# History of botany (1530–1860)/Book 2/Chapter 2

CHAPTER II.
Phytotomy in the Eighteenth Century.
Malpighi had no successor of note in Italy; in England the new light was extinguished with Hooke and Grew, and has so remained, we may almost say, till the present day; in Holland also Leeuwenhoek found none to follow him of equal rank with himself, and the work done in Germany up to the year 1770 is more wretched than can well be imagined. There was in fact no original phytotomic research in the first fifty or sixty years of the last century; the accounts which were given of the structure of plants were taken from Malpighi, Grew, and Leeuwenhoek by persons, who, unable to observe themselves, did not understand their authors and stated things not to be found in their writings. The feebler and obscurer notions of the older writers were preserved with a particular preference, and thus it was Grew's complicated idea of the web-like structure of cell-walls that made most impression on those who reported him. This state of decline must not be ascribed to imperfect microscopes only; these certainly were not good, and still less conveniently fitted up; but no one saw and described clearly even what can be seen with the naked eye or with very small magnifying power; the worst part of the case was that no one tried fully to understand either the little he saw himself or the observations to be found in older works, but contented himself from want of reflection with most misty notions of the inner structure of plants. It is not easy to discover the causes of this decline in phytotomy in the first half of the 18th century; but one of the most important appears to lie in the circumstance, that botanists, following in ​this the example of Malpighi and Grew, did not make the knowledge of structure the sole aim in their anatomical investigations, but sought it chiefly for the purpose of explaining physiological processes. The food and circulation of the sap of plants were more and more the prominent questions, and Hales showed how much may be done in this direction even without the microscope; the interest therefore of the few, who like Bonnet and Du Hamel occupied themselves almost entirely with vegetable physiology, was concentrated on experiment.
Others who knew how to use the microscope, as the Baron von Gleichen-Russworm and Koelreuter, were drawn away from the examination of the structure of vegetable organs by their attention to the processes of fertilisation and especially of propagation. The real botanists, according to the ideas of the time, and specially those who belonged to the Linnaean school, considered physiological and anatomical researches generally to be of secondary importance, if not mere trifling, with which an earnest collector had no need to concern himself. That Linnaeus himself thought little of microscopical phytotomy is sufficiently shown by what has been said of him in the first book.
It is not worth while to notice each of the few small treatises on the subject which appeared towards 1760, for they contain nothing new; a few examples will show the truth of the opinion here expressed on the general condition of phytotomy at this time.
We first of all encounter a writer, whom few would expect to find among the phytotomists, the well-known philosopher Christian Baron von Wolff, who in his two works, 'Vernünftige Gedanken von den Wirkungen der Natur,' Magdeburg (1723) and 'Allerhand nützliche Versuche,' Halle (1721) gives here and there descriptions of microscopes and discusses subjects connected with phytotomy. This he does more particularly in the latter work, where he describes a compound microscope with a focussing lens between the objective and the ocular ​but without a mirror, an instrument which must have served therefore for observing with the light from above on an opaque surface; the objective was a simple lens. But to magnify objects more strongly, he says that he used a simple instead of this compound instrument, as was more the custom at the time. Like a true amateur Wolff submitted all sorts of small and delicate objects to his glass, without examining any of them thoroughly and persistently. His phytotomic gains were small; he observed for instance that starch-flour (powder) consists of grains, but believed from the way in which they refracted light that they were small vesicles filled with a fluid; yet he satisfied himself that these grains are already in the grains of rye and therefore not produced in the grinding. He laid thin sections of portions of plants on glass which was too imperfectly polished to allow of his seeing anything distinctly. His pupil Thümmig in his 'Meletemata' (1736) addressed himself to the subject with still less skill. By the case of these two men we may see plainly that want of success was due much less to the imperfectness of the microscope than to unskilful management and unsuitable preparation. But Wolff and Thümmig at least endeavoured to see something for themselves of the structure of plants; a famous botanist of the time, Ludwig, plainly never made a similar attempt, for in his 'Institutiones regni vegetabilis' (1742) he speaks of the inner structure of the plant in the following manner; 'Laminae or membranous pellicles, so connected together that they form little cavities or small cells and often reticulated by the intervention of fine threads, form the cell-tissue which we see pervading all parts of plants. These are what Malpighi and others call tubes, since they appear in different parts in the form of rows of connected vesicles! 'Boehmer's 'Dissertatio de cellulose contextu' (1785) is still worse; 'White elastic thicker or thinner fibres and threads woven together of differing shape and size form cavities or cells or caverns, and are usually known by the name of cell-tissue.' We see ​what mischief Grew did with his theory of the fibrous structure of the cell-walls, and how the expression cell-tissue literally taken led the botanists here named and others into utterly incorrect ideas. The works of Du Hamel, Comparetti, and Senebier show that such misconceptions were not confined to Germany, and Hill, a countryman of Grew, according to von Mohl's account pictured to himself cells as cups standing one above another, closed below and open above.
Baron von Gleichen-Russworm (1717-1783), privy counsellor to the Margrave of Anspach, gave much attention to the perfecting of the mechanical arrangements of the microscope, but his plates themselves show how strangely unsuitable these arrangements were. With these instruments he made many observations, which are recorded in two voluminous works, 'Das Neueste aus dem Reich der Pflanzen' (1764) and 'Auserlesene mikroskopische Entdeckungen' (1777–1781). But these works contain little or nothing about microscopic anatomy or the structure of vegetable cells. His observations with the microscope are chiefly devoted to processes of fertilisation and to proving that spermatozoa are contained in the pollen and in connection with these subjects he gives magnified figures of many small flowers, some of them beautifully executed; these figures must have made his works very instructive to many in their time. He saw the stomata, which Grew had already discovered, on the leaves of ferns, but took them for the male organs of fertilisation, which at the same time showed that he was still unacquainted with the existence of stomata in phanerogams.
Caspar Friedrich Wolff in his efforts in phytotomy stands ​a solitary figure among his contemporaries, not only because he was the first since Malpighi and Grew who devoted labour and perseverance to the study of the anatomy of plants, but still more because at a time, when the structure even of matured vegetable organs was almost a forgotten subject, he endeavoured to penetrate into the history of the development of this structure and the formation of cellular tissue. Unfortunately he was not directed to this by an exclusive interest in phytotomy, but by a more general question which he endeavoured to set at rest in this manner; he wished to refute the prevailing theory of evolution by demonstrating the development of the organs of plants, and to obtain an inductive basis for his doctrine of epigenesis. Though he was often diverted by these means from the pursuit of purely phytotomic questions, yet his famous work, 'Theoria Generationis' (1759) is nevertheless important in the history of phytotomy; for though it was disregarded by botanists during the succeeding forty years, or at any rate exercised no noticeable influence, yet it was Wolff's doctrine of the formation of cellular structure in plants which was in the main adopted by Mirbel at the beginning of the present century, and the opposition which it encountered contributed essentially to the further advance of phytotomy. This late but lasting influence of Caspar Friedrich Wolff's work was due not to the actual correctness but to the thoughtfulness of his observations, and to the earnest desire which inspired them to ​search out the true nature of vegetable cell-structure and to explain it on physical and philosophical grounds. The observations themselves on this point are highly inexact, and influenced by preconceived opinions, and his account of them is rendered obscure and often quite intolerable by his eagerness to give an immediate philosophic explanation of objects which he had only imperfectly examined. His efforts to follow the course of development in the first beginnings of the formation of cell-tissue were evidently not seconded by sufficient knowledge of the structure of matured organs, and, to judge by his figures and by his theoretical reflections, his microscope was of insufficient power and its definition imperfect. Notwithstanding all these deficiencies, Wolff's treatise is doubtless the most important work on phytotomy that appeared in the period between Grew and Mirbel, not, as has been said, on account of any particular excellence of observation, but because its author was able to make some use of what he saw, and to found a theory upon it.
According to that theory all the youngest parts of plants, the punctum vegetationis in the stem, which Wolff first distinguished, the youngest leaves and parts of the flower, consist of a transparent gelatinous substance; this is saturated with nutrient sap, which is secreted at first in very small drops (we might say vacuoles), and these, as they gradually gain in circumference, expand the intermediate substance and so present enlarged cell-spaces. The intermediate substance therefore answers to what we should now call the cell-walls, only these are at first much thicker, and are constantly becoming thinner with the growth of the cell-spaces. We may compare young vegetable tissue, formed as Wolff imagines, with the porosity of fermenting dough, except that the pores are not filled with gas but with a fluid. It is plain from the above description that the vesicles or pores, as Wolff names the cells, are connected together from the first by the intermediate substance, and that one lamina or cell-membrane ​only lies between each of two adjoining cells, a point which succeeding phytotomists were a long time in determining. As cells are formed by the secretion of drops of sap in the fundamental substance which is at first homogeneous, so vessels, according to Wolff, are produced by longitudinal extension of a drop in the mucilage and formation of a canal; consequently adjoining vessels must be separated from one another by a single lamina of the fundamental substance. Though Wolff expressly mentions the movement of the sap within the firm mucilaginous substance between the cellular cavities and the vascular canals, a movement of diffusion as it might now be termed, he inconsistently enough thinks it necessary to assume the existence of perforations in the bounding-walls of cells and vessels to serve for the movement of sap from cell to cell and vessel to vessel; yet in the single case in which he succeeded in obtaining isolated cells, namely in ripe fruits, he was obliged to allow that the walls were closed.
The growth of the parts of plants, according to Wolff, is effected by expansion of existing cells and vessels, and by the formation of new ones between them in the same way as the first vacuoles were formed in the mucilaginous substance of very young organs; that is to say, the sap which saturates the solid substance between the passages and cavities of the tissue separates in the form of drops, which increase in size and then appear as cells and vessels introduced between the older ones. The substance between the passages and cavities, at first soft and extensible, becomes firmer and harder with increasing age, and at the same time a hardening substance may be deposited on it from the sap which is stagnant in the cell-cavities and in movement in the vascular passages, and this substance in many cases appears as their proper membrane.
This is in all essential points Wolff's theory. We may omit his statements on the subject of the first formation of leaves at the growing point and of the development of the parts of the flower, as well as his physiological views on food and sexuality, ​which remained for a long time without influence on the growth of opinion, and mention only his doctrine of the growth of thickness of the stem. The stem is originally the prolongation of all the leaf-stalks united together. As many bundles of vessels are formed in the developed stem as there are leaves springing from the vegetative axis; each leaf has a single vascular bundle belonging to it in the stem, in modern phraseology an inner leaf-trace. The union of these bundles from the different leaves forms the rind of the stem; but if the leaves are very numerous, their descending bundles form a closed cylinder, and if the stem is perennial, the fresh production of leaves every year produces new zones of wood of this kind every year, which are the yearly rings. This view of Wolff's on the growth of the stem in thickness bears an unmistakable resemblance to the theory afterwards suggested by Du Petit-Thouars, according to which the roots which descend from the buds are supposed to effect the thickening of the stem.
The contests between Mirbel and his German antagonists at the beginning of the present century will bring us back again to the more important points in Wolff's theory of the cell. Contemporary botanists paid less attention to the 'Theoria Generationis' than they did to Hedwig's phytotomic views, not on the formation of cells, but on the structure of mature tissue. Hedwig had given various figures and descriptions of phytotomic subjects in his 'Fundamentum Historiae Muscorum' (1782) and afterwards in his 'Theoria Generationis' ​(1784); but he treats these topics at greater length in his treatise 'De fibrae vegetabilis et animalis ortu,' published in 1789, and known to the author of this work only imperfectly from quotations in later writers. Hedvvig's figures of histological objects appear to be better than those of any of his predecessors; they show that he used strong magnifying powers, and that his glass had a clear field of sight. His defect lay in preconceived opinions and hasty interpretation of what he observed. In order to refute Gleichen's view of the stomata in ferns, he demonstrated the existence of these organs in many phanerogams, and observed the opening of the slits, which he named 'spiracula.' On the epidermis which he had removed for the purpose of these observations he saw plainly the double contour lines bounding the epidermis-cells, and therefore the cell-walls, which are at right angles to the surface. These he took for a particular form of vessel, and called them 'vasa reducentia' or 'lymphatica,' and afterwards 'vasa exhalantia,' and he thought that he had found them again in the interior of parenchymatous tissue, evidently taking the places where three wall-surfaces meet for vessels; such vessels he also saw in the milk-cells of Asclepias, described in 1779 by the elder Moldenhawer, who seems himself to have regarded even the intercellular spaces in the pith of the rose as equivalent to these milk-cells. The word vessel even in the 18th century was used in such an indefinite manner, that the broad air-tubes of the wood and the finest fibres were called vessels. Hedwig's idea of the construction of spiral vessels was strange enough; he took the spiral band itself for the vessel, and supposed it to be hollow because it is coloured by absorption of coloured fluids; in those spiral vessels in which the turns of the spiral band are distant he saw, it is true, the delicate original membrane which lies between the turns, but he supposed that it lay inside the spiral band, which was wound round it on the outside. On the second plate of the first part of the 'Historia Muscorum' he even figures the network of ridges which the ​adjoining cells have left on the wall of the spiral vessel, but explains it as wrinkles caused by desiccation.
Hedwig was without doubt a very practised microscopist, and he constantly recommended the extremest care in the interpretation of all that the instrument reveals; but if an observer so careful and practised, who moreover was provided with a glass of tolerably strong magnifying power, fell into such gross mistakes, it cannot surprise us if others, as P. Schrank, Medicus, Brunn, and Senebier, accomplished still less. These highly unimportant achievements are all that mark the close of the 18th century.
* ↑ This subject will be noticed again in the history of the sexual theory.
* ↑ C. F. Wolff was born at Berlin in 1733. He studied anatomy under Meckel and botany under Gleditsch, in the Collegium Medico-chirurgicum in that city. He afterwards resorted to the University of Halle, and there made acquaintance with the philosophy of Leibnitz and Wolff, which predominates too much in his dissertation, 'Theoria Generationis' (1759). Haller, the representative of the theory of evolution against which this work was directed, replied to it in a kindly spirit and entered into a correspondence with its youthful author. After lecturing on medicine in Breslau, he was admitted to teach physiology and other subjects in the Collegium Medico-chirurgicum in Berlin, but was twice passed over in the appointment to professorships in that institution. He received an appointment in the Academy of St. Petersburg from the Empress Catherine II in 1766, and died in that city in 1794. See Alf. Kirchhoff, 'Idee der Pflanzenmetamorphose,' Berlin, 1867.
* ↑ Johannes Hedwig, the founder of the scientific knowledge of the Mosses, was born at Kronstadt in Siebenbürgen in 1730. Having completed his studies at Leipsic, he returned to his native town, but was not permitted to practice there as a physician because he had not taken a degree in Austria. He consequently went back to Saxony and settled first at Chemnitz, and in 1781 in Leipsic. Here he was appointed in 1784 to the Military Hospital, and became Professor extraordinary of Medicine in 1786 and ordinary Professor of Botany in 1789. He died 1799. He commenced his botanical studies as a student at the University, and continued them in Chemnitz under trying circumstances, till as Professor he was free to devote himself entirely to them.