Page:EB1911 - Volume 21.djvu/764

ANATOMY] undulata the central hydrom cylinder of the aerial stem is a loose tissue, its interstices being filled up with thin-walled, starchy parenchyma. In Dawsonia superba, a large New Zealand moss, the hydroids of the central cylinder of the aerial stem are mixed with thick-walled stereids forming a hydrom-stereom strand somewhat like that of the rhizome in other Polytrichaceae.

The central hydrom strand in the seta of the sporogonium of most mosses has already been alluded to. Besides this there is usually a living conducting tissue, sometimes differentiated as leptom, forming a mantle round the hydrom, and bounded externally by a more or less well-differentiated endodermis, abutting on an irregularly cylindrical lacuna; the latter separates the central conducting cylinder from the cortex of the seta, which, like the cortex of the gametophyte stem, is usually differentiated into an outer thick-walled stereom and an inner starchy parenchyma. Frequently, also, a considerable differentiation of vegetative tissue occurs in the wall of the spore-capsule itself, and in some of the higher forms a special assimilating and transpiring organ situated just below the capsule at the top of the seta, with a richly lacunar chlorophyllous parenchyma and stomata like those of the wall of the capsule in the Anthocerotean liverworts. Thus the histological differentiation of the sporogonium of the higher mosses is one of considerable complexity; but there is here even less reason to suppose that these tissues have any homology (phylogenetic community of origin) with the similar ones met with in the higher plants.

The features of histological structure seen in the Bryophytic series are such as we should expect to be developed in response to the exigencies of increasing adaptation to terrestrial life on soil, and of increasing size of the plant-body. In the liverworts we find fixation of the thallus by water-absorbing rhizoids; in certain forms with a localized region of water-absorption the development of a primitive hydrom or water-conducting system; and in others with rather a massive type of thallus the differentiation of a special assimilative and transpiring system. In the more highly developed series, the mosses, this last division of labour takes the form of the differentiation of special assimilative organs, the leaves, commonly with a midrib containing elongated cells for the ready removal of the products of assimilation; and in the typical forms with a localized absorptive region, a well-developed hydrom in the axis of the plant, as well as similar hydrom strands in the leaf-midribs, are constantly met with. In higher forms the conducting strands of the leaves are continued downwards into the stem, and eventually come into connexion with the central hydrom cylinder, forming a complete cylindrical investment apparently distinct from the latter, and exhibiting a differentiation into hydrom, leptom and amylom which almost completely parallels that found among the true vascular plants. Similar differentiation, differing in some details, takes place independently in the other generation, the sporogonium. The stereom of the moss is found mainly in the outer cortex of the stem and in the midrib of the leaf.

Vascular Plants.—In the Vascular Plants (Pteridophytes, i.e. ferns, horse-tails, club mosses, &c., and Phanerogams or Flowering Plants) the main plant-body, that which we speak of in ordinary language as “the plant,” is called the sporophyte because it bears the asexual reproductive cells or spores. The gametophyte, which bears the sexual organs, is either a free-living thallus corresponding in degree of differentiation with the lower liverworts, or it is a mass of cells which always remains enclosed in a spore and is parasitic upon the sporophyte.

The body of the sporophyte in the great majority of the vascular plants shows a considerable increase in complexity over that found in the gametophyte of Bryophytes. The principal new feature in the external conformation of the body is the acquirement of “true” roots, the nearest approach to which in the lower forms we saw in the “rhizome” of Polytrichaceae. The primary root is a downward prolongation of the primary axis of the plant. From this, as well as from various parts of the shoot system, other roots may originate. The root differs from the shoot in the characters of its surface tissues, in the absence of the green assimilative pigment chlorophyll, in the arrangement of its vascular system and in the mode of growth at the apex, all features which are in direct relation to its normally subterranean life and its fixative and absorptive

functions. Within the limits of the sporophyte generation the Pteridophytes and Phanerogams also differ from the Bryophytes in possessing special assimilative and transpiring organs, the leaves, though these organs are developed, as we have seen, in the gametophyte of many liverworts and of all the mosses. The leaves, again, have special histological features adapted to the performance of their special functions.

Alike in root, stem and leaf, we can trace a three-fold division of tissue systems, a division of which there are indications among the lower plants, and which is the expression of the fundamental

conditions of the evolution of a bulky differentiated plant-body. From the primitive uniform mass of undifferentiated assimilating cells, which we may conceive of as the starting-point of differentiation, though such an undifferentiated body is only actually realized in the thallus of the lower Algae, there is, (1) on the one hand, a specialization of a surface layer regulating the immediate relations of the plant with its surroundings. In the typically submerged Algae and in submerged plants of every group this is the absorptive and the main assimilative layer, and may also by the production of mucilage be of use in the protection of the body in various ways. In the terrestrial plants it differs in the subterranean and subaerial parts, being in the former pre-eminently absorptive, and in the latter protective—provision at the same time being made for the gaseous interchange of oxygen and carbon dioxide necessary for respiration and feeding. This surface layer in the typically subaerial “shoot” of the sporophyte in Pteridophytes and Phanerogams is known as the epidermis, though the name is restricted by some writers, on account of developmental differences, to the surface layer of the shoot of Angiosperms, and by others extended to the surface layer of the whole plant in both these groups. On the other hand, we have (2) an internal differentiation of conducting tissue, the main features of which as seen in the gametophyte of Bryophytes have already been fully described. In the Vascular Plants this tissue is collectively known as the vascular system. The remaining tissue of the plant-body, a tissue that we must regard phylogenetically as the remnant of the undifferentiated tissue of the primitive thallus, but which often undergoes further differentiation of its own, the better to fulfil its characteristically vital functions for the whole plant, is known, from its peripheral position in relation to the primitively central conducting tissue, as (3) the cortex. Besides absorption, assimilation, conduction and protection there is another very important function for which provision has to be made in any plant-body of considerable size, especially when raised into the air, that of support. Special tissues (stereom) may be developed for this purpose in the cortex, or in immediate connexion with the conducting system, according to the varying needs of the particular type of plant-body. The important function of aeration, by which the inner living tissues of the bulky plant-body obtain the oxygen necessary for their respiration, is secured by the development of an extensive system of intercellular spaces communicating with the external air.

In relation to its characteristic function of protection, the epidermis, which, as above defined, consists of a single layer of cells has typically thickened and cuticularized outer walls. These serve not only to protect the plant against slight mechanical injury from without, and against the entry of smaller

parasites, such as fungi and bacteria, but also and especially to prevent the evaporation of water from within.

At intervals it is interrupted by pores (stomata) leading from the air outside to the system of inter cellular spaces below. Each stoma is surrounded by a pair of peculiarly modified epidermal cells called guard-cells (fig. 1, T), which open and close the pore according to the need for transpiration. The structure

of the stomata of the sporophyte of vascular plants is fundamentally the same as that of the stomata on the sporogonium of the true mosses and of the liverwort Anthoceros. Stomata are often situated at the bottom of pits in the surface of the leaf. This arrangement is a method of checking transpiration by creating a still atmosphere above the pore of the stoma, so that water vapour collects in it and diminishes the further outflow of vapour. This type of structure, which is extremely various in its details, is found especially, as we should expect, in plants which have to economize their water