Though Treviranus’ account of vegetable tissues was on the whole weak as far as concerns the history of development, yet Mirbel[77] recognised in him the most dangerous opponent of his own theory, and addressed a public letter to him and not to his other German antagonists, Sprengel, Link and Rudolphi, in defence of the views he had formerly expressed. This letter is the first part of a larger work which appeared in 1808, ‘Exposition et défense de ma théorie de l’organisation végétale,’ in which Mirbel endeavours to meet the objections of his opponents with great adroitness of style and with the results of varied rather than profound observation, and to find new arguments for his theory of vegetable tissue; he admits that his former treatises were in many respects faulty, but demands that his critics should discuss his system as a whole and not take offence at single expressions. Mirbel’s idea of the inner structure of plants is essentially the same as that broached by Caspar Friedrich Wolff. The first and fundamental idea is that all vegetable organisation is formed from one and the same tissue differently modified. The cell-cavities are only hollow spaces of varying form and extension in homogeneous original matter, and have no need therefore of a system of filaments, as Grew supposed, to hold them together. The tracheae only are an exception, which Mirbel, in striking opposition to the much more correct view of Treviranus, considers to be narrow spirally wound laminae, inserted into the tissue and connected with it only at the two ends. If it is asked how interchange of sap is effected in such a cellular tissue as this, it becomes necessary to assume that the membranous substance of plants is pierced by countless invisible pores, through which fluids find their way. But nature has a speedier and more powerful instrument in the larger pores, which the microscope reveals. Mirbel does not discuss the question how the fluids are set in motion, easily disregarding such mechanical difficulties, as was usual in those days, when vital power was always in reserve to be the moving agent. He warmly repels the imputation, which Sprengel had made against him, of having confounded pores and granules, by appealing to his figures; he says that he has depicted prominences on the outside of the walls of the dotted vessels, and an orifice in each of them, which his opponents simply never saw. The question whether these prominences lie on the inside or the outside of the walls of the vessel has no meaning, if we suppose with Mirbel that the dividing wall is single; he is only concerned to enquire whether the perforated projections lie on the one or on the other side of the wall. He refers Treviranus, who had denied the presence of the pores, to his description of scalariform vessels, in which he had himself seen the slits which correspond to the pores.
In comparison with these fundamental questions Mirbel’s further account of matters of detail does not concern us here. He gave a connected view of the whole of his doctrine of tissues in the form of aphorisms, which occupy the second part of his book. Of all that he says on the five kinds into which he distinguishes vessels the most interesting is the statement, that diaphragms pierced like a sieve separate the different members of his ‘beaded’ vessels. We find that the weakest part of Mirbel’s phytotomy, as of that of his opponents, is his description of the true vessels (vasa propria), with which he classes the milk-cells of the Euphorbiae and the resin-ducts of Coniferae, but he saw clearly enough that the latter were canals inclosed in a layer of tissue of their own. The third part of the book is devoted to these forms of tissue, and we learn that he classes not only many kinds of sieve-cell-bundles, but also true bast-fibres, as those of nettle and hemp, with his bundles of true vessels. Like his opponents he makes the growth in thickness in woody stems to be due to transformation of the inner layer of bast; but he gives a new turn to this view, which brings it nearer to the modern theory; during the period of vegetation a delicate tissue with large vessels is developed in Dicotyledons on the confines of the wood and the bark, and these augment the mass of the woody body, while a loose cellular tissue is formed on the other side, destined to replace the constant losses of the outer rind. To later phytotomists, who understood by the word cambium a thin layer of tissue constantly engaged in producing wood and rind, Mirbel’s otherwise indistinct conception of growth in thickness must have become more indistinct from his using the word cambium not for the layer of tissue afterwards so called, but for a highly ‘elaborated and purified sap’ which is intended for the food of the plant and makes its way through all membranes; we see this cambium-sap appear at the spots where it produces new tubes and cells after the manner of the Wolffian theory. The cells appear at first as minute spheres, the tubes are very fine lines; both enlarge and gradually show pores, clefts, etc. This is essentially Wolff’s doctrine, which Mirbel afterwards endeavoured to confirm against his German opponents from the germination of the date-palm with the help of a more powerful microscope.
Mirbel insisted more than the German phytotomists of his day on the idea, that all forms of vegetable tissue are developed originally from young cell-tissue, an idea suggested by Sprengel and following naturally with Mirbel from Wolff’s theory. Both Mirbel and Wolff were hasty in observation and too much under the influence of theory in giving reasons for what they observed, and therefore too ready with far-reaching explanations of phenomena which only long-continued observation could decide.
Treviranus replied, though after some delay, to Mirbel’s polemics by incorporating into his ‘Beiträge zur Pflanzenphysiologie,’ Göttingen (1811), an essay entitled ‘Beobachtungen im Betreff einiger streitigen Puncte der Pflanzenphysiologie,’ in which he again took up the questions in dispute between himself, Mirbel, Link and others, and supported his own views by fresh investigations. It cannot be denied that in this short treatise Treviranus brought some important questions nearer to a decision; he added materially to the knowledge of bordered pits, on which subject his views now approximated more nearly to those of Mirbel; he drew attention to the vesicular nature of vegetable cells, which are often separable from one another, and to the occurrence of true spiral vessels in the neighbourhood of the pith in Conifers also, and among other things discovered the stomata on the capsule of Mosses. On the subject of the theory of cell-formation which he had borrowed from Sprengel, he endeavoured to extricate himself from his difficulty by ingeniously pointing out that though the starch-grains in the seed-leaves of the bean disappear without producing new cells in them, they are dissolved and then serve as fluid material for new cell-formation in other parts of the germinating plant, which however was giving up Sprengel’s theory; yet he cited as a direct proof of that theory the origination of gonidia in the cells of Hydrodictyon, and their development into new nets.
Mirbel and his German opponents moved for the most part in a circle of ideas which had been formed by the speculations of Malpighi, Grew, Hedwig and Wolff, though it must be allowed that the observations of Treviranus did open new points of view. But Johann Jakob Paul Moldenhawer[78] travelled far beyond these older views as early as 1812 in his important work, ‘Beiträge zur Anatomie der Pflanzen.’ He took up from the first a more independent position as regards former opinions than either of the writers hitherto considered. He relied on very detailed, varied, and systematic observations evidently made with a better instrument, abided by what he himself saw, and chose his point of view in accordance with it, while he criticised the views of his predecessors in detail with an unmistakable superiority, and in so doing displayed minute acquaintance with the literature of the subject and varied phytotomical experience. He fixed his eye firmly on the points in question, and made each one the subject of earnest investigation and copious and perspicuous discussion. His figures prove the carefulness of his examination and the greater excellence of his instrument; they are undoubtedly the best that were produced up to 1812. His mode of dealing with his subject and his figures, though they were not executed by himself, remind us in many respects of von Mohl, though it would be more correct to say that von Mohl’s manner reminds us of Moldenhawer, for from the great respect which von Mohl displays for him, especially in his earlier writings, it can scarcely be doubted that he formed himself on Moldenhawer’s ‘Beiträge,’ and first learnt from them the earnestness and carefulness demanded by phytotomic work.
It has been already mentioned that the study of vegetable physiology is indebted to Moldenhawer for one important practical improvement. He was the first who isolated cells and vessels by allowing parts of plants to decay in water and afterwards crushing and dissecting them, a process not much used in modern times, though it may still be applied with advantage in conjunction with what is known as Schulze’s solution, especially if it is carried out with Moldenhawer’s carefulness and circumspection. The isolation of the elementary organs of plants by maceration in water necessarily brought Moldenhawer into direct antagonism with Mirbel, who with Wolff assumed that the partition between any two cells was a single wall; whereas Moldenhawer found that the cells and vessels were closed tubes and sacs after isolation, and must necessarily, as it would seem, so lie one against another in the living plant, that the wall between every two cell-spaces is formed of two membranous laminae, and he expressly says that this is the case even in very thin-walled parenchyma. This result remained unassailable, so long as no one was in a position to conclude from the history of the development of cell-tissue that the partitions are originally single, or by aid of strong magnifying power to prove the true structure of the walls and their later separation, and the differentiation of the once single wall into two separable laminae. If the view based on the results of maceration was still not the true view, yet it was nearer the truth as regards the matured state of the cell-wall than that of Wolff and Mirbel, and the important advantage was gained of being able to study the form of elementary organs and the sculpture on their walls more accurately than before. It is true that Link had occasionally isolated cells by boiling in 1809, and Treviranus had drawn attention in 1811 to the fact that it was possible to isolate parenchyma-cells in their natural condition; but neither of them made systematic use of these observations, and to Moldenhawer belongs the exclusive merit of having first isolated vessels and woody cells; but as usually happens, he did not himself obtain all the possible results from his method of preparation. In his work which indeed embraces the whole of phytotomy, he is continually recurring to one species, maize. This supplies the starting-point in every question to be discussed. The results obtained there are the firm supports on which he leans in the examination of a great variety of plants, which he then compares together at length. This mode of treatment was well chosen both for investigation and instruction in the existing state of the science; it was a particularly happy idea that of choosing the maize-plant for his purpose; former phytotomists had generally had recourse to dicotyledonous stems, and preferred those that had compact wood and complex rind, but the examination of these plants presents difficulties at the present day to a practised observer with a good microscope. Occasionally observers had taken the stem of the gourd, where the large cells and vessels suited small magnifying power, but where many abnormal conditions occurred to interfere with their conclusions. The Monocotyledons, like the Vascular Cryptogams, had hitherto been comparatively neglected. When then Moldenhawer made a monocotyledonous and rapidly growing plant, with very large-celled tissue and comparatively very simple structure, the chief subject of his investigations, he was sure to succeed in making out many things more clearly than his predecessors. It was an important point that he found the fibrous elementary organs in this plant united with the vessels into bundles, which are separated by a strict line of demarcation from the large-celled parenchyma that surrounds them. Thus the peculiar character, the idea, of the vascular bundle was brought prominently into contrast with that of other forms of tissue. This took the place of the distinction between rind, wood, and pith, which had served former phytotomists as the basis of their histological survey, but which is in itself only a secondary result of the later elaboration of certain parts of the plant. Moldenhawer, in laying the chief stress from the first on the contrast between vascular bundles and parenchyma, hit upon a histological fact of more fundamental importance, the right appreciation of which has since enabled the phytotomist to find his way through the histology of the higher plants. For while the construction of Monocotyledons and Ferns must seem abnormal and quite peculiar to any one who starts with examining the rind, wood, and pith of old dicotyledonous stems, those on the contrary who, with Moldenhawer, have recognised a special histological system in the vascular bundles of Monocotyledons, have the way opened to them to seek for a similar one in the Dicotyledons, and to refer the secondary phenomenon of wood and rind to the primary existence of vascular bundles. Moldenhawer did in fact open this way, when he showed how the growth of a dicotyledonous stem may be understood from the structure and position of the originally isolated vascular bundles (Beiträge, p. 49, etc.). But he was thus of necessity led to the rejection of Malpighi’s theory of the growth in thickness of woody stems, which all vegetable anatomists from Grew to Mirbel had adopted. Though Bernhardi and Treviranus made weak attempts to discredit it, Moldenhawer was the first who distinctly rejected the origin of the external layers of wood from the inner bast, and proposed the first really practical basis for the later and correct theory of secondary growth in thickness (p. 35). The removal of this ancient error is in itself a very important result, and one which, apart from all other services, must secure him an honourable place in the history of botany.
But the light must have its attendant shadow, and all his carefulness in observation and cautiousness in judgment did not protect him from one prejudice and its evil consequences. After Moldenhawer had isolated the elementary organs by maceration, he had to answer the question how we are to conceive of their firm coherence in the living plant. He came to the conclusion, as did von Mohl, Schacht, and others after him, that there must be some special connecting medium; but he did not hit upon their idea of a matrix, in which the cells are imbedded, or of a cement which holds them together, but on a much stranger theory, which reminds us at once of Grew’s thread-tissue, and like that rests partly on incorrect observations. These were too hastily accepted as the basis of a theory which in its turn interfered with after observations. He thought that the cells and vessels were surrounded and held together by an extremely delicate network of fine fibres; in some cases he really believed that he saw these fibres, and interpreted in this way the thickened bands in the well-known cells of Sphagnum, and still more strangely he appears to have taken the thickened longitudinal and transverse edges of cells and vessels for such threads. The unfavourable impression produced by this theory is necessarily heightened by the fact that he gave the name of cell-tissue, a term long used in a different sense, to his fancy-structure of reticulated threads which were to hold the cells and vessels together, while he called the parenchyma itself cellular substance, an expression which fortunately no one copied, and which certainly contributed at a later time to discredit the great services which Moldenhawer rendered to phytotomy.
His ‘Beiträge zur Anatomie der Pflanzen’ are divided into two portions; the first treats of the parts surrounding the spiral vessels; the second of the spiral vessels themselves.
The position and collective form of the component parts of the vascular bundle in the stem of the maize-plant are well described in the first section of the work. It is correctly stated that there is a sheath to the whole bundle composed of strongly thickened fibrous cells, that each of these cells has its own membrane and is entirely closed, and that they resemble the bast and the fibrous elements of the wood of Dicotyledons. The segmented wood-cells and the parenchyma-cells of the wood arranged in rows are incidentally noticed. Under the name of fibrous tubes he included the cells of the sclerenchyma-sheath of many vascular bundles and the true bast and wood-fibres, which latter he says are wanting in the Coniferae. He explained the secondary growth in thickness of rind and bast by the example of the shoot of the vine, in which he correctly distinguished the medullary sheath and the spiral vessels. In herbaceous Dicotyledons he found the bundles of vessels to consist of a bast portion and a woody portion, and he attributed the formation of the compact wood of true woody plants to the blending together of the woody portions of these separate bundles.
In discussing the parenchymatous cell-tissue he rejects emphatically and on good grounds the origin of new cells from the granular contents of older ones, which had been the view of Sprengel and Treviranus, as also the theory of Wolff and Mirbel, while he maintains against Mirbel especially, that the separation of fibrous tubes is possible even where no dividing line can be seen between them in the cross section. He considers that both in thin-walled and thick-walled parenchyma the dividing wall is double and the cell-membrane entirely closed. ‘It appears,’ he continues on p. 86, ‘from these observations that cellular substance consists of separate closed tubes, which may be round or oval, or more or less elongated, or almost cylindrical in shape, and these by mutual pressure assume an angular and flattened form, which is either regular like the cells of the comb of bees or more or less irregular. Such an aggregate of separate cells (and here he is certainly quite right) has nothing in common with a tissue, and the word cell-tissue seems therefore less suitable than the term cellular substance, composed of cell-like tubes.’ Further on he rejects Mirbel’s idea of the existence of visible holes in the walls of cells, and points out that they are not necessary for the movement of sap. The dispute between Mirbel and his opponents respecting the porousness of cell-walls was extended at the same time to the stomata of the epidermis[79], the slits in them being supposed to be apertures in the epidermis regarded as a simple membrane. Moldenhawer took occasion to examine the anatomy of stomata more closely, and produced the first accurate descriptions and figures of these organs, showing especially that the apertures are not surrounded by a simple border, as most previous observers believed, but lie between two cells, and that therefore they are not examples of the existence of pores in cell-walls, as Mirbel imagined. It may be observed here by the way, that Mirbel afterwards considered stomata to be short broad hairs; Amici in 1824, Treviranus in 1821, demonstrated their true structure by cross sections, and von Mohl at a later period investigated it thoroughly. Moldenhawer on the present occasion also enquired into the faculty attributed to stomata of opening and closing alternately, which, first observed by Comparetti, was then much discussed by the German phytotomists, and has been made the subject of repeated investigation in modern times. The whole of this discussion was in connection with the question of the pitting of cell-walls, the true nature of which Moldenhawer however never clearly understood.