In connection with his researches into the importance of the constituents of the ash, de Saussure proposed the question whether roots take up the solutions of salts and other substances exactly in the form in which they offer themselves. He found first of all that very various and even poisonous matters are absorbed by them, and that there is therefore no such power of choice, as Jung had once supposed; on the other hand, it appeared that the solutions do not enter unchanged into the roots, for in his experiments in every case the proportion of water to the salt absorbed was greater than the proportion between them in the solution, and that some salts enter the plant in larger, some in smaller quantities, under circumstances in other respects the same. But at this time, and for a long time after, it was not possible to understand and rightly explain these facts; the theory of diffusions was not yet known, and fifty or sixty years were to elapse before light was thrown on the questions thus raised by de Saussure.
Such were the most important contents of de Saussure’s publication in 1804. His later contributions to the knowledge of some important questions in vegetable physiology will be mentioned further on. A comparison of the contents of the ‘Recherches chimiques’ with what was known of the chemistry of the food of plants before 1780 excites the liveliest astonishment at the enormous advance made in these twenty-four years. The latter years of the 18th century had proved still more fruitful, if possible, as regards the theory of nutrition than the latter years of the 17th; both periods have this in common, that they developed an extraordinary abundance of new points of view in every branch of botanical science. They resemble each other also in the circumstance that they were both followed by a longer period of inactivity; the time from Hales to Ingen-Houss was highly unproductive, and so also were the thirty years that followed the appearance of de Saussure’s great work, though it must be admitted that some good work was done during that period in France, while in Germany the new theory was grossly misunderstood by the chief representatives of botany, as we shall see in the following section. It should be mentioned however that one of these misconceptions, which was not removed till after 1860, was caused by de Saussure himself. He had observed that the red leaves of a variety of the garden Orache disengage oxygen from carbon dioxide, as much as the green leaves of the common kind. In this case he was hasty, and concluded from this single observation that the green colour is not an essential character of the parts which decompose carbonic acid; if he had only removed the epidermis of the red leaves he would have found that the inner tissue is coloured as dark green as the ordinary green leaves. He who was usually so extremely careful as an observer was for once negligent, and later writers, as is apt to happen, fixed exactly on this one weak point, and repeatedly called in question one of the most weighty facts of vegetable physiology, namely, that only cells which contain chlorophyll eliminate oxygen.
5. Vital force. Respiration and heat of plants. Endosmose.
1804-1840.
During the twenty years that followed the appearance of de Saussure’s chemical researches the theory of the nutrition of plants can scarcely be said to have been advanced in any one direction, while much that had already been accomplished was not even understood. Various circumstances worked together to introduce misconceptions in this province of botany; above all others the inclination, more strongly pronounced than ever at this period, to attribute to organisms a special vital principle or force, which was supposed to possess a variety of wonderful powers, so that it could even produce elementary substances, heat, and other things out of nothing. Whenever any process in such organisms was difficult to explain by physical or chemical laws, the vital force was simply called in to bring about the phenomena in question in some inexplicable manner. It was not that the question was now raised, which at a later time engaged the attention of profounder thinkers, whether there was a special agent operating in organic bodies beside the general forces which govern inorganic nature; for a careful examination of this question would certainly have led to the most earnest efforts to explain all the phenomena of life by physical or chemical laws. On the contrary, it was found convenient to assume this vital force as proved, and to assign it as the cause of a variety of phenomena, thus escaping the necessity of explaining the way in which the effects were produced; in a word, the assumption of a vital force was not a hypothesis to stimulate investigation, but a phantom that made all intellectual efforts superfluous.
Another hindrance to the progress of physiology, especially where questions of nutrition turned on the movement of the sap, was the backward condition of the study of the inner structure of plants, as described in the second book. For instance, the question of the descending sap was complicated in the strangest way by Du Petit-Thouars’s theory of bud-roots that descend between the bark and the wood; Reichel’s unfounded idea of the rising of the sap in the tubes of the wood was generally accepted, and a still worse error was maintained by some, that the intercellular spaces of the parenchyma are true sap-conveying organs. In 1812 Moldenhawer had to insist, but without producing any general conviction, that the vessels of the wood contain air, and Treviranus in 1821 that the stomata serve for the entrance and exit of air. We need not notice here what nature-philosophers like Kieser said about nutrition and the movement of the sap; but even those who were far from adopting the extravagancies of this school were incapable of either making use of or carrying on the labours of Ingen-Houss, Senebier, and de Saussure. We may adduce in proof of this statement the remarks of Link on the function of leaves in his ‘Grundlehren der Anatomie und Physiologie,’ 1807. He says at p. 202 that their function is according to Hales transpiration, according to Bonnet absorption, according to Bjerkander the exudation and secretion of a variety of fluids, according to Hedwig the storing up of juices, and inasmuch as leaves increase the green surfaces of plants, bear stomata and hairs, and hold a quantity of juices in their abundant parenchyma, we may ascribe all these functions, but none of them exclusively, to leaves; the only thing peculiar to them is that they convey elaborated juices to the young parts. Their great work, the decomposition of carbon dioxide, he does not mention. But this neglect of the doctrines of Ingen-Houss, Senebier, and de Saussure was common, especially in Germany; it is seen in the efforts made to prove once more the existence of a descending sap in the rind, just as it had been proved in the two previous centuries, by the result of removing a ring of bark from the stem, and by similar experiments; whereas the simple consideration that it is only in the green leaves that carbonaceous vegetable substance is formed, would have made the existence of what was known as a descending sap appear to be a matter of course, and must have led to a much clearer conception of the matter. But this consideration was either quite overlooked or only mentioned incidentally by those who occupied themselves with experiments on the movement of the descending sap. This is the case in Heinrich Cotta’s ‘Naturbeobachtungen über die Bewegung und Function des Saftes in den Gewächsen,’ 1806, in many respects an instructive work, and in Knight’s otherwise serviceable experiments on the growth in thickness of trees. It was not till after 1830 that De Candolle and Dutrochet perceived that the fact that the green leaves are assimilating organs must be decisive of the question of the movement of the sap in the stem.
No progress was made with the general doctrine of nutrition between 1820 and 1840 except in one point, the absorption of oxygen by all parts of plants; here something was done to consolidate the theory and to enrich it with new facts; it was indeed a subject more adapted to the views of the day, because it at once suggested a variety of analogies with the respiration of animals. Grischow showed in 1819 that Fungi never decompose carbon dioxide, but absorb oxygen and give off carbon dioxide. Marcet carried the subject further in 1834, after de Saussure had published in 1822 an excellent investigation into the absorption of oxygen by flowers; in this work we have the basis laid for the theory of vegetable heat, to which we shall return. But Dutrochet was the first who made an elaborate comparison of the respiration of plants and animals (1837), and showed that not only growth, as de Saussure had already perceived, but also the sensitiveness of plants depends on the presence of oxygen, that is on their respiration. The recognition of the fact, that the inhalation of oxygen plays the same part in plants that it does in animals, prepared the way for the view that heat in plants is simply a result of their respiration, as it is in animals. It is not necessary to describe at length the experiments which were made on heat in plants before 1822; they were one and all vitiated by a want of clearness in the statement of the question, which made success impossible; it was assumed that this heat by raising the temperature of the plant would make itself felt by surrounding objects, and it was sought for exactly where it is least to be found, in the wood, in fruits and tubers, and generally in resting, inactive parts. Moreover the previous experiments, collected in Goeppert’s book ‘Ueber die Wärmeentwicklung der Pflanzen,’ 1830, were so unskilfully managed that they could not possibly lead to any result. Nor could the question whether plants really develope internal heat, as animals do, be determined by a few cases of active development of heat in flowers, because an idea was prevalent at the time in connection with the theory of a vital force, that flowers as the organs of reproduction alone possessed the power of generating heat.
Lavoisier had clearly perceived in 1777 that the combustion of substances containing carbon by inhaled oxygen was the source of animal heat, and had proved it by experiments. Senebier, who first observed the rise of temperature in the inflorescence of Arum by the thermometer, had at least suggested in his work on physiology of 1800 (iii. p. 315) that a vigorous absorption of oxygen might be the cause of the phenomenon. Bory de St. Vincent reported in 1804 that Hubert, the owner of a plantation in Madagascar, had observed among other things that the air in which the flowering spike of one of the Aroideae had developed its heat could support neither animal respiration nor combustion. These indications were however disregarded, until de Saussure in 1822 proved directly the connection between the absorption of oxygen and the rise of temperature in flowers. It was however a long time before heat in plants was conceived of as a general fact necessarily connected with their respiration. This conception would have swept away the whole mass of facts accumulated by Goeppert in his book of 1830, from which he tried to prove (p. 228) that plants at no period of their life possess the power of generating heat—a view which he retracted however in 1832, when he had observed a rise of temperature in germinating plants, bulbs, tubers, and in green plants, when collected into heaps. How difficult it was for physiologists under the dominion of the ‘vital force’ to hold firmly to the simple principle of natural heat, and not to be led away by isolated observations, is shown by the expressions of De Candolle in 1835, and still more by those of Treviranus in 1838. It is therefore refreshing to see Meyen in his ‘Neues System’ (1838), vol. ii, warmly asserting this principle, and making the development of heat in plants a necessary consequence of their respiration and of other chemical processes. Meyen himself produced no new observations; but Vrolik and De Vriese showed by laborious experiments in 1836 and 1839 the dependence of the generation of heat in the flowers of Aroideae on the absorption of oxygen. A higher importance as regards the general principle attaches to the attempt of Dutrochet in 1840 to prove that even growing shoots generate small quantities of heat, as shown by a thermo-electric apparatus. Some of the details in these observations are open to objection; but it cannot be denied that they are based on a clear recognition of the general principle, though they ignore the consideration that the generation of heat in plants is not necessarily accompanied with a rise in temperature, since cooling causes may be acting at the same time with greater effect. However the doctrine of the natural heat of plants was in the main established by the observations of de Saussure, Vrolik, De Vriese, and Dutrochet, and by Meyen’s and Dutrochet’s assertion of the principle laid down by Lavoisier, though thirty years elapsed before it became an accepted truth in vegetable physiology.
The crude idea of a vital force was deprived of one of its chief supports when it was recognised that the natural heat of organisms was a product of chemical processes induced by respiration, for this had been regarded since the time of Aristotle as more peculiarly an effect of the principle of life. And now another discovery was made, equally calculated to promote the reference to mechanical principles of those general and important phenomena of life which had hitherto been indolently ascribed to the operation of the vital force. It appears to be a matter of indifference whether Professor Fischer of Breslau is or is not to be considered as the true discoverer of endosmose in 1822, for it is certain that it was Dutrochet[135] who first studied the subject with exactness, and above all perceived its extraordinary value for the explanation of certain phenomena in living organisms. He repeatedly called attention to this value in the years between 1826 and 1837, and endeavoured to refer a variety of phenomena in vegetation to this agency. He had first observed the operation of endosmose in its mechanical effects in living bodies; the escape of the zoospores of an aquatic Fungus and the ejection of the sperm from the spermathecae of snails first led him to the hypothesis, that the more concentrated solutions inclosed in organic membranes exercise an attraction on surrounding water, which, forcing its way into the inclosed space, is there able to exert considerable powers of pressure. To Dutrochet must always belong the merit of having brought into notice this mechanical effect of endosmose and of employing it to explain a number of vital phenomena. Many things in which a mechanical explanation had not been hitherto thought of could now be traced to a mechanical principle, the effects of which could be exhibited and more accurately studied by means of artificial apparatus. Dutrochet rightly attached a special value to the fact, that all states of tension in vegetable tissue could be at once explained by endosmose and exosmose, though, as so often happens in such matters, he may have extended his new principle to cases where it was not applicable, as we shall see below. His account of the nature of endosmose itself must now be considered to be obsolete, nor did the mathematician Poisson or the physicist Magnus about 1830 succeed in framing a satisfactory theory on the subject. It was discovered in the course of the succeeding twenty or thirty years, that the phenomena observed by Dutrochet, and which he called endosmose and exosmose, were only complicated cases of hydro-diffusion, which with the diffusion of gas forms an important part of molecular physics. Dutrochet, like his immediate successors, conducted his investigations into osmose with animal and vegetable membranes, the latter being of a complex structure; with these he always observed in addition to the endosmotic flow of water into the more concentrated solution, an escape of the solution itself, and from this he concluded that there must always be two currents in opposite directions through the membrane which separates the two fluids, that, as he expresses it, the endosmose is always accompanied with exosmose. This error, which was even developed later into a theory of the endosmotic equivalent, has had much to do till recently with making it impossible or difficult to refer certain phenomena of vegetation to the processes of hydro-diffusion. To mention only one case, Schleiden rightly observed that if endosmose, as Dutrochet understood it, is the sole cause why water is absorbed by the roots, there must also be a corresponding exosmose at the roots; and this, which was called root-discharge, Macaire Prinsep thought he had actually discovered, and even Liebig firmly believed in its existence till a recent period, although the researches of Wiegman and Polstorff (1842) and later more careful investigations showed, that there was no noticeable discharge by exosmose to answer to the great quantity of water with substances in solution in it which is taken up by the roots. Again, Dutrochet’s theory of endosmose did not fully explain the way in which the several substances which feed the plant find their way into and are disseminated in it. But notwithstanding these and other defects it deserved the greatest consideration, because it gave the first impulse to the further development of the theory of diffusion, and contained a mechanical principle which might serve to explain very various phenomena in vegetation as yet unexplained. Dutrochet hastened to apply it to this purpose, where it was at all possible to do so, and chiefly in his treatise on the ascending and descending sap (‘Mémoires,’ 1837, i. p. 365), which was superior to anything which had been written on the movement of the sap in plants in its clear conception of the question and in perspicuity of treatment. It should be especially mentioned that Dutrochet formed a true estimate of the functions of the leaves as regards both the ascending and descending sap, and to some extent pointed out the fault which lies at the bottom of the earlier experiments with coloured fluids. After communicating a number of good observations on the paths of the ascending and descending sap, and noticing particularly that in the vine the vessels of the wood serve for the movement of the sap only in spring, when vines bleed, but that they are air-passages in summer, when transpiration causes the most copious flow of water in the wood, he proceeds to consider the forces which effect the movement of the ascending sap in the wood both in spring and summer. He first of all judiciously distinguishes two things which had been before always mixed up together, the weeping of severed root-stocks and the rise of the sap in the wood in transpiring plants. The first is caused, he thinks, by impulsion, the other by attraction; we should now say, that in weeping root-stocks the water is pressed upwards, in transpiring plants drawn up. He then refers the phenomenon of impulsion to endosmose in the roots, and without going much into detail as regards the anatomical conditions, he compares a weeping root-stock to his own endosmometer, in the tube of which the fluid that has been sucked in rises by endosmose and even flows over; it is true that no very thorough understanding of the matter was gained in this way, but at any rate the principle which was to explain it was indicated. He then endeavours to explain the movement of the water which ascends in the wood of transpiring plants by the action of endosmose from cell to cell. In this he failed entirely, as was afterwards perceived; but he succeeded in showing that all the mechanical explanations that had been previously attempted were incorrect, and the whole treatise, though unsatisfactory in its main result, contains a great number of ingenious experiments and acute remarks.
With the exception of Théodore de Saussure, who occupied himself exclusively with chemical questions in physiology, Dutrochet was the only vegetable physiologist in the period between 1820 and 1840 who studied all its more important questions thoroughly and experimentally; his treatise on the respiration of plants, which has been already mentioned, is excellent in itself, and was of the greatest importance at the time it appeared, because it brought the chemical processes in respiration, the entrance and exit of the gases, for the first time into correct connection with the air-passages in the plant, with the stomata, the vessels, and the intercellular spaces, and submitted the composition of the air contained in the cavities of plants to careful examination. It was the best work on the respiration of plants in the year 1837 and for a long time after; and if Dutrochet made the mistake of regarding the oxygen which is disengaged from the plant itself in the light as the chief agent in respiration, and the oxygen directly absorbed from the atmosphere as only subsidiary to this, he compensated for it by recognising the importance of the fact, that only cells which contain chlorophyll decompose carbon dioxide, and still more by correctly distinguishing between respiration by the absorption of oxygen and the decomposition of carbonic dioxide in light; these two processes were at that time and afterwards very inappropriately distinguished as the diurnal and nocturnal respiration of plants, and this misleading expression maintained itself in spite of Garreau’s protest in 1851 till after 1860, when a modern German physiologist succeeded in establishing the true distinction between respiration and assimilation in plants. Another mischievous complication arose about 1830 connected with the expression, circulation of the sap; it was thought that an argument for such a circulation even in the higher plants was to be found in the ‘circulation of the sap’ (protoplasm) in the cells of the Characeae, which had been detected by Corti and more exactly described by Amici; Dutrochet (Mémoires, I. p. 431) exposed this confusion of ideas, and has the merit of refuting at the same time the absurd theory of the ‘circulation of the vital sap,’ for which Schultz-Schultzenstein had received a prize from the Academy of Paris.
We shall recur in the next chapter to Dutrochet’s minute investigations into the movements connected with irritability in plants, which he also endeavoured to refer to endosmotic changes in the turgidity of the tissues, but he did not do justice to the anatomical conditions of the problem. And here we may take occasion to remark, that Dutrochet’s works were often undervalued, especially in Germany, greatly to the detriment of vegetable physiology. His younger German contemporaries, von Mohl and Schleiden, and at a later time Hofmeister, were right in pointing out what was erroneous and sometimes arbitrary in his mechanical explanations of various movements in plants, and it cannot be denied that he was sometimes led into obscure and doubtful views, as for instance when without any apparent connection he regarded the inhalation of oxygen as a mechanical condition of the rising of the sap and also of heliotropic curvatures, and that his attempts at explanation were not seldom forced and improbable; but all this does not prevent it from being true, that an attentive reader will still gain much instruction from his physiological writings and be excited by them to examine for himself. Dutrochet was a decidedly able man and an independent thinker, who it is true was often led astray by his prejudices, but at the same time manfully protested against the old traditional way of dealing with physiological ideas, and substituted careful examination both of his own and others’ investigations for the accumulation and comfortable retailing of isolated observations which was then the fashion. After de Saussure’s ‘Recherches chimiques’ Dutrochet’s ‘Mémoires pour servir a l’histoire anatomique et physiologique des végétaux et des animaux,’ 1837, are without doubt the best production, which physiological literature has to show in the long period from 1804 to 1840. If later botanists, instead of dwelling on his faults, had developed with care and judgment all that was really good in his general view of vegetable physiology, this branch of botanical science would not have declined as it did in the interval between 1840 and 1860. We shall discover the greatness of Dutrochet as a vegetable physiologist by comparing his work above-mentioned with the best text-books of the subject of the same time, those of De Candolle, Treviranus, and Meyen; not one of them comes up to Dutrochet’s Mémoires in acuteness or depth.