walls are frequently impregnated with some dark-coloured substance which, in exposed situations, must counteract the influence of too direct sunlight and be of service in sheltering the gonidia. Lichen acids—sometimes very brightly coloured—and oxalic acid are deposited in the cortical tissues in great abundance and aid in retaining moisture; but the two chief functions to which the cortex is specially adapted are the checking of transpiration and the strengthening of the thallus against external strains.

d. Cortical Hairs or Trichomes. Though somewhat rare, cortical hairs are present on the upper surface of several foliose lichens. They take rise, in all the instances noted, as a prolongation of one of the cell-rows forming a plectenchymatous cortex.

In Peltidea (Peltigera) aphthosa they are especially evident near the growing edges of the thallus; and they take part in the development of the superficial cephalodia[354] which are a constant feature of the lichen. They tend to disappear with age and leave the central older parts of the thallus smooth and shining. In several other species of Peltigera (P. canina, etc.) they are present and persist during the life of the cortex. In these lichens the cells of the cortical tissue are thin-walled, all except the outer layer, the membranes of which are much thicker. The hairs rising from them are also thick-walled and septate. Generally they branch in all directions and anastomose with neighbouring hairs so that a confused felted tangle is formed; they vary in size but are, as a rule, about double the width of the medullary hyphae as are the cortical cells from which they rise. They disappear from the thallus, frequently in patches, probably by weathering, but over large surfaces, and especially where any inequality affords a shelter, they persist as a soft down.

Hairs are also present on the upper surface of some Parmeliae. Rosendahl[355] has described and figured them in P. glabra and P. verruculifera—short pointed unbranched hyphae, two or more septate and with thickened walls. They are most easily seen near the edge of the thallus, though they persist more or less over the surface; they also grow on the margins of the apothecia. In P. verruculifera they arise from the soredia; in P. glabra a few isolated hairs are present on the under surface.

In Nephromium tomentosum there is a scanty formation of hairs on the upper surface. They are abundant on the lower surface, and function as attaching organs. A thick tomentum of hairs is similarly present on the lower surface of many of the Stictaceae either as an almost unbroken covering or in scattered patches. In several species of Leptogium they grow out from the lower cortical cells and attach the thin horizontal fronds; and very occasionally they are present in Collema.