Page:EB1911 - Volume 21.djvu/762

Rh is connected with its neighbours belonging to the same thread by two depressions or pits, one at each end. The common wall separating the pits of the two adjoining cells is pierced by strands of protoplasm. The whole structure, consisting of the two pits and the wall between is known as a genetic pit. Other pits, connecting cells not belonging to the same branch, are, however, formed at a later stage.

Many of the lower forms of Brown Seaweeds (Phoeophyceae) have a thallus consisting of simple or branched cell threads, as in the green and red forms. The lateral union of the branches to form a solid thallus is not, however, so common, nor is it carried to so high a pitch of elaboration as in the Rhodophyceae. In a few of the lower forms (Sphacelariaceae), and in the higher forms which possess a solid thallus, often of very large size, the plant-body is no longer formed entirely of branched cell-threads, but consists of what is called a true parenchymatous tissue, i.e. a solid mass of cells, formed by cell division in all directions of space. In the Laminariaceae this tissue is formed by cell division at what is called an intercalary growing point, i.e. a meristematic (cell-dividing) region occupying the whole of a certain transverse zone of the thallus, and cutting off new cells to add to the permanent tissue on both sides. In the Fucaceae, on the other hand, there is a single prismatic apical cell situated at the bottom of a groove at the growing apex of the thallus, which cuts off cells from its sides to add to the peripheral, and from its base to add to the central permanent cells. The whole of the tissue of the plant is formed by the division of this apical cell. In whatever way the tissues are originally formed, however, the main features of their differentiation are the same. According to a law which, as we have seen, applies also to the green and red forms, the superficial cells are packed with chromatophores and form the assimilating tissue (fig. 1, F). In these brown types with bodies of considerable thickness (Laminariaceae and Fucaceae), there is, however, a further differentiation of the internal tissues. The cells immediately subjacent to the superficial assimilating layer form a colourless, or nearly colourless, parenchymatous cortex, which acts as a food storage tissue (fig. 1, G), and surrounds a central medulla of elongated conducting cells. The latter are often swollen at the ends, so that the cross-wall separating two successive cells has a larger surface than if the cells were of uniform width along their entire length. Cells of this type are often called trumpet-hyphae (though they have no connexion with the hyphae of Fungi), and in some genera of Laminariaceae those at the periphery of the medulla simulate the sieve-tubes of the higher plants in a striking degree, even (like these latter) developing the peculiar substance callose on or in the perforated cross-walls or sieve-plates. A specialized conducting tissue of this kind, used mainly for transmitting organic substances, is always developed in plants where the region of assimilative activity is local in the plant-body, as it is in practically all the higher plants. This is the case in the Fucaceae, and in a very marked degree in the Laminariaceae in question, where the assimilative frond is borne at the end of an extremely long supporting and conducting stipe. A similar state of things exists in some of the more highly differentiated Red Seaweeds. The tissue developed to meet the demands for conduction in such cases always shows some of the characters described. It is known as leptom, each constituent cell being a leptoid (fig. 1, H). In addition to the cell types described, it is a very common occurrence in these bulky forms for rhizoid-like branches of the cells to grow out, mostly from the cells at the periphery of the medulla, and grow down between the cells, strengthening the whole tissue, as in the Rhodophyceae. This process may result in a considerable thickening of the thallus. In many Laminariaceae the thallus also grows regularly in thickness by division of its surface layer, adding to the subjacent permanent tissue and thus forming a secondary meristem.

The simpler Fungi, like the simpler Green Algae, consist of single cells or simple or branched cell-threads, but among the

higher kinds a massive body is often formed, particularly in connexion with the formation of spores, and this may exhibit considerable tissue-differentiation. A characteristic feature of the fungal vegetative plant-body (mycelium) is its formation from independent coenocytic tubes or cell-threads. These branch, and may be packed or interwoven to form a very solid structure; but each grows in length independently of the others and retains its own individuality, though its growth in those types with a definite external form is of course correlated with that of its neighbours and is subject to the laws governing the general form of the body. Such an independent coenocytic branch or cell-thread is called a hypha. Similar modes of growth occur among the Siphoneous Green Algae and also among the Red Seaweeds. A solid fungal body may usually be seen to consist of separate hyphae, but in some cases these are so bent and closely interwoven that an appearance like that of ordinary parenchymatous tissue is obtained in section, the structure being called pseudoparenchyma. By the formation of numerous cross-walls the resemblance to parenchyma is increased. The surface-layer of the body in the massive Fungi differs in character according to its function, which is not constant throughout the class, as in the Algae, because of the very various conditions of life to which different Fungi are exposed. In many forms its hyphae are particularly thick-walled, and may strikingly resemble

the epidermis of a vascular plant. This is especially the case in the lichens (symbiotic organisms composed of a fungal mycelium in association with algal cells), which are usually exposed to very severe fluctuations in external conditions. The formation of a massive body naturally involves the localization of the absorptive region, and the function of absorption (which in the simpler forms is carried out by the whole of the vegetative part of the mycelium penetrating a solid or immersed in a liquid substratum) is subserved by the outgrowth of the hyphae of the surface-layer of that region into rhizoids, which, like those of the Algae living on soil, resemble the root-hairs of the higher plants. The internal tissue of the body of the solid higher Fungi, particularly the elongated stalks (stipes) of the fructifications of the Agarics, consists of hyphae running in a longitudinal direction, which no doubt serve for the conduction of organic food substances, just as do the “trumpet-hyphae,” similar in appearance, though not in origin, of the higher Brown Seaweeds. (In one genus (Lactarius) “milk-tubes,” recalling the laticiferous tubes of many vascular plants, are found.) These elongated hyphae are frequently thick-walled, and in some cases form a central strand, which may serve to resist longitudinal pulling strains. This is particularly marked in certain lichens of shrubby habit. The internal tissues, either consisting of obvious hyphae or of pseudoparenchyma, may also serve as a storehouse of plastic food substances.

Looking back over the progress of form and tissue-differentiation in the Thallophyta, we find that, starting from the simplest unicellular forms with no external differentiation of the body, we can trace an increase in complexity of organization everywhere determined by the principles of the division of physiological labour and of the adaptation of the organism to the needs of its environment. In the first place there is a differentiation of fixing organs, which in forms living on a soft nutrient substratum penetrate it and become absorbing organs. Secondly, in the Algae, which build up their own food from inorganic materials, we have a differentiation of supporting axes from assimilating appendages, and as the body increases in size and becomes a solid mass of cells or interwoven threads, a corresponding differentiation of a superficial assimilative system from the deep-lying parts. In both Algae and Fungi the latter are primarily supporting and food-conducting, and in some bulky Brown Seaweeds, where assimilation is strongly localized, some of the deep cells are highly specialized for the latter function. In the higher forms a storage and a mechanically-strengthening system may also be developed, and in some aerial Fungi an external protective tissue. The “hyphal” mode of growth, i.e. the formation of the thallus, whatever its external form, by branched, continuous or septate, coenocytic tubes (Siphoneae and Fungi), or by simple or branched cell-threads (Red and many Green Algae), in both cases growing mainly or entirely at the apex of each branch, is almost universal in the group, the exceptions being met with almost entirely among the higher Brown Seaweeds, in which is found parenchyma produced by the segmentation of an apical cell of the whole shoot, or by cell division in some other type of meristem.

Bryophyta.—The Bryophyta [including the Liverworts (Hepaticae) and Mosses (Musci)], the first group of mainly terrestrial plants, exhibit considerably more advanced tissue differentiation, in response to the greater complexity in the conditions of life on land. In a general way this greater complexity may be said to consist (1) in the restriction of regular absorption of water to those parts of the plant-body embedded in the soil, (2) in the evaporation of water from the parts exposed to the air (transpiration). But these two principles do not find their full expression till we come, in the ascending series, to the Vascular Plants. In the Bryophytes water is still absorbed, not only from the soil but also largely from rain, dew, &c., through the general surface of the subaerial body (thallus), or in the more differentiated forms through the leaves. The lowest Hepaticae have an extremely simple vegetative structure, little more advanced than that found in some of the higher Green Algae and very much simpler than in the large Red and Brown Seaweeds, The plant-body (thallus) is always small and normally lives in very damp air, so that the demands of terrestrial life are at a minimum. It always consists of true parenchyma, and is entirely formed by the cutting off of segments from an apical cell.