History of Botany (1530-1860)

by Julius Sachs

That plants take up certain substances from their environment for the purpose of building up their own structures could not be a matter of doubt even in the earliest times; it was also obvious, that movements of the nutrient material must be connected with this proceeding. But it was not so easy to say, what was the nature of this food of plants, in what manner it finds its way into and is distributed in them, and what are the forces employed; it was even for a long time undecided, whether the food taken up from without suffers any change inside the plant, before it is applied to purposes of growth. Such were the questions which had engaged the attention of Aristotle, and which formed the chief subject of Cesalpino’s physiological meditations.

But the questions respecting the nutrition of plants acquired a much more definite shape in the latter half of the 17th century, when the various phenomena of vegetation began to be more closely observed, and some attempt was made to understand their relations to the outer world. Malpighi, the founder of phytotomy, was the first who undertook to explain the share which belongs to the different organs of the plant in the whole work of nutrition; guided by analogy, he perceived that the green leaves are the organs which prepare the food, and that the material so prepared by them passes into all parts of the plant, there to be stored up or employed for purposes of growth. But this gave no insight into the nature of the substances from which plants prepare their food. On this point Mariotte endeavoured to give such information as could be obtained from the chemistry of his day; and he has the merit of having shown, in opposition to the old Aristotelian notion, that plants convert the food-material which they derive from the ground into new chemical combinations, while the earth and the water supply the same elements of nutrition to the most different kinds of plants. It could not escape the notice of physiologists even of that time, that the water which plants take up from the ground introduces into them but very small quantities of matter in solution. Van Helmont in the first half of the 17th century had shown this by an experiment, the results of which, however, led him to think that plants were able to produce both the combustible and incombustible parts of their substance from water. Hales at the beginning of the 18th century formed a different opinion, being led by the evolution of the gases in the dry distillation of plants to conclude, that a considerable part of their substance was absorbed in a gaseous form from the atmosphere.

The views propounded by Malpighi, Mariotte, and Hales contained the most important elements of a theory of the nutrition of plants; fully understood they would have taught that one part of the food of plants comes from the earth and the water, and another part from the air; that the leaves change the materials thus obtained in such a manner as to produce from them the substance of plants and to apply this to the purposes of growth; but the ideas were not combined in this way, for during some years after their time botanists were chiefly engaged in observations on the movement of the sap in plants, and they arrived even on this point at very obscure and even contradictory results, because they overlooked the function of the leaves which had already been recognised by Malpighi. All insight not only into the chemical processes in the nutrition of plants, but also into the mechanical laws of the movement of the sap, and generally into the whole internal economy of plants, depends on a knowledge of the fact, that it is only the cells which contain chlorophyll, and therefore in the higher plants the leaves chiefly as consisting largely of such cells, which have the power of converting the gaseous food supplied by the atmosphere into the substance of the plant with the aid of the materials taken up from the soil. This fact is of fundamental importance to the whole theory of the nutrition of plants; it is only by a knowledge of it that we can explain the movement of material connected with nutrition and growth, the dependence of vegetation on light, and to a great extent also the function of the roots.

But this principle could not be discovered till the new chemical system founded by Lavoisier took the place of the old phlogistic chemistry, and it is remarkable that the discoveries, which laid the foundation of modern chemistry in the period between 1760 and 1780, contributed essentially to the establishment at the same time of the modern doctrine of the nutrition of plants. Ingen-Houss, in reliance on Lavoisier’s antiphlogistic views on the composition of air, water, and the mineral acids, succeeded in proving that all parts of plants are continually absorbing oxygen and forming carbon dioxide, but that the green organs at the same time under the influence of light absorb carbon dioxide and exhale oxygen; and as early as 1796 he considered it probable that plants obtain the whole mass of their carbon from the carbon dioxide of the atmosphere. Soon after (1804) de Saussure proved, that plants, while they decompose carbon dioxide, increase in weight by a greater amount than that of the carbon which they retain, and that this is to be explained by the fact that they at the same time fix the elements of water. He likewise showed that the small quantities of saline compounds, which plants take up from the soil, are a necessary part of their food, and that it was at least probable, that the nitrogen of the atmosphere does not contribute to the formation of nitrogenous substances in plants. Senebier had before insisted on the fact, that the decomposition of carbon dioxide under the influence of light only takes place in green organs.

Thus the most important points in the nutrition of plants were discovered by Ingen-Houss, Senebier and de Saussure. But, as often happens in the case of discoveries of such magnitude, their ideas were for a long time exposed to great misunderstanding. They were better appreciated in France than in any other country; Dutrochet and De Candolle were able to see the importance of the interchange of gases in the green organs to the general nutrition and respiration; but others, and especially German botanists, were not content with these simple chemical processes as the foundation of the whole system of nutrition and consequently of the whole life of the plant; the theory of the vital force, which was elaborated in connection with the nature-philosophy during the first years of the 19th century, and was generally accepted by philosophers and physiologists, chemists and physicists, preferred to supply the plant with a mysterious substance for its food, which had its source in the life itself and which it called humus. The most obvious considerations, which must at once have shown that this humus-theory was absurd, were entirely overlooked; and thus in the face of de Saussure’s results the food of plants was once more referred entirely to the soil and the roots, as it was in the earliest times; one of the consequences of this humus-theory in combination with the vital force was that the ash-constituents of plants were supposed to be merely accidental admixtures or stimulants, or to be directly produced in the plant by the vital force.

In the period between 1820 and 1840 the reaction set in from different quarters against the theory of vital force; chemists succeeded in producing by artificial means certain organic compounds, which had hitherto been regarded as products of that force; Dutrochet discovered in endosmose a process, which served to refer various vital phenomena in plants to physico-mechanical principles; de Saussure and others showed that the heat of plants is a product of respiration, and by 1840 the earlier theory of a vital force might be looked upon as antiquated and obsolete. It remained to restore to their rights the observations of Ingen-Houss and de Saussure, which under the influence of that theory and of the notions respecting the humus had been so utterly misconstrued. Liebig set aside the humus-theory in 1840, and referred the carbon of plants entirely to the carbon dioxide of the atmosphere, and their nitrogenous contents to ammonia and its derivatives; he claimed the components of the ash as essential factors in the nutrition, and taking his stand on the general laws of chemistry endeavoured to obtain chiefly by the method of deduction an insight into the chemical processes of assimilation and metabolism. The whole theoretical value of the facts discovered by Ingen-Houss, Senebier and de Saussure was first made apparent by the connection which Liebig succeeded in establishing between the phenomena of nutrition. The doctrine of nutrition burst suddenly into new life; firm ground was gained, and the botanist, no longer distracted by the difficulties raised by the vital force but resting on physical and chemical principles, might now resume the task of investigation. Oxygen-respiration denied by Liebig was first of all re-established by von Mohl and others. Liebig’s views on the source of nitrogen in plants and on the importance of the ash-constituents rested chiefly on general considerations and observations and on calculation, and had now to be tested by systematic investigation and especially by experiments on vegetation in individual plants. And here the place of honour must be assigned to Boussingault, who pursued the path of pure induction as contrasted with Liebig’s deductive mode of proceeding, gradually improved the methods for experimenting on vegetation, and soon succeeded in so producing plants in a purely mineral soil free from all humus, that he finally settled the question of the derivation of the carbon from the atmosphere and of the source of the nitrogen also. He showed from the plants thus artificially nourished, and with due consideration of the many sources of error which beset the question, that the uncombined nitrogen of the atmosphere does not contribute to the nutrition of plants, but that a normal increase in the nitrogenous substances in a plant takes place when the roots take up nitrates as well as the necessary constituents of the ash.

With the exception of some doubts which still remained respecting the necessity of certain constituents of the ash, such as sodium, chlorine and silicic acid, the source of the materials which take a part in the chemistry of the nutrition of plants was known before 1860; but the knowledge obtained with regard to processes in the interior of the plant, the origination of organic substances in the processes of assimilation, and the further changes which they undergo was still fragmentary and uncertain, and led to no general and conclusive results.

1. Cesalpino.

Aristotle had sought to determine the nature of the materials which plants take up as food, and had laid down the proposition, that the food of all organisms is not simple but composed of various substances. This view was correct, but he united with it the erroneous notion, that the food of plants is elaborated beforehand in the earth, as in a stomach, and is made applicable to purposes of growth, so as to exclude the necessity of any separation of excrements in the plant; this error was refuted by Jung, as we shall see, but nevertheless it continued to live as late as into the 18th century, and ultimately quite spoilt Du Hamel’s theory of nutrition.

Cesalpino, whom we have learnt to regard as a faithful and gifted disciple of Aristotle, directed his speculations to the mechanical rather than to the chemical side of the question, and chiefly tried to explain the movement of the nutrient sap in plants. He had a larger stock of material drawn from experience at his disposition than his master, and it is instructive therefore to make a nearer acquaintance with his views, because they show how far the old philosophy was in a condition to turn better empirical knowledge than Aristotle possessed to a satisfactory use; they will also show that Cesalpino’s first essays led him to views which can no longer be said to be strictly Aristotelian.