Darwin and Modern Science

by A. C. Seward

The result of a more thorough investigation of the determining conditions appears to produce at first sight a confused impression of all sorts of possibilities. Even closely allied species exhibit differences in regard to the connection between their development and external conditions. It is especially noteworthy that the same form in development may be produced as the result of very different alterations in the environment. At the same time we can undoubtedly detect a certain unity in the multiplicity of the individual phenomena.

If we compare the factors essential for the different stages in development, we see that the question always resolves itself into one of modification of similar conditions common to all life-processes. We should rather have inferred that there exist specific external stimuli for each developmental stage, for instance, certain chemical agencies. Experiments hitherto made support the conclusion that QUANTITATIVE alterations in the general conditions of life produce different types of development. An alga or a fungus grows so long as all the conditions of nutrition remain at a certain optimum for growth. In order to bring about asexual reproduction, e.g. the formation of zoospores, it is sometimes necessary to increase the degree of intensity of external factors; sometimes, on the other hand, these must be reduced in intensity. In the case of many algae a decrease in light-intensity or in the amount of salts in the culture solution, or in the temperature, induces asexual reproduction, while in others, on the contrary, an increase in regard to each of these factors is required to produce the same result. This holds good for the quantitative variations which induce sexual reproduction in algae. The controlling factor is found to be a reduction in the supply of nutritive salts and the exposure of the plants to prolonged illumination or, better still, an increase in the intensity of the light, the efficiency of illumination depending on the consequent formation of organic substances such as carbohydrates.

The quantitative alterations of external conditions may be spoken of as releasing stimuli. They produce, in the complex equilibrium of the cell, quantitative modifications in the arrangement and distribution of mass, by means of which other chemical processes are at once set in motion, and finally a new condition of equilibrium is attained. But the commonly expressed view that the environment can as a rule act only as a releasing agent is incorrect, because it overlooks an essential point. The power of a cell to receive stimuli is only acquired as the result of previous nutrition, which has produced a definite condition of concentration of different substances. Quantities are in this case the determining factors. The distribution of quantities is especially important in the sexual reproduction of algae, for which a vigorous production of the materials formed during carbon-assimilation appears to be essential.

In the Flowering plants, on the other hand, for reasons already mentioned, the whole problem is more complicated. Investigations on changes in the course of development of fertilised eggs have hitherto been unsuccessful; the difficulty of influencing egg-cells deeply immersed in tissue constitutes a serious obstacle. Other parts of plants are, however, convenient objects of experiment; e.g. the growing apices of buds which serve as cuttings for reproductive purposes, or buds on tubers, runners, rhizomes, etc. A growing apex consists of cells capable of division in which, as in egg-cells, a complete series of latent possibilities of development is embodied. Which of these possibilities becomes effective depends upon the action of the outer world transmitted by organs concerned with nutrition.

Of the different stages which a flowering plant passes through in the course of its development we will deal only with one in order to show that, in spite of its great complexity, the problem is, in essentials, equally open to attack in the higher plants and in the simplest organisms. The most important stage in the life of a flowering plant is the transition from purely vegetative growth to sexual reproduction—that is, the production of flowers. In certain cases it can be demonstrated that there is no internal cause, dependent simply on the specific structure, which compels a plant to produce its flowers after a definite period of vegetative growth. (Klebs, "Willkurliche Entwickelungsanderungen", Jena 1903; see also "Probleme der Entwickelung", I. II. "Centralbl." 1904.)

One extreme case, that of exceptionally early flowering, has been observed in nature and more often in cultivation. A number of plants under certain conditions are able to flower soon after germination. (Cf. numerous records of this kind by Diels, "Jugendformen und Bluten", Berlin, 1906.) This shortening of the period of development is exhibited in the most striking form in trees, as in the oak (Mobius, "Beitrage zur Lehre von der Fortpflanzung", Jena, 1897, page 89.), flowering seedlings of which have been observed from one to three years old, whereas normally the tree does not flower until it is sixty or eighty years old.

Another extreme case is represented by prolonged vegetative growth leading to the complete suppression of flower-production. This result may be obtained with several plants, such as Glechoma, the sugar beet, Digitalis, and others, if they are kept during the winter in a warm, damp atmosphere, and in rich soil; in the following spring or summer they fail to flower. (Klebs, "Willkurliche Aenderungen", etc. Jena, 1903, page 130.) Theoretically, however, experiments are of greater importance in which the production of flowers is inhibited by very favourable conditions of nutrition (Klebs, "Ueber kunstliche Metamorphosen", Stuttgart, 1906, page 115) ("Abh. Naturf. Ges. Halle", XXV.) occurring at the normal flowering period. Even in the case of plants of Sempervivum several years old, which, as is shown by control experiments on precisely similar plants, are on the point of flowering, flowering is rendered impossible if they are forced to very vigorous growth by an abundant supply of water and salts in the spring. Flowering, however, occurs, if such plants are cultivated in relatively dry sandy soil and in the presence of strong light. Careful researches into the conditions of growth have led, in the cases Sempervivum, to the following results: (1) With a strong light and vigorous carbon-assimilation a considerably increased supply of water and nutritive salts produces active vegetative growth. (2) With a vigorous carbon-assimilation in strong light, and a decrease in the supply of water and salts active flower-production is induced. (3) If an average supply of water and salts is given both processes are possible; the intensity of carbon-assimilation determines which of the two is manifested. A diminution in the production of organic substances, particularly of carbohydrates, induces vegetative growth. This can be effected by culture in feeble light or in light deprived of the yellow-red rays: on the other hand, flower-production follows an increase in light-intensity. These results are essentially in agreement with well-known observations on cultivated plants, according to which, the application of much moisture, after a plentiful supply of manure composed of inorganic salts, hinders the flower-production of many vegetables, while a decrease in the supply of water and salts favours flowering.

ii. INFLUENCE OF THE ENVIRONMENT ON THE FORM OF SINGLE ORGANS. (A considerable number of observations bearing on this question are given by Goebel in his "Experimentelle Morphologie der Pflanzen", Leipzig, 1908. It is not possible to deal here with the alteration in anatomical structure; cf. Kuster, "Pathologische Pflanzenanatomie", Jena, 1903.)

If we look closely into the development of a flowering plant, we notice that in a given species differently formed organs occur in definite positions. In a potato plant colourless runners are formed from the base of the main stem which grow underground and produce tubers at their tips: from a higher level foliage shoots arise nearer the apex. External appearances suggest that both the place of origin and the form of these organs were predetermined in the egg-cell or in the tuber. But it was shown experimentally by the well-known investigator Knight (Knight, "Selection from the Physiological and Horticultural Papers", London, 1841.) that tubers may be developed on the aerial stem in place of foliage shoots. These observations were considerably extended by Vochting. (Vochting, "Ueber die Bildung der Knollen", Cassel, 1887; see also "Bot. Zeit." 1902, 87.) In one kind of potato, germinating tubers were induced to form foliage shoots under the influence of a higher temperature; at a lower temperature they formed tuber-bearing shoots. Many other examples of the conversion of foliage-shoots into runners and rhizomes, or vice versa, have been described by Goebel and others. As in the asexual reproduction of algae quantitative alteration in the amount of moisture, light, temperature, etc. determines whether this or that form of shoot is produced. If the primordia of these organs are exposed to altered conditions of nutrition at a sufficiently early stage a complete substitution of one organ for another is effected. If the rudiment has reached a certain stage in development before it is exposed to these influences, extraordinary intermediate forms are obtained, bearing the characters of both organs.

The study of regeneration following injury is of greater importance as regards the problem of the development and place of origin of organs. (Reference may be made to the full summary of results given by Goebel in his "Experimentelle Morphologie", Leipzig and Berlin, 1908, Section IV.) Only in relatively very rare cases is there a complete re-formation of the injured organ itself, as e.g. in the growing-apex. Much more commonly injury leads to the development of complementary formations, it may be the rejuvenescence of a hitherto dormant rudiment, or it may be the formation of such ab initio. In all organs, stems, roots, leaves, as well as inflorescences, this kind of regeneration, which occurs in a great variety of ways according to the species, may be observed on detached pieces of the plant. Cases are also known, such, for example, as the leaves of many plants which readily form roots but not shoots, where a complete regeneration does not occur.