scoparius, Potentilla procumbens, Galium hercynicum (＝G. saxatile),
Gnaphalium sylvaticum, Digitalis purpurea. Other plants occur
indifferently both on calcareous and on non-calcareous soils.
It is sometimes said that lime acts as a poison on some plants and not on others, and sometimes that it is the physiological dryness of calcareous soils that is the important factor. In relation to the latter theory, it is pointed out that some markedly calcicole species occur on sand dunes; but this may be due to the lime which is frequently present in dune sand as well as to the physical dryness of the soil. Further, no theory of calciolous and calcifugous plants can be regarded as satisfactory which fails to account for the fact that both kinds of plants occur among aquatic as well as among terrestrial plants. Schimper (1903: 102) thinks that in the case of aquatic plants, the difference must depend on the amount of lime in the water, for the physical nature of the substratum is the same in each case. Again, acidic humus does not form in calcareous soils; and hence one does not expect to find plants characteristic of acidic peat or humus on calcareous soils. Some such species are Blechnum boreale, Aira flexuosa, Calluna vulgaris, Vaccinium, Myrtillus, Rubus, Chamaemorus, Empetrum nigrum, Drosera spp. Some, at least, of these species possess mycorhiza in their roots, and are perhaps unable to live in soils where such organisms are absent.
In England, the number of calcicole species is greater than the number of silicolous species. It would therefore be curious if it were proved that lime acts on plants as a poison. It is said that some plants may be calcicoles in one geographical district and not in another. However, until more is known of the exact chemical composition of natural—as contrasted with agricultural—soils, and until more is known of the physiological effects of lime, it is impossible to decide the vexed question of the relation of lime-loving and lime-shunning plants to the presence or absence of calcium carbonate in the soil. From such points of view as this, it is indeed true, as Warming has recently stated, “that ecology is only in its infancy.” (C. E. M.)
Cytology of Plants
The elementary unit of plant structure, as of animal structure, is the cell. Within it or its modifications all the vital phenomena of which living organisms are capable have their origin. Upon our knowledge of its minute structure or cytology, combined with a study of its physiological activities, depends the ultimate solution of all the important problems of nutrition and growth, reception and conduction of stimuli, heredity, variation, sex and reproduction.
The Cell Theory.—For a general and historical account of the cell theory see Cytology. It is sufficient to note here that cells were first of all discovered in various vegetable tissues by Robert Hooke in 1665 (Micrographia); Malpighi and Grew (1674-1682) gave the first clear indications of the importance of cells in the building up of plant tissues, but it was not until the beginning of the 19th century that any insight into the real nature of the cell and its functions was obtained. Hugo von Mohl (1846) was the first to recognize that the essential vital constituent of the plant cell is the slimy mass—protoplasm—inside it, and not the cell wall as was formerly supposed. The nucleus was definitely recognized in the plant cell by Robert Brown in 1831, but its presence had been previously indicated by various observers and it had been seen by Fontana in some animal cells as early as 1781. The cell theory so far as it relates to plants was established by Schleiden in 1838. He showed that all the organs of plants are built up of cells, that the plant embryo originates from a single cell, and that the physiological activities of the plant are dependent upon the individual activities of these vital units. This conception of the plant as an aggregate or colony of independent vital units governing the nutrition, growth and reproduction of the whole cannot, however, be maintained. It is true that in the unicellular plants all the vital activities are performed by a single cell, but in the multicellular plants there is a more or less highly developed differentiation of physiological activity giving rise to different tissues or groups of cells, each with a special function. The cell in such a division of labour cannot therefore be regarded as an independent unit. It is an integral part of an individual organization and as such the exercise of its functions must be governed by the organism as a whole.
General Structure and Differentiation of the Vegetable Cell.—The simplest cell forms are found in embryonic tissues, in reproductive cells and in the parenchymatous cells, found in various parts of the plant. The epidermal, conducting and strengthening tissues show on the other hand considerable modifications both in form and structure.
The protoplasm of a living cell consists of a semifluid granular substance, called the cytoplasm, one or more nuclei, and sometimes centrosomes and plastids. Cells from different parts of a plant differ very much in their cell-contents. Young cells are full of cytoplasm, old cells generally contain a large vacuole or vacuoles, containing cell-sap, and with only a thin, almost invisible layer of cytoplasm on their walls. Chlorophyll grains, chromatophores, starch-grains and oil-globules, all of which can be distinguished either by their appearance or by chemical reagents, may also be present. Very little is known of the finer structure of the cytoplasm of a vegetable cell. It is sometimes differentiated into a clearer outer layer, of hyaloplasm, commonly called the ectoplasm, and an inner granular endoplasm. In some cases it shows, when submitted to a careful examination under the highest powers of the microscope, and especially when treated with reagents of various kinds, traces of a more or less definite structure in the form of a meshwork consisting of a clear homogeneous substance containing numerous minute bodies known as microsomes, the spaces being filled by a more fluid ground-substance. This structure, which is visible both in living cells and in cells treated by reagents, has been interpreted by many observers as a network of threads embedded in a homogeneous ground-substance. Bütschli, on the other hand, interprets it as a finely vacuolated foam-structure or emulsion, comparable to that which is observed when small drops of a mixture of finely powdered potash and oil are placed in water, the vacuoles or alveoli being spaces filled with liquid, the more solid portion representing the mesh-work in which the microsomes are placed. Evidence is not wanting, however, that the cytoplasm must be regarded as, fundamentally, a semifluid, homogeneous substance in which by its own activity, granules, vacuoles, fibrils, &c., can be formed as secondary structures. The cytoplasm is largely concerned in the formation of spindle fibres and centrosomes, and such structures as the cell membrane, cilia, or fiagella, the coenocentrum, nematoplasts or vibrioids and physodes are also products of its activity.
Protoplasmic Movements.—In the cells of many plants the cytoplasm frequently exhibits movements of circulation or rotation. The cells of the staminal hairs of Tradescantia virginica contain a large sap-cavity across which run, in all directions, numerous protoplasmic threads or bridges. In these, under favourable conditions, streaming movements of the cytoplasm in various directions can be observed. In other forms such as Elodea, Nitella, Chara, &c., where the cytoplasm is mainly restricted to the periphery of the sap vacuole and lining the cell wall, the streaming movement is exhibited in one direction only. In some cases both the nucleus and the chromatophores may be carried along in the rotating stream, but in others, such as Nitella, the chloroplasts may remain motionless in a non-motile layer of the cytoplasm in direct contact with the cell wall.[1]
Desmids, Diatoms and Oscillaria show creeping movements probably due to the secretion of slime by the cells; the swarm-spores and plasmodium of the Myxomycetes exhibit amoeboid movements; and the motile spores of Fungi and Algae, the spermatozoids of mosses, ferns, &c., move by means of delicate prolongations, cilia or flagella of the protoplast.
Chromatophores.—The chromatophores or plastids are protoplasmic structures, denser than the cytoplasm, and easily distinguishable from it by their colour or greater refractive power. They are spherical, oval, fusiform, or rod-like, and are always found in the cytoplasm, never in the cell-sap. They appear to be permanent organs of the cell, and are transmitted from one cell to another by division. In young cells the chromatophores are small, colourless, highly refractive bodies, principally located around the nucleus. As the cell grows they may become converted into leucoplasts (starch-formers), chloroplasts (chlorophyll-bodies), or chromoplasts (colour-bodies). And all three structures may be converted one into the other (Schimper). The chloroplasts are generally distinguished by their green colour, which is due to the presence of chlorophyll; but in many Algae this is masked by another colouring matter—Phycoerythrin in the Florideae, Phycophaein in the Phaeophyceae, and Phycocyanin
- ↑ Ewart, On the Physics and Physiology of Protoplasmic Streaming in Plants. (Oxford, 1903), gives an excellent account of the phenomena of protoplasmic streaming with a full discussion of the probable causes to which it is due.
