Chapter X: "On the Imperfection of the Geological Record," and Chapter (2)

The seeds of the Pteridosperms, unlike those of the Lycopod stock, have not yet been found in statu nascendi--in all known cases they were already highly developed organs and far removed from the cryptogamic sporangium. But in two respects we find that these seeds, or some of them, had not yet realised their possibilities. In the seed of Lyginodendron and other cases the micropyle, or orifice of the integument, was not the passage through which the pollen entered; the open neck of the pollen-chamber protruded through the micropyle and itself received the pollen. We have met with an analogous case, at a more advanced stage of evolution, in the Bennettiteae, where the wall of the gynaecium, though otherwise closed, did not provide a stigma to catch the pollen, but allowed the micropyles of the ovules to protrude and receive the pollen in the old gymnospermous fashion. The integument in the one case and the pistil in the other had not yet assumed all the functions to which the organ ultimately became adapted. Again, no Palaeozoic seed has yet been found to contain an embryo, though the preservation is often good enough for it to have been recognised if present. It is probable that the nursing of the embryo had not yet come to be one of the functions of the seed, and that the whole embryonic development was relegated to the germination stage.

In these two points, the reception of the pollen by the micropyle and the nursing of the embryo, it appears that many Palaeozoic seeds were imperfect, as compared with the typical seeds of later times. As evolution went on, one function was superadded on another, and it appears impossible to resist the conclusion that the whole differentiation of the seed was a process of adaptation, and consequently governed by Natural Selection, just as much as the specialisation of the rostellum in an Orchid, or of the pappus in a Composite.

Did space allow, other examples might be added. We may venture to maintain that the glimpses which the fossil record allows us into early stages in the evolution of organs now of high systematic importance, by no means justify the belief in any essential distinction between morphological and adaptive characters.

Another point, closely connected with Darwin's theory, on which the fossil history of plants has been supposed to have some bearing, is the question of Mutation, as opposed to indefinite variation. Arber and Parkin, in their interesting memoir on the Origin of Angiosperms, have suggested calling in Mutation to explain the apparently sudden transition from the cycadean to the angiospermous type of foliage, in late Mesozoic times, though they express themselves with much caution, and point out "a distinct danger that Mutation may become the last resort of the phylogenetically destitute"!

The distinguished French palaeobotanists, Grand'Eury (C. Grand'Eury, "Sur les mutations de quelques Plantes fossiles du Terrain houiller". "Comptes Rendus", CXLII. page 25, 1906.) and Zeiller (R. Zeiller "Les Vegetaux fossiles et leurs Enchainements", "Revue du Mois", III. February, 1907.), are of opinion, to quote the words of the latter writer, that the facts of fossil Botany are in agreement with the sudden appearance of new forms, differing by marked characters from those that have given them birth; he adds that these results give more amplitude to this idea of Mutation, extending it to groups of a higher order, and even revealing the existence of discontinuous series between the successive terms of which we yet recognise bonds of filiation. (Loc. cit. page 23.)

If Zeiller's opinion should be confirmed, it would no doubt be a serious blow to the Darwinian theory. As Darwin said: "Under a scientific point of view, and as leading to further investigation, but little advantage is gained by believing that new forms are suddenly developed in an inexplicable manner from old and widely different forms, over the old belief in the creation of species from the dust of the earth." ("Origin of Species", page 424.)

It most however be pointed out, that such mutations as Zeiller, and to some extent Arber and Parkin, appear to have in view, bridging the gulf between different Orders and Classes, bear no relation to any mutations which have been actually observed, such as the comparatively small changes, of sub-specific value, described by De Vries in the type-case of Oenothera Lamarckiana. The results of palaeobotanical research have undoubtedly tended to fill up gaps in the Natural System of plants--that many such gaps still persist is not surprising; their presence may well serve as an incentive to further research but does not, as it seems to the writer, justify the assumption of changes in the past, wholly without analogy among living organisms.

As regards the succession of species, there are no greater authorities than Grand'Eury and Zeiller, and great weight must be attached to their opinion that the evidence from continuous deposits favours a somewhat sudden change from one specific form to another. At the same time it will be well to bear in mind that the subject of the "absence of numerous intermediate varieties in any single formation" was fully discussed by Darwin. ("Origin of Species", pages 275-282, and page 312.); the explanation which he gave may go a long way to account for the facts which recent writers have regarded as favouring the theory of saltatory mutation.

The rapid sketch given in the present essay can do no more than call attention to a few salient points, in which the palaeontological records of plants has an evident bearing on the Darwinian theory. At the present day the whole subject of palaeobotany is a study in evolution, and derives its chief inspiration from the ideas of Darwin and Wallace. In return it contributes something to the verification of their teaching; the recent progress of the subject, in spite of the immense difficulties which still remain, has added fresh force to Darwin's statement that "the great leading facts in palaeontology agree admirably with the theory of descent with modification through variation and natural selection." (Ibid. page 313.)

XIII. THE INFLUENCE OF ENVIRONMENT ON THE FORMS OF PLANTS. By Georg Klebs, PH.D.

Professor of Botany in the University of Heidelberg.

The dependence of plants on their environment became the object of scientific research when the phenomena of life were first investigated and physiology took its place as a special branch of science. This occurred in the course of the eighteenth century as the result of the pioneer work of Hales, Duhamel, Ingenhousz, Senebier and others. In the nineteenth century, particularly in the second half, physiology experienced an unprecedented development in that it began to concern itself with the experimental study of nutrition and growth, and with the phenomena associated with stimulus and movement; on the other hand, physiology neglected phenomena connected with the production of form, a department of knowledge which was the province of morphology, a purely descriptive science. It was in the middle of the last century that the growth of comparative morphology and the study of phases of development reached their highest point.

The forms of plants appeared to be the expression of their inscrutable inner nature; the stages passed through in the development of the individual were regarded as the outcome of purely internal and hidden laws. The feasibility of experimental inquiry seemed therefore remote. Meanwhile, the recognition of the great importance of such a causal morphology emerged from the researches of the physiologists of that time, more especially from those of Hofmeister (Hofmeister, "Allgemeine Morphologie", Leipzig, 1868, page 579.), and afterwards from the work of Sachs. (Sachs, "Stoff und Form der Pflanzenorgane", Vol. I. 1880; Vol. II. 1882. "Gesammelte Abhandlungen uber Pflanzen-Physiologie", II. Leipzig, 1893.) Hofmeister, in speaking of this line of inquiry, described it as "the most pressing and immediate aim of the investigator to discover to what extent external forces acting on the organism are of importance in determining its form." This advance was the outcome of the influence of that potent force in biology which was created by Darwin's "Origin of Species" (1859).

The significance of the splendid conception of the transformation of species was first recognised and discussed by Lamarck (1809); as an explanation of transformation he at once seized upon the idea--an intelligible view--that the external world is the determining factor. Lamarck (Lamarck, "Philosophie zoologique", pages 223-227. Paris, 1809.) endeavoured, more especially, to demonstrate from the behaviour of plants that changes in environment induce change in form which eventually leads to the production of new species. In the case of animals, Lamarck adopted the teleological view that alterations in the environment first lead to alterations in the needs of the organisms, which, as the result of a kind of conscious effort of will, induce useful modifications and even the development of new organs. His work has not exercised any influence on the progress of science: Darwin himself confessed in regard to Lamarck's work--"I got not a fact or idea from it." ("Life and Letters", Vol. II. page 215.)

On a mass of incomparably richer and more essential data Darwin based his view of the descent of organisms and gained for it general acceptance; as an explanation of modification he elaborated the ingeniously conceived selection theory. The question of special interest in this connection, namely what is the importance of the influence of the environment, Darwin always answered with some hesitation and caution, indeed with a certain amount of indecision.

The fundamental principle underlying his theory is that of general variability as a whole, the nature and extent of which, especially in cultivated organisms, are fully dealt with in his well-known book. (Darwin, "The variation of Animals and Plants under domestication", 2 vols., edition 1, 1868; edition 2, 1875; popular edition 1905.) In regard to the question as to the cause of variability Darwin adopts a consistently mechanical view. He says: "These several considerations alone render it probable that variability of every kind is directly or indirectly caused by changed conditions of life. Or, to put the case under another point of view, if it were possible to expose all the individuals of a species during many generations to absolutely uniform conditions of life, there would be no variability." ("The variation of Animals and Plants" (2nd edition), Vol. II. page 242.) Darwin did not draw further conclusions from this general principle.

Variations produced in organisms by the environment are distinguished by Darwin as "the definite" and "the indefinite." (Ibid. II. page 260. See also "Origin of Species" (6th edition), page 6.) The first occur "when all or nearly all the offspring of an individual exposed to certain conditions during several generations are modified in the same manner." Indefinite variation is much more general and a more important factor in the production of new species; as a result of this, single individuals are distinguished from one another by "slight" differences, first in one then in another character. There may also occur, though this is very rare, more marked modifications, "variations which seem to us in our ignorance to arise spontaneously." ("Origin of Species" (6th edition), page 421.) The selection theory demands the further postulate that such changes, "whether extremely slight or strongly marked," are inherited. Darwin was no nearer to an experimental proof of this assumption than to the discovery of the actual cause of variability. It was not until the later years of his life that Darwin was occupied with the "perplexing problem... what causes almost every cultivated plant to vary" ("Life and Letters", Vol. III. page 342.): he began to make experiments on the influence of the soil, but these were soon given up.

In the course of the violent controversy which was the outcome of Darwin's work the fundamental principles of his teaching were not advanced by any decisive observations. Among the supporters and opponents, Nageli (Nageli, "Theorie der Abstammungslehre", Munich, 1884; cf. Chapter III.) was one of the few who sought to obtain proofs by experimental methods. His extensive cultural experiments with alpine Hieracia led him to form the opinion that the changes which are induced by an alteration in the food-supply, in climate or in habitat, are not inherited and are therefore of no importance from the point of view of the production of species. And yet Nageli did attribute an important influence to the external world; he believed that adaptations of plants arise as reactions to continuous stimuli, which supply a need and are therefore useful. These opinions, which recall the teleological aspect of Lamarckism, are entirely unsupported by proof. While other far-reaching attempts at an explanation of the theory of descent were formulated both in Nageli's time and afterwards, some in support of, others in opposition to Darwin, the necessity of investigating, from different standpoints, the underlying causes, variability and heredity, was more and more realised. To this category belong the statistical investigations undertaken by Quetelet and Galton, the researches into hybridisation, to which an impetus was given by the re-discovery of the Mendelian law of segregation, as also by the culture experiments on mutating species following the work of de Vries, and lastly the consideration of the question how far variation and heredity are governed by external influences. These latter problems, which are concerned in general with the causes of form-production and form-modification, may be treated in a short summary which falls under two heads, one having reference to the conditions of form-production in single species, the other being concerned with the conditions governing the transformation of species.

I. THE INFLUENCE OF EXTERNAL CONDITIONS ON FORM-PRODUCTION IN SINGLE SPECIES.

The members of plants, which we express by the terms stem, leaf, flower, etc. are capable of modification within certain limits; since Lamarck's time this power of modification has been brought more or less into relation with the environment. We are concerned not only with the question of experimental demonstration of this relationship, but, more generally, with an examination of the origin of forms, the sequences of stages in development that are governed by recognisable causes. We have to consider the general problem; to study the conditions of all typical as well as of atypic forms, in other words, to found a physiology of form.

If we survey the endless variety of plant-forms and consider the highly complex and still little known processes in the interior of cells, and if we remember that the whole of this branch of investigation came into existence only a few decades ago, we are able to grasp the fact that a satisfactory explanation of the factors determining form cannot be discovered all at once. The goal is still far away. We are not concerned now with the controversial question, whether, on the whole, the fundamental processes in the development of form can be recognised by physiological means. A belief in the possibility of this can in any case do no harm. What we may and must attempt is this--to discover points of attack on one side or another, which may enable us by means of experimental methods to come into closer touch with these elusive and difficult problems. While we are forced to admit that there is at present much that is insoluble there remains an inexhaustible supply of problems capable of solution.

The object of our investigations is the species; but as regards the question, what is a species, science of to-day takes up a position different from that of Darwin. For him it was the Linnean species which illustrates variation: we now know, thanks to the work of Jordan, de Bary, and particularly to that of de Vries (de Vries, "Die Mutationstheorie", Leipzig, 1901, Vol. I. page 33.), that the Linnean species consists of a large or small number of entities, elementary species. In experimental investigation it is essential that observations be made on a pure species, or, as Johannsen (Johannsen, "Ueber Erblichkeit in Populationen und reinen Linien", Jena, 1903.) says, on a pure "line." What has long been recognised as necessary in the investigation of fungi, bacteria and algae must also be insisted on in the case of flowering plants; we must start with a single individual which is reproduced vegetatively or by strict self-fertilisation. In dioecious plants we must aim at the reproduction of brothers and sisters.

We may at the outset take it for granted that a pure species remains the same under similar external conditions; it varies as these vary. IT IS CHARACTERISTIC OF A SPECIES THAT IT ALWAYS EXHIBITS A CONSTANT RELATION TO A PARTICULAR ENVIRONMENT. In the case of two different species, e.g. the hay and anthrax bacilli or two varieties of Campanula with blue and white flowers respectively, a similar environment produces a constant difference. The cause of this is a mystery.

According to the modern standpoint, the living cell is a complex chemico-physical system which is regarded as a dynamical system of equilibrium, a conception suggested by Herbert Spencer and which has acquired a constantly increasing importance in the light of modern developments in physical chemistry. The various chemical compounds, proteids, carbohydrates, fats, the whole series of different ferments, etc. occur in the cell in a definite physical arrangement. The two systems of two species must as a matter of fact possess a constant difference, which it is necessary to define by a special term. We say, therefore, that the SPECIFIC STRUCTURE is different.

By way of illustrating this provisionally, we may assume that the proteids of the two species possess a constant chemical difference. This conception of specific structure is specially important in its bearing on a further treatment of the subject. In the original cell, eventually also in every cell of a plant, the characters which afterwards become apparent must exist somewhere; they are integral parts of the capabilities or potentialities of specific structure. Thus not only the characters which are exhibited under ordinary conditions in nature, but also many others which become apparent only under special conditions (In this connection I leave out of account, as before, the idea of material carriers of heredity which since the publication of Darwin's Pangenesis hypothesis has been frequently suggested. See my remarks in "Variationen der Bluten", "Pringsheim's Jahrb. Wiss. Bot." 1905, page 298; also Detto, "Biol. Centralbl." 1907, page 81, "Die Erklarbarkeit der Ontogenese durch materielle Anlagen".), are to be included as such potentialities in cells; the conception of specific structure includes the WHOLE OF THE POTENTIALITIES OF A SPECIES; specific structure comprises that which we must always assume without being able to explain it.

A relatively simple substance, such as oxalate of lime, is known under a great number of different crystalline forms belonging to different systems (Compare Kohl's work on "Anatomisch-phys. Untersuchungen uber Kalksalze", etc. Marburg, 1889.); these may occur as single crystals, concretions or as concentric sphaerites. The power to assume this variety of form is in some way inherent in the molecular structure, though we cannot, even in this case, explain the necessary connection between structure and crystalline form. These potentialities can only become operative under the influence of external conditions; their stimulation into activity depends on the degree of concentration of the various solutions, on the nature of the particular calcium salt, on the acid or alkaline reactions. Broadly speaking, the plant cell behaves in a similar way. The manifestation of each form, which is inherent as a potentiality in the specific structure, is ultimately to be referred to external conditions.

An insight into this connection is, however, rendered exceedingly difficult, often quite impossible, because the environment never directly calls into action the potentialities. Its influence is exerted on what we may call the inner world of the organism, the importance of which increases with the degree of differentiation. The production of form in every plant depends upon processes in the interior of the cells, and the nature of these determines which among the possible characters is to be brought to light. In no single case are we acquainted with the internal process responsible for the production of a particular form. All possible factors may play a part, such as osmotic pressure, permeability of the protoplasm, the degree of concentration of the various chemical substances, etc.; all these factors should be included in the category of INTERNAL CONDITIONS. This inner world appears the more hidden from our ken because it is always represented by a certain definite state, whether we are dealing with a single cell or with a small group of cells. These have been produced from pre-existing cells and they in turn from others; the problem is constantly pushed back through a succession of generations until it becomes identified with that of the origin of species.

A way, however, is opened for investigation; experience teaches us that this inner world is not a constant factor: on the contrary, it appears to be very variable. The dependence of VARIABLE INTERNAL on VARIABLE EXTERNAL conditions gives us the key with which research may open the door. In the lower plants this dependence is at once apparent, each cell is directly subject to external influences. In the higher plants with their different organs, these influences were transmitted to cells in course of development along exceedingly complex lines. In the case of the growing-point of a bud, which is capable of producing a complete plant, direct influences play a much less important part than those exerted through other organs, particularly through the roots and leaves, which are essential in nutrition. These correlations, as we may call them, are of the greatest importance as aids to an understanding of form-production. When a bud is produced on a particular part of a plant, it undergoes definite internal modifications induced by the influence of other organs, the activity of which is governed by the environment, and as the result of this it develops along a certain direction; it may, for example, become a flower. The particular direction of development is determined before the rudiment is differentiated and is exerted so strongly that further development ensues without interruption, even though the external conditions vary considerably and exert a positively inimical influence: this produces the impression that development proceeds entirely independently of the outer world. The widespread belief that such independence exists is very premature and at all events unproven.

The state of the young rudiment is the outcome of previous influences of the external world communicated through other organs. Experiments show that in certain cases, if the efficiency of roots and leaves as organs concerned with nutrition is interfered with, the production of flowers is affected, and their characters, which are normally very constant, undergo far-reaching modifications. To find the right moment at which to make the necessary alteration in the environment is indeed difficult and in many cases not yet possible. This is especially the case with fertilised eggs, which in a higher degree than buds have acquired, through parental influences, an apparently fixed internal organisation, and this seems to have pre-determined their development. It is, however, highly probable that it will be possible, by influencing the parents, to alter the internal organisation and to switch off development on to other lines.

Having made these general observations I will now cite a few of the many facts at our disposal, in order to illustrate the methods and aim of the experimental methods of research. As a matter of convenience I will deal separately with modification of development and with modification of single organs.

I. EFFECT OF ENVIRONMENT UPON THE COURSE OF DEVELOPMENT.

Every plant, whether an alga or a flowering plant passes, under natural conditions, through a series of developmental stages characteristic of each species, and these consist in a regular sequence of definite forms. It is impossible to form an opinion from mere observation and description as to what inner changes are essential for the production of the several forms. We must endeavour to influence the inner factors by known external conditions in such a way that the individual stages in development are separately controlled and the order of their sequence determined at will by experimental treatment. Such control over the course of development may be gained with special certainty in the case of the lower organisms.

With these it is practicable to control the principal conditions of cultivation and to vary them in various ways. By this means it has been demonstrated that each developmental stage depends upon special external conditions, and in cases where our knowledge is sufficient, a particular stage may be obtained at will. In the Green Algae (See Klebs, "Die Bedingung der Fortpflanzung... ", Jena, 1896; also "Jahrb. fur Wiss. Bot." 1898 and 1900; "Probleme der Entwickelung, III." "Biol. Centralbl." 1904, page 452.), as in the case of Fungi, we may classify the stages of development into purely vegetative growth (growth, cell-division, branching), asexual reproduction (formation of zoospores, conidia) and sexual processes (formation of male and female sexual organs). By modifying the external conditions it is possible to induce algae or fungi (Vaucheria, Saprolegnia) to grow continuously for several years or, in the course of a few days, to die after an enormous production of asexual or sexual cells. In some instances even an almost complete stoppage of growth may be caused, reproductive cells being scarcely formed before the organism is again compelled to resort to reproduction. Thus the sequence of the different stages in development can be modified as we may desire.

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.

The widely spread power of reacting to wounding affords a very valuable means of inducing a fresh development of buds and roots on places where they do not occur in normal circumstances. Injury creates special conditions, but little is known as yet in regard to alterations directly produced in this way. Where the injury consists in the separation of an organ from its normal connections, the factors concerned are more comprehensible. A detached leaf, e.g., is at once cut off from a supply of water and salts, and is deprived of the means of getting rid of organic substances which it produces; the result is a considerable alteration in the degree of concentration. No experimental investigation on these lines has yet been made. Our ignorance has often led to the view that we are dealing with a force whose specific quality is the restitution of the parts lost by operation; the proof, therefore, that in certain cases a similar production of new roots or buds may be induced without previous injury and simply by a change in external conditions assumes an importance. (Klebs, "Willkurliche Entwickelung", page 100; also, "Probleme der Entwickelung", "Biol. Centralbl." 1904, page 610.)

A specially striking phenomenon of regeneration, exhibited also by uninjured plants, is afforded by polarity, which was discovered by Vochting. (See the classic work of Vochting, "Ueber Organbildung im Pflanzenreich", I. Bonn, 1888; also "Bot. Zeit." 1906, page 101; cf. Goebel, "Experimentelle Morphologie", Leipzig and Berlin, 1908, Section V, Polaritat.) It is found, for example, that roots are formed from the base of a detached piece of stem and shoots from the apex. Within the limits of this essay it is impossible to go into this difficult question; it is, however, important from the point of view of our general survey to emphasise the fact that the physiological distinctions between base and apex of pieces of stem are only of a quantitative kind, that is, they consist in the inhibition of certain phenomena or in favouring them. As a matter of fact roots may be produced from the apices of willows and cuttings of other plants; the distinction is thus obliterated under the influence of environment. The fixed polarity of cuttings from full grown stems cannot be destroyed; it is the expression of previous development. Vochting speaks of polarity as a fixed inherited character. This is an unconvincing conclusion, as nothing can be deduced from our present knowledge as to the causes which led up to polarity. We know that the fertilised egg, like the embryo, is fixed at one end by which it hangs freely in the embryo-sac and afterwards in the endosperm. From the first, therefore, the two ends have different natures, and these are revealed in the differentiation into root-apex and stem-apex. A definite direction in the flow of food-substances is correlated with this arrangement, and this eventually leads to a polarity in the tissues. This view requires experimental proof, which in the case of the egg-cells of flowering plants hardly appears possible; but it derives considerable support from the fact that in herbaceous plants, e.g. Sempervivum (Klebs, "Variationen der Bluten", "Jahrb. Wiss. Bot." 1905, page 260.), rosettes or flower-shoots are formed in response to external conditions at the base, in the middle, or at the apex of the stem, so that polarity as it occurs under normal conditions cannot be the result of unalterable hereditary factors. On the other hand, the lower plants should furnish decisive evidence on this question, and the experiments of Stahl, Winkler, Kniep, and others indicate the right method of attacking the problem.

The relation of leaf-form to environment has often been investigated and is well known. The leaves of bog and water plants (Cf.Goebel, loc. cit. chapter II.; also Gluck, "Untersuchungen uber Wasser- und Sumpfgewachse", Jena, Vols. I.-II. 1905-06.) afford the most striking examples of modifications: according as they are grown in water, moist or dry air, the form of the species characteristic of the particular habitat is produced, since the stems are also modified. To the same group of phenomena belongs the modification of the forms of leaves and stems in plants on transplantation from the plains to the mountains (Bonnier, "Recherches sur l'Anatomie experimentale des Vegetaux", Corbeil, 1895.) or vice versa. Such variations are by no means isolated examples. All plants exhibit a definite alteration in form as the result of prolonged cultivation in moist or dry air, in strong or feeble light, or in darkness, or in salt solutions of different composition and strength.

Every individual which is exposed to definite combinations of external factors exhibits eventually the same type of modification. This is the type of variation which Darwin termed "definite." It is easy to realise that indefinite or fluctuating variations belong essentially to the same class of phenomena; both are reactions to changes in environment. In the production of individual variations two different influences undoubtedly cooperate. One set of variations is caused by different external conditions, during the production, either of sexual cells or of vegetative primordia; another set is the result of varying external conditions during the development of the embryo into an adult plant. The two sets of influences cannot as yet be sharply differentiated. If, for purposes of vegetative reproduction, we select pieces of the same parent-plant of a pure species, the second type of variation predominates. Individual fluctuations depend essentially in such cases on small variations in environment during development.

These relations must be borne in mind if we wish to understand the results of statistical methods. Since the work of Quetelet, Galton, and others the statistical examination of individual differences in animals and plants has become a special science, which is primarily based on the consideration that the application of the theory of probability renders possible mathematical statement and control of the results. The facts show that any character, size of leaf, length of stem, the number of members in a flower, etc. do not vary haphazard but in a very regular manner. In most cases it is found that there is a value which occurs most commonly, the average or medium value, from which the larger and smaller deviations, the so-called plus and minus variations fall away in a continuous series and end in a limiting value. In the simpler cases a falling off occurs equally on both sides of the curve; the curve constructed from such data agrees very closely with the Gaussian curve of error. In more complicated cases irregular curves of different kinds are obtained which may be calculated on certain suppositions.

The regular fluctuations about a mean according to the rule of probability is often attributed to some law underlying variability. (de Vries, "Mutationstheorie", Vol. I. page 35, Leipzig, 1901.) But there is no such law which compels a plant to vary in a particular manner. Every experimental investigation shows, as we have already remarked, that the fluctuation of characters depends on fluctuation in the external factors. The applicability of the method of probability follows from the fact that the numerous individuals of a species are influenced by a limited number of variable conditions. (Klebs, "Willkurl. Ent." Jena, 1903, page 141.) As each of these conditions includes within certain limits all possible values and exhibits all possible combinations, it follows that, according to the rules of probability, there must be a mean value, about which the larger and smaller deviations are distributed. Any character will be found to have the mean value which corresponds with that combination of determining factors which occurs most frequently. Deviations towards plus and minus values will be correspondingly produced by rarer conditions.

A conclusion of fundamental importance may be drawn from this conception, which is, to a certain extent, supported by experimental investigation. (Klebs, "Studien uber Variation", "Arch. fur Entw." 1907.) There is no normal curve for a particular CHARACTER, there is only a curve for the varying combinations of conditions occurring in nature or under cultivation. Under other conditions entirely different curves may be obtained with other variants as a mean value. If, for example, under ordinary conditions the number 10 is the most frequent variant for the stamens of Sedum spectabile, in special circumstances (red light) this is replaced by the number 5. The more accurately we know the conditions for a particular form or number, and are able to reproduce it by experiment, the nearer we are to achieving our aim of rendering a particular variation impossible or of making it dominant.

In addition to the individual variations of a species, more pronounced fluctuations occur relatively rarely and sporadically which are spoken of as "single variations," or if specially striking as abnormalities or monstrosities. These forms have long attracted the attention of morphologists; a large number of observations of this kind are given in the handbooks of Masters (Masters, "Vegetable Teratology", London, 1869.) and Penzig (Penzig, "Pflanzen-Teratologie", Vols I. and II. Genua, 1890-94.) These variations, which used to be regarded as curiosities, have now assumed considerable importance in connection with the causes of form-development. They also possess special interest in relation to the question of heredity, a subject which does not at present concern us, as such deviations from normal development undoubtedly arise as individual variations induced by the influence of environment.

Abnormal developments of all kinds in stems, leaves, and flowers, may be produced by parasites, insects, or fungi. They may also be induced by injury, as Blaringhem (Blaringhem, "Mutation et traumatismes", Paris, 1907.) has more particularly demonstrated, which, by cutting away the leading shoots of branches in an early stage of development, caused fasciation, torsion, anomalous flowers, etc. The experiments of Blaringhem point to the probability that disturbances in the conditions of food-supply consequent on injury are the cause of the production of monstrosities. This is certainly the case in my experiments with species of Sempervivum (Klebs, "Kunstliche Metamorphosen", Stuttgart, 1906.); individuals, which at first formed normal flowers, produced a great variety of abnormalities as the result of changes in nutrition, we may call to mind the fact that the formation of inflorescences occurs normally when a vigorous production of organic compounds, such as starch, sugar, etc. follows a diminution in the supply of mineral salts. On the other hand, the development of inflorescences is entirely suppressed if, at a suitable moment before the actual foundations have been laid, water and mineral salts are supplied to the roots. If, during the week when the inflorescence has just been laid down and is growing very slowly, the supply of water and salts is increased, the internal conditions of the cells are essentially changed. At a later stage, after the elongation of the inflorescence, rosettes of leaves are produced instead of flowers, and structures intermediate between the two kinds of organs; a number of peculiar plant-forms are thus obtained (Cf. Lotsy, "Vorlesungen uber Deszendenztheorien", Vol. II. pl. 3, Jena, 1908.) Abnormalities in the greatest variety are produced in flowers by varying the time at which the stimulus is applied, and by the cooperation of other factors such as temperature, darkness, etc. In number and arrangement the several floral members vary within wide limits; sepals, petals, stamens, and carpels are altered in form and colour, a transformation of stamens to carpels and from carpels to stamens occurs in varying degrees. The majority of the deviations observed had not previously been seen either under natural conditions or in cultivation; they were first brought to light through the influence of external factors.

Such transformations of flowers become apparent at a time, which is separated by about two months from the period at which the particular cause began to act. There is, therefore, no close connection between the appearance of the modifications and the external conditions which prevail at the moment. When we are ignorant of the causes which are operative so long before the results are seen, we gain the impression that such variations as occur are spontaneous or autonomous expressions of the inner nature of the plant. It is much more likely that, as in Sempervivum, they were originally produced by an external stimulus which had previously reached the sexual cells or the young embryo. In any case abnormalities of this kind appear to be of a special type as compared with ordinary fluctuating variations. Darwin pointed out this difference; Bateson (Bateson, "Materials for the study of Variation", London, 1894, page 5.) has attempted to make the distinction sharper, at the same time emphasising its importance in heredity.

Bateson applies the term CONTINUOUS to small variations connected with one another by transitional stages, while those which are more striking and characterised from the first by a certain completeness, he names DISCONTINUOUS. He drew attention to a great difficulty which stands in the way of Lamarck's hypothesis, as also of Darwin's view. "According to both theories, specific diversity of form is consequent upon diversity of environment, and diversity of environment is thus the ultimate measure of diversity of specific form. Here then we meet the difficulty that diverse environments often shade into each other insensibly and form a continuous series, whereas the Specific Forms of life which are subject to them on the whole form a Discontinuous Series." This difficulty is, however, not of fundamental importance as well authenticated facts have been adduced showing that by alteration of the environment discontinuous variations, such as alterations in the number and form of members of a flower, may be produced. We can as yet no more explain how this happens than we can explain the existence of continuous variations. We can only assert that both kinds of variation arise in response to quantitative alterations in external conditions. The question as to which kind of variation is produced depends on the greater or less degree of alteration; it is correlated with the state of the particular cells at the moment.

In this short sketch it is only possible to deal superficially with a small part of the subject. It has been clearly shown that in view of the general dependence of development on the factors of the environment a number of problems are ready for experimental treatment. One must, however, not forget that the science of the physiology of form has not progressed beyond its initial stages. Just now our first duty is to demonstrate the dependence on external factors in as many forms of plants as possible, in order to obtain a more thorough control of all the different plant-forms. The problem is not only to produce at will (and independently of their normal mode of life) forms which occur in nature, but also to stimulate into operation potentialities which necessarily lie dormant under the conditions which prevail in nature. The constitution of a species is much richer in possibilities of development than would appear to be the case under normal conditions. It remains for man to stimulate into activity all the potentialities.

But the control of plant-form is only a preliminary step--the foundation stones on which to erect a coherent scientific structure. We must discover what are the internal processes in the cell produced by external factors, which as a necessary consequence result in the appearance of a definite form. We are here brought into contact with the most obscure problem of life. Progress can only be made pari passu with progress in physics and chemistry, and with the growth of our knowledge of nutrition, growth, etc.

Let us take one of the simplest cases--an alteration in form. A cylindrical cell of the alga Stigeoclonium assumes, as Livingstone (Livingstone, "On the nature of the stimulus which causes the change of form, etc." "Botanical Gazette", XXX. 1900; also XXXII. 1901.) has shown, a spherical form when the osmotic pressure of the culture fluid is increased; or a spore of Mucor, which, in a sugar solution grows into a branched filament, in the presence of a small quantity of acid (hydrogen ions) becomes a comparatively large sphere. (Ritter, "Ueber Kugelhefe, etc." "Ber. bot. Gesell." Berlin, XXV. page 255, 1907.) In both cases there has undoubtedly been an alteration in the osmotic pressure of the cell-sap, but this does not suffice to explain the alteration in form, since the unknown alterations, which are induced in the protoplasm, must in their turn influence the cell-membrane. In the case of the very much more complex alterations in form, such as we encounter in the course of development of plants, there do not appear to be any clues which lead us to a deeper insight into the phenomena. Nevertheless we continue the attempt, seeking with the help of any available hypothesis for points of attack, which may enable us to acquire a more complete mastery of physiological methods. To quote a single example; I may put the question, what internal changes produce a transition from vegetative growth to sexual reproduction?

The facts, which are as clearly established from the lower as for the higher plants, teach us that quantitative alteration in the environment produces such a transition. This suggests the conclusion that quantitative internal changes in the cells, and with them disturbances in the degree of concentration, are induced, through which the chemical reactions are led in the direction of sexual reproduction. An increase in the production of organic substances in the presence of light, chiefly of the carbohydrates, with a simultaneous decrease in the amount of inorganic salts and water, are the cause of the disturbance and at the same time of the alteration in the direction of development. Possibly indeed mineral salts as such are not in question, but only in the form of other organic combinations, particularly proteid material, so that we are concerned with an alteration in the relation of the carbohydrates and proteids. The difficulties of such researches are very great because the methods are not yet sufficiently exact to demonstrate the frequently small quantitative differences in chemical composition. Questions relating to the enzymes, which are of the greatest importance in all these life-processes, are especially complicated. In any case it is the necessary result of such an hypothesis that we must employ chemical methods of investigation in dealing with problems connected with the physiology of form.

II. INFLUENCE OF ENVIRONMENT ON THE TRANSFORMATION OF SPECIES.