History of botany (1530–1860)/Book 2/Chapter 3

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Examination of the Matured Framework of Cell-membrane in Plants.


There is no sharp line of division between the 18th and the 19th centuries; the phytotomists who appear on the scene during the first years of the new century are scarcely more successful than Hedwig and Wolff; careful and judicious interpretation of their own and others' observations is still rare, and they are often misled by preconceived opinions.

In one respect indeed a very great improvement appeared with the commencement of the 19th century; the number of phytotomists working contemporaneously, checking and criticising one another, became all at once much larger. Hitherto ten or twenty years had intervened between every two works on phytotomy; but in the course of the twelve years after 1800 nearly as many publications followed one another, and scientific discussion enlivened enquiry. Now we meet with a Frenchman for the first time in the field of phytotomy, Brisseau Mirbel, who brought out his 'Traité d' Anatomie et de Physiologic Végétale' in 1802, and raised a series of questions in the discussion of which several German botanists, Kurt Sprengel (1802), Bernhardi (1805), Treviranus (1806), Link and Rudolphi (1807), at once took part. It was a step in advance and one affecting all botanical studies, that with the exception of Rudolphi all these men, like Hedwig before them, were botanists by profession; it was at last felt that the examination of the internal structure of plants, as well as the describing them according to Linnaean patterns, was a part of botanical enquiry; it is at the same time true that the botanical knowledge of these observers was often of service to them in their phytotomical investigations, and directed their work decidedly and from the first towards that which was worth knowing, and towards the objects which claimed the first attention. This remark applies to the younger Moldenhawer even more than to the botanists above-named; his 'Beitrage,' published in 1812, may be taken as closing the first section of this century, during which time he improved the methods of observation, compared his own observations and those of others with great acuteness of judgment, and did all that could be expected with the microscopes of the time.

The period of sixteen years after Moldenhawer, from 1812 to 1828, has nothing of material importance to show in phytotomy. On the other hand, it produced a series of the most important improvements that the compound microscope has undergone since its invention.

As early as 1784 Aepinus had produced objectives of flint and crown glass, and in 1807 Van Deyl[1] made similar ones with two achromatic lenses, and still the phytotomists complained of the condition of their instruments. Their figures show that they could not see clearly with them, though the magnifying powers were not high; Link says expressly in the preface to his prize-essay of 1807, that he usually observed with a lens that magnified a hundred and eighty times. Moldenhawer in 1812 gives the preference over all the microscopes he had used to one by Wright, which was serviceable with a magnifying power of four hundred times, while the German instruments, especially those by Weickert, could not be used with higher powers than from one hundred and seventy to three hundred.

A certain interval elapsed each time between an improvement in the instrument and the appearance of the advantages which phytotomy derived from it; thus in 1824, Selligue exhibited to the Academy of Paris an excellent microscope with double lenses, several of which could be screwed on one over the other, and which could be used with ordinary daylight and a magnifying power of five hundred times; in 1827 Amici made the first achromatic and aplanatic objectives with three double lenses screwed on one over the other, the flat sides being turned to the object. And yet still in 1836 a practised phytotomist like Meyen spoke with disapproval of the instruments of his time, and gave the preference to an old English microscope by James Man, though he allowed that the newest instruments by Ploessl were a little better. In his work on phytotomy, which appeared in 1830, all the figures were magnified two hundred and twenty times, as were the very beautiful figures in his prize essay of 1836; but in his 'Neues System' (1837), he had already adopted powers that magnified to over five hundred times. How rapid the progress was in the years before and after 1830 is shown by comparing von Mohl's work on climbing plants of 1827 and its antiquated illustrations, with his publications of 1831 and 1833, where the figures have a thoroughly modern appearance.

The art also of preparing anatomical objects rose by degrees with the improvement of the microscope. It was not in a very advanced state at the beginning of the century, if we judge by the language of writers and by their figures. It was a great step in advance when the younger Moldenhawer in 1812 isolated cells by maceration and decay in water, and was thus enabled to view cells and vessels on every side and in a perfect condition, to see their real shape, and to survey the manner of their combination more exactly than had hitherto been done. But even Moldenhawer still made the mistake of submitting delicate microscopic objects to observation in a dry state, though Rudolphi and Link in 1807 had urged the advisability of keeping every part of the preparations moist, especially the surface towards the object-glass, which shows that they did not then use covering glasses. Nor was sufficient attention shown to the use of sharp knives of suitable form, such as the razor, which is now almost exclusively employed, or to practice in making transverse and longitudinal sections of the utmost possible delicacy, two things which, through the example of Meyen's and von Mohl's practice, were afterwards recognised as indispensable helps to phytotomy; even in their time observers were satisfied with crushing and picking their preparations to pieces.

Drawing from the microscope kept even pace on the whole with increasing skill in making preparations, and with the improvement of the instrument. If we compare together the drawings of Mirbel and Kurt Sprengel in the beginning of the century, those of Link and Treviranus in 1807, Moldenhawer's in 1812, and Meyen's and von Mohl's from 1827 to 1840, we shall obtain a rapid and instructive survey of the history of phytotomy during this period of forty years. The figures testify at once to constant increase in the magnifying powers, to the greater clearness of the field of sight, and still more to the constant improvement in the arts of preparing and observing objects. But a curious misconception crept in among the phytotomists at this time; they believed that more correct and trustworthy figures would be obtained, if the observer and writer did not himself make them, but employed other eyes and other hands for that purpose; they imagined that in this way every kind of prejudice,of preconceived opinion would be eliminated from the drawings. Thus both Mirbel and Moldenhawer had their figures drawn by a woman, and many later phytotomists entrusted the execution of their drawings to hired draughtsmen, as Leeuwenhoek had done before them. A drawing from the microscope, like every other copy of an object in natural history, cannot pretend to take the place of the object itself, but is intended to give an exact and clear rendering of what the observer has perceived, and by so doing illustrate the verbal description. The drawing will be perfect in proportion to the practised skill of the eye that observes and of the mind that interprets the forms. The copy should only show to another person what has passed through the mind of the observer, for then only can it serve the purpose of a mutual understanding. There is also another point to be considered; it is exactly in the process of drawing a microscopic object that the eye is compelled to dwell on the individual lines and points and to grasp their true connection in all dimensions of space; it will often happen that in this process relations will be perceived, which previous careful observation had disregarded, and which may be decisive of the question under examination or even open up new ones. As the microscope trains the eye to scientific sight, so the careful drawing of objects makes the educated eye become the watchful adviser of the investigating mind; but this advantage is lost to the observer who has his drawings made by another hand. It is not one of the least of von Mohl's merits, that he practised microscopic drawing under the influence of the views here indicated, and sought to make his figures no mere undigested copies of the objects, but an expression of his own opinions about them.

Enough has been said to show that an important portion of the history of phytotomy lies between the beginning and the end of the period under consideration. The distance between the knowledge of the structure of vegetable tissue which existed at the beginning of the century, and that of Meyen and von Mohl on the same subject in 1840, is wonderfully great; in the one case an uncertain groping about among obscure ideas, in the other a complete exposition of the inner architecture of the mature plant. But in spite of this great difference between beginning and end, it is better to review the efforts of this period of forty years as a connected process of historical development, and, notwithstanding the interval between the appearance of Moldenhawer's contributions in 1812 and Meyen's and von Mohl's labours about 1840, to consider the latter as the settlement of the questions taken up at the commencement of the century. Moreover after 1840, with the appearance of Schleiden and Nägeli on the scene, new points of view were suddenly disclosed, and new aims were proposed in phytotomic investigation; it is no objection to this view of the subject, that the most productive portion of von Mohl's labours falls in the succeeding twenty years, and that during this later period his position is one of equal authority with the new tendency and of participation in it. Up to 1845 his discoveries were the culminating point of the older phytotomy; they put the finishing stroke to the work which Mirbel, Link, Treviranus, and Moldenhawer had begun. The object almost exclusively pursued during all this period was to frame as true a scheme as possible of the inner structure of the mature organs of the plant; it was requisite to gain a right understanding of the diversities of cells and forms of tissues, to classify them and supply them with names, and to secure well-conceived definitions of these names. Hence almost exclusive attention was paid to the configuration of the solid framework of cell-membrane, and of this chiefly in the matured state, to the form of the several elementary organs and their combination in the tissue, to the sculpture of the wall-surfaces, and to the connection of cell-spaces by pores or their separation by closed walls. There was much discussion, especially at first, on the contents of vessels and cells, and on supposed movements of sap in connection with anatomical research, but there was no careful connected investigation of the cell-contents; it was not yet recognised that the true living body of the vegetable cell is only a definite part of the contents inclosed by the cell-wall; the solid walls, the framework of the whole building, were regarded as of primary importance in the structure of the cell. It was not till the following period that in the light of historical development another view asserted itself, namely, that the solid framework of vegetable tissue with all its importance is yet in the genetic sense only a secondary product of the phenomena of vegetative life, that the true cell-body, the cell=protoplasm is prior in time and in conception, and can claim the higher position.

Mirbel, to whom we now return, had in 1801 laid down a theory of cell-formation which agreed in the main with that of Caspar Friedrich Wolff; he supposed with Wolff that each cell-space was separated from its neighbour by a single wall, and relying on fresh observations asserted the existence of visible pores in the dividing walls of parenchyma and of vessels, and also maintained some new views on the nature and formation of vessels. The essential points of this theory found an opponent in Germany in the person of Kurt Sprengel, the well-known historian of botany, and one of the most variously accomplished botanists of his time, who had published in 1802 a work written in diffuse and familiar style under the title of 'Anleitung zur Kenntniss der Gewächse.' He relied on his own observations, but these were evidently made with small magnifying powers, an obscure field of sight, and indifferent preparations. The cell-tissue, says Sprengel, consists of cavities of very various shape communicating with one another, the dividing walls being in some places broken through and in others wanting. He took the starch-granules which he saw in the seed-leaves of beans and other plants for vesicles, which increase in size by absorption of water and so form new tissue; but he did not explain how we are to conceive of the growth of organs with such a mode of cell-formation. His account of the vessels is extremely obscure, even more obscure than Hedwig's, though he has the merit of refuting the latter's strange theory of reconducting vessels in the epidermis; he also suggested, though only incidentally, the happy idea that spiral passages and even vessels might arise from cell-tissue, since the youngest parts of plants have only the latter; but he did not attempt to explain how or where the process takes place. Like Malpighi and Grew he supposed that the spiral vessels had no wall of their own, but thought that the closely-rolled spiral threads formed a wall; the constrictions in broad short-membered vessels he regarded as real contractions in their substance, caused by the increased tightening of the spiral threads through a sort of peristaltic movement, a mistaken notion often entertained at the beginning of the century, by Goethe among others, and connected with ideas of vital power prevalent at the time. In the stomata, to which he gave the name still in use, Sprengel like Grew, Gleichen, and Hedwig, saw a circular cushion instead of the two guard-cells; but he notices the observation first made by Comparetti, that the orifice closes and opens alternately, being wide open in the morning and closed in the evening. But he considered the stomata to be organs of absorption.

Sprengel in enunciating his own theory of cell-formation accused Mirbel of mistaking the starch-grains in the cells for the pores of the cell-walls. On this point, so important in the doctrine of the cell and in physiology, he was followed by the three candidates for the Gottingen prize, though Bernhardi had in 1805 defended Mirbel's view, and had pointed out how little likely it was, that so skilful an observer as Mirbel should fall into so gross an error. Bernhardi's short treatise, 'Beobachtungen über Pflanzengefässe,' Erfurt (1805[2]'), was in general distinguished by a variety of new and correct observations, and was the work of a simple and straightforward understanding, which takes things as they are presented to the eye without allowing itself to be led astray by preconceived opinions. His observations are certainly the best in the whole period from Malpighi and Grew to the younger Moldenhawer; his method of dealing with questions of phytotomy is much more to the purpose than that of the three rivals for the Gottingen prize.

In the work just mentioned Bernhardi treats of other forms of tissue as well as vessels, and endeavours to distinguish and classify them more exactly than had hitherto been done. He contrasts favourably with his contemporaries in the fact, that he sought to define the histological terms employed as precisely as possible, a great step in advance at a time when phytotomic conceptions were in a very misty condition. He distinguishes three chief forms of vegetable tissue, pith, bast, and vessels.

By pith he means the tissue which Grew had named parenchyma, and which is still so called; it remained a question with him whether the cells of the pith are pierced by visible pores. By the word bast he understood not only the fibrous elements of the rind, but those of the wood also, and in general what is now known as prosenchyma; this agrees very well with Malpighi's view, which was adopted by Bernhardi and by all his contemporaries, that the inner layers of the bast are changed into the exterior layers of wood to make the increase in thickness of the woody stem; but he did not admit the same origin in the case of the innermost portion of the wood, for this is formed from the first in the young shoots, which alone contain true spiral vessels with threads that may be wound off.

Bernhardi distinguishes vessels into two main groups, air-vessels and vessels properly so called. He calls the first group air-vessels for the same reason that led Grew to give them that name, namely, that they are filled with air during a part at least of the period of vegetation; they are found in the wood, and, where there is no closed woody body, there the woody bundles are formed both of vessels and also of bast strands which enclose vascular canals. These latter he next divides into three chief kinds; annular vessels, which he was the first to discover, true spiral vessels with a band which can be unwound, and scalariform vessels, by which term he understood not only those with broad slits, such as are found in Ferns, but also the pitted vessels in secondary wood. His idea of annular and spiral vessels was perfectly correct, and he mentions Hedwig's notion already described, and shows that its exact opposite is true, namely, that the spiral band is surrounded by a membrane on the outside, a fact which was afterwards denied by Link, Sprengel, and Moldenhawer. On the other hand he did not understand the sculpturing on the scalariform vessels; he took the pits in the dotted vessels for thickenings of the wall, such as are seen in the transverse ridges between the slits in true scalariform vessels, and the slits he thought were closed. If there was much that was erroneous in these views, yet Bernhardi contributed essentially to the clearing up of the subject by his effort to distinguish the different forms of air-vessels, and especially by pointing attention to the fact that neither spiral nor annular vessels are found in secondary wood. The resemblance between different forms of vessels misled many of his contemporaries into supposing that they are due to metamorphosis of true spiral vessels. Bernhardi showed that different forms of wall are found inside one vascular tube, but that this does not depend on modification with age; observation rather teaches that every kind of vessel receives its character in its young state, and especially that the youngest scalariform vessels do not present the form of spiral vessels.

Under the head of vessels proper he reckoned all tubular forms filled with a peculiar juice, milk-cells and true milk-vessels, and also resin-ducts and the like, and he made many good and still valuable observations on their distribution and sap-contents. He could not see the differences of structure in these various fluid-conveying vessels with the low magnifying power of his glass, and therefore attended chiefly to the structure of the large resin-ducts, which on the whole he rightly understood.

The question whether there are any other forms of vessels in the plant beside those already named gave him occasion to define a vessel better than it had yet been defined, namely as an uninterrupted tube or canal, and at the same time he found himself obliged to consider whether his bast-threads are vessels; but he did not give a decided answer to the question. He declared however distinctly against Hedwig's reconducting vessels in the epidermis, as Sprengel had done, and it is worthy of recognition that he understood the true nature of the corners where three longitudinal walls of the parenchyma meet, while later observers found difficulties in them.

Before the appearance of Bernhardi's work the Royal scientific Society of Göttingen proposed a subject for a prize in the year 1804, which shows very plainly what uncertainty was felt at that time on all points of phytotomy. For this reason it will be well to give it at length from the preface to Rudolphi's 'Anatomic der Pflanzen' (1807): 'Since some modern physiologists deny the peculiar construction of vessels in plants which is attributed to them by other and especially the older observers, it would be well to institute new microscopical investigations, which shall either confirm the observations of Malpighi, Grew, Du Hamel, Mustel, and Hedwig, or prove that plants have a special organisation of their own which is more simple than that of animals, whether that organisation is supposed to originate in simple peculiar fibres and threads (Medicus) or with cellular and tubular tissue (tissu tubulaire of Mirbel). Attention should also be given to the following subordinate questions: 1. How many kinds of vessels may certainly be distinguished from the first period of their development? The existence of certain forms having been established; 2. Are the twisted fibres which are called spiral vessels (vasa spiralia) themselves hollow, and do they therefore form vessels, or do they serve by their convolutions for the formation of closed cavities, and how? 3. Do fluids as well as gases move in these cavities? 4. Do the scalariform ducts arise from adherence of the twisted threads (Sprengel), or do the threads owe their origin to the ducts (Mirbel)? Do alburnum and woody fibres originate in the scalariform ducts, or in true vessels, or in tubular tissue?'

We see in this case as in many similar ones, that the subject was proposed by persons who understood little of it, and who were unable to judge of what had been written about it; how else could they have placed the opinions of a Mustel and a Medicus side by side with those of Malpighi and Grew? Had Bernhardi or Mirbel set the question, it would certainly have been better conceived. It was in keeping that the three essays sent in, all inferior to Bernhardi's work already mentioned, though they contradicted one another on the most important points, were nevertheless all accepted; not less so that Treviranus' essay obtained only the second place, though it was decidedly better than the other two, and very much better than Rudolphi's. The best result of the whole affair was that it stirred up the phytotomists of the day, and led Mirbel to submit the three prize treatises to a searching criticism, especially that of Treviranus, which Mirbel with professional acumen recognised as the best. Link's essay appeared in 1807 under the title 'Grundlehren der Anatomie und Physiologie der Pflanzen,' that of Rudolphi as 'Anatomie der Pflanzen,' also in 1807, each forming a handsome octavo volume. The work of Treviranus had already appeared in 1806 with the title, 'Vom inwendigen Bau der Gewächse.'

If we compare the works of Link and Rudolphi[3], which both received a prize, and which had all the appearance of text-books of general vegetable phytotomy and physiology, we miss in both any clear exposition of the conceptions connected with the words used, and the train of thought therefore is constantly obscure and vacillating. Yet it is easy to see that they are opposed to one another in all essential points, Link[4] generally hitting on the correct, or at least the correcter view. For instance, Rudolphi denies altogether the vegetable nature of Fungi and Lichens, because he finds no resemblance between their hyphae and vegetable cell-tissue, and he supposes them to arise by spontaneous generation; even of the Confervae he says that the microscope has shown him nothing that agrees with the structure of plants, evidently a sign of bad observation or of incapacity to understand what he saw. Link on the other hand regards all Thallophytes as plants, and sees that the filaments of Lichens and Fungi consist of cells, and that cells occur at least in many Algae. Rudolphi praises in the same breath the views of Wolff and Sprengel on cell-tissue, although they are directly opposed to one another, and although he adopts Sprengel's strange theory of cell-formation without alteration. Link on the contrary declares against Sprengel's theory, and on good grounds, and shows that the vesicles which Sprengel took for young cells are starch-grains; at the same time he makes new cells be formed between the old ones. Rudolphi is of opinion that cells open into one another, as is plainly shown by the passage of coloured fluids. Link maintains that cells are closed bodies, and proves it well by the occurrence of cells with coloured juice in the middle of colourless tissue. Rudolphi represents the orifices of the stomata as encircled by a roundish rim, which he takes without hesitation for a closing muscle because the apertures enlarge and diminish. Link is more happy in taking the part that surrounds the aperture for a cell, or a group of cells. Rudolphi considers the great cavities in hollow stems and in the tissue of water-plants as the only air-passages in plants; Link explains these cavities as gaps caused by the irregular growth of cellular tissue. With Rudolphi the word vessel means not only vascular forms in wood, but milk-vessels and resin-ducts also, and to the former he even transfers Malpighi's view of the structure of spiral vessels. Link designates the tubes of the wood only as vessels, combining the most various forms of them under the term spiral vessels; he excludes milk-vessels, resin-ducts, and the like from the conception of a vessel, and in this he is somewhat inconsistent, since he assumes with Rudolphi that a vessel, in plants as in animals, is a canal for the conveyance of nutrient sap.

With all these contradictions, the two essays agree in adopting the old Malpighian view of the growth in thickness of stems, according to which the new layers of wood are formed from the inner layers of bast, while between the bast-cells, which are here taken to be identical with woody fibre, new spiral vessels arise contemporaneously, and, as Link expressly says, from juices which pour out between the bast-cells.

It is hard to understand how two treatises, so contradictory as they have been shown to be, could have both received a prize at the same time, or how the great difference could have been overlooked between Link's sensible and well-arranged account of his subject, and Rudolphi's uncritical statements, which everywhere rely more on old authority than on his own observation. It is however certain that Link's better production is inferior to Bernhardi's treatise, unless we choose to consider the greater copiousness of detail in Link, the number of his observations, and his aquaintance with the literature of the subject, as giving him the advantage. His figures, as well as Rudolphi's, are not so good as those of Bernhardi.

The work of Treviranus[5] to which the judges at Göttingen awarded the second place, is much less comprehensive than those of his competitors; the style is inferior to Link's, and may even be called clumsy. But the much better figures show at once that Treviranus was the more accurate observer, and his work, in spite of the inferiority of its style, is of far higher value on account of the attention paid in it to the history of development; Treviranus laid greater stress on this method than either Link or Rudolphi, and it led him to form views on some of the fundamental questions of phytotomy, in which we see the germs of theories afterwards perfected by von Mohl. His account of the formation of cell-tissue is mainly that of Sprengel, and therefore an unfortunate one; but nevertheless his observations on the composition of wood and the nature of vessels were as good and correct as could be expected from the condition of the microscope at the time. He made one discovery of considerable value, that of the intercellular spaces in parenchyma, but he lessened its merit by filling these passages with sap, and even describing its movement. Woody fibres are due, he thinks, to strong

  1. See P. Harting, 'Das Mikroskop,' § 433 and 434.
  2. Johann Jakob Bernhardi, born in 1774, was Professor of Botany in Erfurt, and died there in 1850.
  3. Karl Asmus Rudolphi, born at Stockholm in 1771, was Professor of Anatomy and Physiology in Berlin, and died there in 1832.
  4. Heinrich Friedrich Link was born at Hildesheim in 1767, and became Doctor of Medicine of Göttingen in 1788. In 1792 he became Professor of Zoology, Botany, and Chemistry in Rostock, Professor of Botany in 1811 in Breslau, and in 1815 in Berlin, where he died in 1851. He was a clever man of very varied accomplishment, but not a very accurate observer of details, and was held in estimation by many chiefly as a good teacher and philosophic author of popular works on natural science. He was one of the few German botanists in the early part of the present century who aimed at a general knowledge of plants, and combined anatomical and physiological enquiries with solid researches in systematic botany. Of his many treatises on all branches of botanical science, zoology, physics, chemistry, and other subjects, his Göttingen prize essay must be considered to have contributed most to the advancement of science. Von Martius somewhat overrates his scientific importance in his 'Denkrede auf H. F. Link' in the 'Gelehrte Anzeigen,' Munchen (1851), 58-69.
  5. Ludolf Christian Treviranus, born at Bremen in 1779, became Doctor of Medicine of Jena in 1801, and practised at first in his native town, where he became a teacher at the Lyceum in 1807. In 1812 he accepted the professorship in Rostock vacated by Link, and was again his successor in Breslau. In 1830 he exchanged posts with C. G. Nees von Esenbeck, who was a professor in Bonn; he died in that town in 1864. In the first part of his life he occupied himself chiefly with vegetable anatomy and physiology, afterwards with the determination and correction of species. His first works, which are noticed in the text, and the treatises on sexuality and the embryology of the Phanerogams, published between 1815 and 1828, are the most important in a historical point of view. His 'Physiologie der Gewächse' in two volumes (1835-1838) is still of value for its accurate information on the literature of the subject; but it can scarcely be said to have contributed to the advance of physiology, for its author adhered in it to the old views, and especially to the notion of the vital force, at a time when new ideas were already asserting themselves. The 'Botanische Zeitung' for 1864, p. 176, contains a notice of his life.