History of Botany (1530-1860)

by Julius Sachs

For the purpose of these observations he looked about in every case for the most suitable objects for examination, a point to which former phytotomists, with the exception of Moldenhawer, had paid little attention; he then studied these objects thoroughly, and thus prepared the way for the examination of others, which presented greater difficulties. Every monograph of this kind was a nucleus, round which a larger number of observations might afterwards gather. In a long series of such solid productions he treated conclusively all the more important questions of phytotomy.

Von Mohl’s extraordinary carefulness was not however able to guard him, calm observer though he was, from some serious mistakes, at least in his earlier years, such as those which occur in his first theory of intercellular substance (1836), and in his earliest views on the nature of the cell-membrane of the pollen-grain (1834). These and some other errors on the part of a gifted and truly inductive enquirer are instructive, since they show that observation without any groundwork of theory is psychologically impossible; it is a delusion to suppose that an observer can take the phenomena into himself as photographic paper takes the picture; the sense-perception encounters views already formed by the observer, preconceived opinions with which the perception involuntarily associates itself. The only means of escaping errors thus produced lies in having a distinct consciousness of these prepossessions, testing their logical applicability and distinctly defining them. When von Mohl laid down his theory of intercellular substance, there evidently floated before his mind indistinct, half-conscious ideas of the kind that Wolff and Mirbel entertained of the structure of the vegetable cell; and as he considered the cell-membrane of the pollen-grain to consist of a cell-layer, he summarised its obscure structural relations under the then very obscure conception of the cell. As a true investigator of nature, who adheres always and firmly to the results of further observation, and endeavours to clear his ideas by their aid, conceding only a relative value to every view, von Mohl soon escaped from these errors, and himself supplied proofs of the incorrectness of his former opinion. The number of really erroneous statements in his works is wonderfully small considering the very large number of investigations in which he engaged.

In examining the part which von Mohl played in the general development of phytotomy we can distinguish satisfactorily two periods in his scientific career, the first of which extends from 1827 to about 1845. Before 1845 he was acknowledged to be the first of phytotomists, decidedly superior to all rivals; his authority, though often attacked by unimportant persons, grew from year to year. This period may be said to close with the publication of his ‘Vermischte Schriften’ in 1845. Up to that time investigations into the form of the solid framework of cell-membrane had chiefly attracted the interest of phytotomists, and in this subject there was no one who could measure himself with von Mohl. Yet he began soon after 1830 to study the history of development in plants; in 1833 he described the development of spores in a great variety of Cryptogams, in 1835 the multiplication of cells by division in an alga, and the cell-division in the formation of stomata in 1838; in this period appeared Mirbel’s first observations on the formation of pollen-cells (1833). Von Mohl too was the first, if we disregard Treviranus’ somewhat imperfect notices of the origin of vessels in 1806 and 1811, who explained the history of the development of those organs; and his theory of the thickening of cell-membranes, the principles of which are to be found in his treatise on the pores in cellular tissue (1828), may also be regarded as a mode of conceiving the sculpture of the cell-membrane from the point of view of the history of development.

Ever since 1838 Schleiden had raised the history of development to the first rank in botanical investigation, but he had proposed a thoroughly faulty theory of cell-formation, to which von Mohl at first at least did not withhold his assent in spite of previous and much better observations; but after 1842 Nägeli devoted himself still more thoroughly and with more lasting results to the study of the development both of vegetable cells and tissue-systems, and of the external organs. He introduced new elements into phytotomic research, and it soon became apparent that even the questions hitherto examined must be grappled with in a different fashion. Von Mohl did not hold aloof from the new direction, but completed a series of excellent investigations connected with the new questions in the theory of cell-formation. The most important of these was his enquiry into the nature of protoplasm, to which he gave the name still in use. In his treatise, ‘Die Vegetabilische Zelle,’ which came out in 1851 in Wagner’s Dictionary of Physiology, he even gave an excellent account of the modern theory of cell-formation; but notwithstanding all this, and the great authority which he rightly continued to enjoy, he was no longer the guide who led the way in the domain of phytotomy, as he had been before 1845.

His zeal as an observer had at all times been chiefly attracted to the solid framework of vegetable structure in its matured condition, though a number of his most important works were devoted to the study of cell-contents.

Except in his ‘Anatomic der Palmen’ (1831), where he expended much and to some extent even unnecessary labour on figures representing the general appearance of the tissue (histologic habit), von Mohl’s microscopic drawings do not aim at giving the collective impression, but at facilitating the understanding of the delicate structure of single cells and their combinations by aid of the simplest possible lines. He always despised pictures from the microscope, such as were introduced at a later time by Schacht,—a kind of artistic restoration of the originals and to some extent a playing with science; and in his later publications he was more sparing of illustrations or omitted them altogether, in proportion as he acquired the power of giving clear verbal explanations of even difficult structural conditions.

Von Mohl’s scientific activity was so wonderfully productive that it is not easy to present the reader with a clear account of it; but we must endeavour at least to furnish such a summary of its chief results as may serve to give a general idea of his importance in the history of our science. We may here pass over such of his treatises as do not bear on the main questions of phytotomy, and notice only those that relate to the structure of the solid framework of plants, because the historical significance of his investigations into the history of development can only be understood in connection with the questions to be treated in the following chapter. But we shall not limit ourselves to publications which appeared before 1845, though we may be thus compelled to notice researches which in succession of time belong to the next period, and indeed almost to the present moment.

1. The view that the cell is the sole and fundamental element in vegetable structure had been already maintained by Sprengel and Mirbel, but not supported by exact observations. Treviranus too had shown that the vessels in wood are formed by the union of rows of cell-like tubes, but he had never arrived at a thoroughly clear conception of the matter. On the one side was the theory that the plant consists entirely of cells, on the other, and for long the old and strange view, that the spiral thread was an independent elementary organ of vegetable structure,—a view which Meyen still maintained in 1830. Von Mohl must be regarded as the first who took up the all-important position, that not only the fibrous elements of bast and wood, which had long been considered to be elongated cells, but the vessels of the wood also are formed from cells; and we may on this point give great weight to his own assertion that he was the first who observed the formation of vessels from rows of closed cells. This discovery happened in the year 1831, and he describes distinctly, though briefly, the decisive observations in his treatise on the structure of the palm-stem. At the points of constriction in the vessels he saw the dividing walls, the existence of which had been denied by all former phytotomists; ‘these dividing walls,’ he says, ‘are entirely different from the rest of the membranes of the plant, being formed of a network of thick fibres with openings between them.’ He studied the history of the development of these vessels both in palms and in dicotyledonous plants. ‘In the young shoot,’ he says, ‘are found at the spots, where afterwards there are large vessels, perfectly closed large cylindrical tubes with a transparent and very delicate membrane.’ He then shows how by degrees the sculpture peculiar to the walls of vessels is formed on the inside of these tubes, and he takes the opportunity of saying that a metamorphosis in time from one form of vessel into another is entirely out of the question, as Treviranus also and Bernhardi had maintained. ‘The dividing walls (transverse septa),’ he continues, ‘are formed in a precisely similar manner to the side-walls of vessels; only the original tender membrane of the septa is usually lost in the meshes of the network of fibres.’ From that time no phytotomist capable of an independent judgment has had any doubt with regard to this view of the formation of the vessels in wood. It is however striking enough that von Mohl, who thought it so important to show that the cell is the sole foundation of vegetable structure, never extended the proof to milk-vessels and other secretion-canals in order to show whether and how these also are formed from the cells. In his treatise on the vegetable cell (1851) he still expressed doubt about Unger’s assertion, that the milk-vessels are also formed from rows of cells that coalesce with one another, and held rather to the view of an anonymous writer in the ‘Botanische Zeitung’ of 1846, page 833, that these vessels are membranous linings of gaps in the cell-tissue. He might well lose his taste for the examination of these and similar organs after Schultz Schultzenstein had by his various treatises, written after 1824, on the so-called vital sap and the circulation which he attributed to it, made this part of phytotomy a very quagmire of error, and had not refrained from replying in an unbecoming manner to von Mohl, who repeatedly opposed his views; moreover Schultz’s essay ‘Ueber die Circulation des Lebenssafter’ (1833), which teems with absurdities, had received a prize from the Academy of Paris.

2. The growth in thickness of the cell-membrane, and the sculpture caused by it was a subject that is more or less connected with most of von Mohl’s writings. He developed the chief features of his view in 1828 in his first work, ‘Die Poren des Pflanzengewebes!’ The way in which he represented to himself the growth in thickness of cell-membranes at a later time may be expressed as follows. All elementary organs of a plant are originally very thin-walled perfectly closed cells, which in the tissue are separated by walls formed of two laminae[82]; on the inside of these primary cell-membranes, after they have ceased to increase in circumference, new layers of membranous substance are formed, which lying one upon another adhere closely together, and represent the whole amount of secondary thickening layers; on the inner side of the membrane thus thickened by apposition there may usually[83] be perceived a tertiary layer of a different character.

But there are certain sharply defined spots on the original cell-wall, where this thickening does not take place; in such spots the cell is still bounded only by the primary membrane; it is these thin spots which bear the name of pits, and which Mirbel, and in some cases Moldenhawer, took for holes, but von Mohl considered that it was only in very exceptional cases that they were really changed into holes by resorption of the thin primary wall. In accordance with this theory, the spiral, annular, and reticulated vessels are produced by deposition of thickening matter in the form suitable to each case on the inside of the originally smooth thin cell-wall. But like Schleiden and other phytotomists, von Mohl was not quite clear in his views either of the origin or mode of formation of matured bordered pits; it was supposed that the two laminae of the dividing wall parted from one another at certain spots in such a manner that a lenticular hollow space was formed between them, and that this space answered to the outer border of the pit, while the inner border was the result of ordinary pit-formation. This view, which could be shown to be incorrect by the history of development, arose in fact from inexact observation,—a rare case with von Mohl; the true course of events in the formation of bordered pits was first described by Schacht in 1860.