History of botany (1530–1860)/Book 1/Chapter 5
Morphology and Systematic Botany under the Influence of the History of Development and the Knowledge of the Cryptogams.
In the years immediately before and after 1840 a new life began to stir in all parts of botanical research, in anatomy, physiology, and morphology. Morphology was now specially connected with renewed investigations into the sexuality of plants and into embryology, and attention was no longer confined to the Phanerogams but was extended to the higher and later on to the lower Cryptogams. These researches into the history of development first became possible, when von Mohl had restored the study of anatomy, and Nägeli had founded and elaborated the theory of cell-formation about the year 1845. The success of both these enquirers was due to the previous development of the art of microscopy; it was the microscope which revealed the facts on which the foundations of the new research were laid, while its promoters at the same time started from other philosophical principles than those which had hitherto prevailed among botanists. Investigation by means of the microscope enforces on the observer the very highest strain of attention and its concentration on a definite object, while at the same time a definite question to be decided by the observation has always to be kept before the mind; there are sources of error on all sides to be avoided, and possible deceptions to be taken into consideration; the securing of the facts demands all the powers which specially display the individual character of the observer. Thus serious attention to microscopy was one of the causes which introduced the best observers to the practice of inductive enquiry, and gave them an insight into its nature; and in a few years' time when the actual results of these investigations began to appear, and when a wholly new world disclosed itself to botanists, especially in the Cryptogams, then questions arose on which the dogmatic philosophy had not essayed its ancient strength; the facts and the questions were new and untouched, and presented themselves to unprejudiced observation in a purer form than those, which during the first three centuries had been so mixed up with the old philosophy and with the principles of scholasticism. Von Mohl, who only occasionally occupied himself with morphological subjects, was a firm adherent of the inductive method, and was bent on the establishment of individual facts rather than of general principles; but the founders also of the new morphology, Schleiden and Nägeli, started from philosophical points of view, which, different as they were in the two men, had yet two things in common, a demand for severely inductive investigation as the foundation of all science, and the rejection of all teleological modes of explaining phenomena, in which latter point their opposition to the idealistic nature-philosophy school was most distinctly manifested. They had indeed one very important point of contact with this school, the belief in the constancy of organic forms; but this belief, not being connected with the Platonic doctrine of ideas, was with them only a recognition of evcry-day observations, and was therefore of less fundamental importance, being felt merely as an inconvenient element in the science. Treating the question in this way, and influenced by the results of the new researches, they either inclined to entertain the idea of descent before the appearance of Darwin's great work, or gave a ready assent to the principle of the new doctrine, though they expressed some doubts respecting matters of detail. Hofmeister's researches in morphology and embryology ('Vergleichende Untersuchungen,' 1851) threw an entirely new light on the relations of affinity between the great groups in the vegetable kingdom, and were leading more and more to the view, that there must be some special peculiarity in the question of the constancy of organic forms. But the idea of evolution in the vegetable kingdom was brought more distinctly home to men's minds by palaeontological researches; Sternberg (1820-1838), Brongniart (1828-1837), Goeppert (1837-1845), and Corda (1845) made the flora of former ages the subject of careful study, and compared fossil plants with living allied forms. Unger especially, while advancing the knowledge of the structure of cells and of vegetable anatomy and physiology, and generally taking a prominent part in the development of the new botany, applied the results of its investigations to the examination of primeval vegetation, and showed the morphological and systematic relations between past and existing floras. After twenty years of preliminary study he declared distinctly in 1852, that the immutability of species is an illusion, that the new species which have made their appearance in geological periods are organically connected, the younger having arisen from the elder. It was shown in the former chapter, how about the same time the leading representative of idealistic views, Alexander Braun, was driven to the hypothesis, though in a more indefinite form, of an evolution of the vegetable kingdom: and in the year that Darwin's book on the origin of species appeared, Nägeli ('Beitrage,' ii. p. 34) wrote:—'External reasons, supplied by the comparison of the floras of successive geological periods, and internal reasons given in physiological and morphological laws of development and in the variability of the species, leave scarcely a doubt that species have proceeded one from another.'
Though these words might not contain a theory of descent capable at once of scientific application, yet they show that the latest researches and candid appreciation of facts were compelling the most eminent representatives of the botany of the day to give up the constancy of forms. At the same time in the genetic morphology which had developed itself mainly under Nägeli's guidance since 1844, and still more in embryology, which in Hofmeister's hands was leading to results of the greatest systematic importance, there lay a fruitful element destined to correct and enrich Darwin's doctrine of descent in one essential point. That doctrine in its original form sought to show that selection, the result of the struggle for existence, combined with perpetual variation was the sole cause of progressive improvement in organic forms; but Nägeli, relying on the results of German morphology, was able as early as 1865 to point out that this explanation was not satisfactory, because it leaves unnoticed certain morphological relations, especially between the large divisions of the vegetable kingdom, which scarcely seem explainable by mere selection in breeding. While Nägeli allowed that Darwin's principle of selection was well adapted to explain fully the adaptation of organisms to their environment and the suitableness and physiological peculiarities of their structure, he pointed out that in the nature of plants themselves there are intimations of laws of variation, which lead to a perfecting of organic forms and to their progressive differentiation, independently of the struggle for existence and of natural selection; the importance of this result of morphological research has since been recognised by Darwin. Thus Nägeli supplied what was wanting in the theory of descent and gave it the form, in which it is adequate to explain the problem already recognised by the systematists of the old persuasion, namely, how it is possible for the morphological affinity of species in the system to be in so high a degree independent of their physiological adaptation to their environment.
The modern teaching on vegetable cells, modern anatomy, and morphology, and the improved form of the theory of selection are the product of inductive enquiry since 1840, a product, the full importance of which will be described in the following portions of our history. At present we have to deal only with morphological and systematic results, and therefore with a part only of the abundant labours of the botanists who will be noticed in this chapter; the remainder will be reserved for succeeding books, which contain the history of the anatomy and physiology of plants.
It is one of the characteristic features of this period of botany, that morphology enters into the closest connection with the doctrine of the cell, with anatomy and embryology, and that researches, especially into the process of fecundation and the formation of the embryo, form to some extent the central point of morphological and systematic investigations. A strict separation of these various enquiries, which are all ultimately applicable to the purposes of systematic botany, can therefore scarcely be maintained, and least of all in dealing with the lower Cryptogams.
The condition of botanical literature about the year 1840 was highly unsatisfactory; it is true that eminent service was rendered in the several domains of systematic botany, morphology, anatomy, and physiology, and a number of von Mohl's best works were produced in this period; Meyen also, Dutrochet, Ludolph Treviranus and others were cultivating vegetable anatomy and physiology, and it has been already stated that good and noticeable work was done in the previous years in morphology and systematic botany. But there was no one to put together, to criticise and apply the knowledge which had been accumulated in all parts of the science; no one really knew what a wealth there was at that time of important facts; least of all was it possible to form a judgment on the matter from the text-books of the period, which were deficient in ideas and facts, and crammed with a superfluous terminology; their mode of treating their subject was trivial and tasteless, and whatever was specially worth knowing and important to the student they did not contain. Those who undertook really scientific enquiries separated themselves from those who dealt with botany after the old schematism of the Linnaean school; but botanical instruction, the propagation of knowledge, was almost everywhere in the hands of this school, though it was the one least fitted for the task; and thus a mass of lifeless phrases was the instruction offered to the majority of students under the name of botany, with the inevitable effect of repelling the more gifted natures from the study. This was the evil result of the old and foolish notion, that the sole or chief business of every botanist is to trifle away time in plant-collecting in wood and meadow and in rummaging in herbaria, proceedings which could do no good to systematic botany even as understood by the Linnaean school. Even the better sort lost the sense for higher knowledge while occupying themselves in this way with the vegetable world; the powers of the mind could not fail after a time to deteriorate, and every text-book of the period on every page supplies proof of this deterioration.
But such a condition of things is dangerous for every science; of what profit is it, that single men of superior merit advance this or that part of the science when a connected view of the whole is wanting, and the beginner has no opportunity of studying the best things in their mutual relations. However, the right man was found at the right moment to rouse easy indolence from its torpor, and to show his contemporaries, not in Germany only but in all countries where botany was studied, that no progress was possible in this way. This man was Matthias Jacob Schleiden, born at Hamburg in 1804, and for many years Professor in Jena. Endowed with somewhat too great love of combat, and armed with a pen regardless of the wounds it inflicted, ready to strike at any moment, and very prone to exaggeration, Schleiden was just the man needed in the state in which botany then was. His first appearance on the scene was greeted with joy by the most eminent among those who afterwards contributed to the real advance of the science, though their paths it is true diverged considerably at a later period, when the time of reconstruction was come. If we were to estimate Schleiden's merit only by the facts which he discovered, we should scarcely place him above the level of ordinarily good botanists; we should have to reckon up a list of good monographs, numerous refutations of ancient errors and the like; the most important of the theories which he proposed, and over which vigorous war was waged among botanists during many years, have long since been set aside. His true historical importance has been already intimated; his great merit as a botanist is due not to what he did as an original investigator, but to the impulse he gave to investigation, to the aim and object which he set up for himself and others, and opposed in its greatness to the petty character of the text-books. He smoothed the way for those who could and would do really great service; he created, so to speak, for the first time an audience for scientific botany capable of distinguishing scientific work from frivolous dilettanteism. Whoever wished from this time forward to take part in the discussion of botanical subjects must address all his powers to the task, for he would be judged by another standard than had hitherto prevailed.
Schleiden, who had commenced his botanical labours with some important researches in anatomy and the history of development, the most valuable of which in matter and form was an enquiry into the development of the ovule before fertilisation (1837), composed also a comprehensive text-book of general botany, which appeared first in 1842–3, and in much improved editions in 1845 and 1846, and in two subsequent years. The difference between this and all previous text-books is the difference between day and night; in the one, an indolent carelessness and an absence of ideas; in the other, a fulness of life and thought, calculated to influence young minds all the more, because it was in many respects incomplete and still in a state of fermentation. On every page of this remarkable work, by the side of facts really worth knowing, the student found interesting reflections, a lively and generally coarse polemic, and praise and blame of others. It was not a book to be studied quietly and comfortably, but one that excited the reader everywhere to take a side for or against, and to seek for further instruction.
The work is generally quoted as 'Grundzüge der wissenschaftlichen Botanik,' but its chief title is 'Die Botanik als inductive Wissenschaft,' which indicates the point on which Schleiden laid most stress. His great object was to place the study, which had been so disfigured in the text-books as scarcely to wear the semblance of a natural science, on the same footing with physics and chemistry, in which the spirit of genuine inductive enquiry into nature had already asserted itself in opposition to the nature-philosophy of the immediately preceding years. It may seem strange to us now to see a text-book of botany introduced by a formal essay, 131 pages long, on the inductive method of investigation as opposed to dogmatic philosophy, and to find the principles of induction set forth again and again in connection with a great variety of subjects in the book itself. Many objections may be raised to the contents of this introduction; it may be said that many philosophical dicta are misunderstood in it; that Schleiden himself has frequently offended against the rules there laid down, for instance, when he substitutes a formative impulse (nisus formativus) for the vital force which he rejects, which is only introducing vital force again under another name; that it is superfluous to present the history of development as a 'maxim' in Kant's use of the word, instead of showing that the history of development enters naturally and of itself into inductive investigation, and so on. All this will not lessen the historical importance of this philosophic introduction; the traditional way in which descriptive botany was at that time presented to the student was so thoroughly dogmatic and scholastic, trivial and uncritical, that it was necessary to impress upon him in many words, that this is not the method of true investigation of nature.
Passing on to the more special problems of botanical enquiry, Schleiden next dwells on the history of development as the foundation of all insight into morphology, though he overshot the mark when he rejected as unfruitful the simple comparative method, which had produced considerable results in the hands of De Candolle, and was virtually the fruitful element in the doctrine of phyllotaxis of Schimper and Braun. Still he took an active part himself in the study of development in plants, and gave special prominence to embryology; he also discussed the doctrine of metamorphosis from the point of view of the history of development, and pointed to Caspar Friedrich Wolff's treatment of that subject as much clearer than that which had been introduced by Goethe. Finally, Schleiden's mode of dealing with the natural system must be reckoned among the good services which he rendered to method; not because his classification of the vegetable kingdom presents any specially interesting features or brought to light any new affinities, but because we see an attempt made for the first time to give detailed characters drawn from morphology and the history of development to the primary divisions, and because by this means the positive and distinct nature of the Cryptogams was from the first clearly brought out. The old way of treating morphology, as though there were only Phanerogams in the world, and then having recourse to unmeaning negatives in dealing with the Cryptogams, was thus set aside, much to the profit of the immediate future, which directed its attention specially to the Cryptogams.
Schleiden however did not succeed in securing firm ground for the morphology of the Cryptogams as founded on the history of their development. His investigations into the morphology of the Phanerogams were more successful. His theory of the flower and fruit is an admirable performance for the time, even though we abandon his view of the stalk nature of placentas and some other notions, as we obviously must. As Robert Brown founded the history of the development of the ovule, so Schleiden founded that of the flower, and his example influenced other botanists. Soon investigations into the genesis of the flower was one of the chief occupations of morphologists, and the results of enquiry into development proved to be of great value for the systematic arrangement of the Phanerogams, especially when more exact attention was paid to the sequence of development in the organs of an inflorescence, to abortion, doubling and branching of the stamens, and to the like matters. Duchartre, Wigand, Gelesnoff and many others, were soon working in the same direction with great success. Payer deserves special mention for his enormous perseverance in examining the development of the flower in all the more important families in his 'Organogenic de la fleur,' 1857, and thus producing a standard work, distinguished alike for the certainty of the observations, the simple unbiassed interpretation of the things observed, and the beauty and abundance of the figures—a work which became more important every year for the morphology of the flower.
Schleiden's text-book was the first of its kind that supplied the student with really good figures based on careful observations. With all its many and obvious defects it had one merit which cannot be rated too highly; its appearance at once put botany on the footing of a natural science in the modern sense of the word, and placed it upon a higher platform, extending its horizon by raising its point of view. Botany appeared all at once as a science rich in subject-matter; Schleiden had not only himself made many investigations and broached new theories, but he everywhere drew attention to what was already before the world and was important; for it is not sufficient as regards the literature of a science that there should be good investigators; it is as necessary that the scientific public, and especially the rising generation of professed students, should be well and sufficiently instructed in the art of distinguishing important from unimportant contributions. It must be distinctly affirmed in this place, that if Schleiden's theory of cell-formation, his strange notion about the embryology of Phanerogams and the like were very quickly shown to be untenable, this does not in the least affect the great historical importance which his writings possess in the sense here indicated.
That others besides Schleiden in the period following 1840 felt strongly, that botany must thenceforward give up its complacent resting in the old ideas, was shown among other things by the addition at this time of new periodicals to the old journal 'Flora.' The 'Botanische Zeitung' was founded by von Mohl and Schlechtendal in 1843, and the 'Zeitschrift für wissenschaftliche Botanik' by Schleiden and Nägeli. The latter, however, only lived three years, from 1844 to 1846, and was filled almost entirely with Nägeli's contributions. Both publications expressly set themselves the task of representing the new aims in the science. The immediate consequence was that 'Flora' braced up its energies, and endeavoured to do more justice to the modern spirit; excellent notices of botanical works now appeared in it under the exclusive management of Fürnrohr.
Schleiden's productivity in the higher sense of the word expended itself in his labours on the elements of scientific botany. His later somewhat discursive writings exerted no great influence on the further development of the science. The ideal which he had set up for scientific botany and had sketched in its larger outlines, could only be realised by the most persevering labour not of one man only, but of whole generations of observers and thinkers, nor did he apply himself with painful unremitting industry to the attainment of this exalted aim.
Soon after Schleiden's 'Grundzüge' first stirred the scientific world, a man of a very different character of mind began to address himself to the great task. This was Carl Nägeli, whose researches from this time onwards laid the foundations of knowledge in every department of botany. He showed what points were the most immediately attainable, and aided in perfecting the inductive method of enquiry and in advancing the study of the history of development. He did not make discoveries here and there by desultory efforts, but worked with earnest endurance at every question which he took up till he had arrived at a positive result; and this was almost always an enlargement of previous knowledge, and a new foundation on which others might build, and a copious literature be developed.
Nägeli like others felt the necessity of first determining his position with respect to the philosophical principles of the investigation of nature, but he did not proceed to give a general exposition of the inductive method as opposed to the dogmatism of the idealistic school. He went straight to the application of the laws of induction to the most general problems of organic nature, and specially of vegetation. It is easy to say that the task of natural science is simply to deduce conceptions and laws from the facts of experience by aid of exact observation. Many considerations present themselves as soon as the attempt is made to satisfy this demand; for it is not enough merely to accumulate individual facts, the point to which the inductive enquiry is to lead must be kept constantly and clearly before the mind. Nägeli insisted that it is only in this way that facts and observations have any scientific value; that the one important thing is to make every single conception obtained by induction find its place in the scheme of all the rest of our knowledge. With greater consistency of reasoning than Schleiden, and in entire accordance with the nominalist view of genuine investigation of nature in its sternest opposition to the idealistic school, Nägeli's first principle is not only to deduce conceptions from the observation of phenomena, to classify them and establish their subordination, but to treat these conceptions as mere subjective products of the understanding and employ them as instruments of thought and communication, and to be always ready to modify them as soon as inductive enquiry renders such modification necessary. Till this happens, the conception once laid down and connected with a word is to be strictly adhered to, and every arbitrary change or confusion with another conception is strictly forbidden. Since in nature everything is in movement, and every phenomenon is transitory, presenting itself to us in organic life especially as the history of development, all due regard must be paid to this condition of constant motility in forming scientific conceptions. The history of development is not merely to be treated generally as one of various means of investigation, but as identical with investigation into organic nature. These views are expressed in Nägeli's detailed observations on method in the first and second volume of the journal which he brought out in conjunction with Schleiden in 1844 and 1855, where the chief hindrance to his carrying them out fully and consistently is also to be found; for, like all his contemporaries, Nägeli believed at that time in the constancy of species, and consistently with this view he looked upon the natural system as a framework of conceptions, though these do not take the form of Platonic ideas with him as with the systematists of the idealistic school. It is equally consistent with his philosophical position, which refused to regard a change in our conceptions as a change in things themselves, that 'the idea of metamorphosis' in the sense of Goethe and Alexander Braun disappears in Nägeli from the field of scientific observation. It has been shown in the previous chapter that what Goethe called the normal or ascending metamorphosis has no scientific meaning unless species are supposed to be variable. It appeared moreover that if the Cryptogams are made the chief subjects of investigation, as Nägeli made them, the so-called metamorphosis of the leaves is a phenomenon of secondary importance, and only attains to its full importance in the Phanerogams. If Schleiden, illogically from his point of view, conceived of metamorphosis as the principle of development, Nägeli on the contrary scarcely employed the word. He regarded the history of development as the law of growth of the organs, and, in accordance with the theory of the constancy of species, the law of growth of every species and every organ was invariable in the same sense in which we apply the term to natural laws in physics and chemistry. In a word, Nägeli's considerations on the 'present task of natural history' in the work above cited, are not only logically and entirely consistent on the principles of the inductive method, but they are also consistent where others have been misled by the theory of the constancy of species into illogical conclusions.
Nägeli set himself in earnest to meet the demands of inductive enquiry, such as he had himself described them. It will be shown more in detail in the history of phytotomy, how he satisfied these demands in his refutation of Schleiden's doctrine of the cell, and in the establishment of his own, and at a later time in the framing of his theory of molecular structure and of the growth of organised bodies, and how he made these investigations true models of genuine inductive enquiry. Here we are concerned only with what he effected in this way for morphology and systematic botany. In this field of research he introduced two innovations of the profoundest importance, which affected both the aim and method of enquiry for some years. He connected his own morphological investigations, as far as possible, with the lower Cryptogams, extending them afterwards to the higher Cryptogams and to the Phanerogams; that is, he proceeded from simple and plain facts to the more difficult, thus not only introducing the Cryptogams into the field of systematic investigation, but making them its starting-point. In this way morphology not only secured a foundation in exact historical development, but it assumed a different aspect, inasmuch as the morphological ideas hitherto drawn from the Phanerogams were now examined by the light of the history of development in the Cryptogams. This was one innovation; the second, closely connected with it, was the way in which Nägeli made the new doctrine of the cell the starting-point of morphology. Both the first commencement of organs and their further growth were carried back to the formation of the separate cells; and the remarkable result was to show, that in the Cryptogams especially, whose growth is intimately connected with cell-division, precise conformity to law obtains in the succession and direction of the dividing walls, and that the origin and further growth of every organ is effected by cells of an absolutely fixed derivation. The most remarkable thing was, that every stem and branch, every leaf or other organ has a single cell at its apex, and that all succeeding cells are formed by division of this one cell according to fixed laws, so that the origin of all cell-tissue can be traced back to an apical cell; and as early as the years 1845 and 1846 Nägeli described in the 'Zeitschrift für wissenschaftliche Botanik' the three main forms, according to which the segmentation of an apical cell proceeds, namely, in one, two, and three rows (Delesseria, Echinomitrium, Phascum, Jungermannia, Moss-leaves). In this way the separate points in the history of growth in the Cryptogams were brought out with unusual clearness and decision; but on the other hand, Nägeli showed in 1844 in the case of a genus of Algae (Caulerpa) that the growth of a plant may show the usual morphological differentiation into axis, leaf, and root, when the propagative cell undergoes no cell-divisions in the process of development and further growth, and similar conditions were for the first time demonstrated in 1847 in Valonia, Udotea, and Acetabularia. Beside other results it was established by these facts, that morphological differentiation during growth must not be regarded as an effect of cell-divisions, and from such cases as these the conception of the cell experienced a notable expansion.
Moreover, Nägeli was not satisfied with seeking instructive examples for general morphological axioms in the lower Cryptogams; he devoted special study to the Algae for systematic and descriptive purposes; and his 'Neuen Algensysteme,' which appeared in 1847, and 'Gattungen einzelliger Algen,' of 1849, were the first successful attempts to substitute serious investigation for the mere zeal of the collector in this part of the vegetable kingdom, which had not indeed been hitherto neglected, but had not been systematically worked since the time of Vaucher. In the same spirit Alexander Braun also in his 'Verjüngung' contributed a rich material of new observations on the mode of life of the Algae and the morphological conditions connected with it, and his labours were followed in the succeeding years by the important researches of Thuret, Pringsheim, De Bary, and others, to which we shall recur in a later portion of this history.
But before the examination of the Algae, and soon after of the Fungi also, led to such great results, the systematic botany of the higher plants underwent important changes through the methodical study of the embryology of the Muscineae and Vascular Cryptogams. These groups had been frequently and carefully examined by good observers since the last century, and the systematists, without penetrating deeply into the peculiarities of their organisation, had brought the species and genera, the families and even the higher divisions into tolerable order. Comprehensive and methodically arranged catalogues of these plants had been formed, and attempts had been made to explain their morphology by that of the Phanerogams; Schmidel published valuable observations on the Liverworts in the year 1750, Hedwig especially on the Mosses in 1782; these works were followed by Mirbel's thorough examination of Marchantia in 1835, by Bischoff's of Marchantieae and Riccieae, by Schimper's study of the Mosses in 1850, and by Lantzius Beninga's contributions to the knowledge of the structure of the moss-capsule in 1847. The organisation, and to some extent the germination, of the Vascular Cryptogams had become better known since 1828 through Bischoff's researches; Unger had as early as 1837 described the spermatozoids in the antheridia of various Mosses, Nägeli had discovered them on an organ of the Ferns which had up to that time been taken for the cotyledonary leaf of these plants, and on the same part of the plant Suminski in 1848 observed the female sexual organs and the entrance of the spermatozoids into them. The history of the germination of the Rhizocarps, from which Schleiden thought that he had proved his erroneous theory of fertilisation with more than usual certainty, had been examined some years before by Nägeli, and also by Mettenius, in great detail; here too Nägeli detected the spermatozoids. Thus important fragments of the life and organisation of these plants had been described up to the year 1848, but until they were more fully understood and connected together they had but little scientific value, the one fact perhaps excepted, that fertilisation in the Cryptogams as in animals was effected by spermatozoids. A perfect insight into the embryological conditions in question could only be obtained when the embryology of the Phanerogams especially had been cleared up, for according to Schleiden's theory, which made the pollen-tube enter the embryo-sac in the ovule and develop into the embryo, the ovule was no longer to be regarded as a female sexual organ, but only as a place of incubation for the embryo, which was thus really produced asexually. This important question was set at rest by Wilhelm Hofmeister's work, 'Die Entstehung des Embryos der Phanerogamen,' which appeared in 1849. In this work, and in a series of subsequent treatises, he showed that the egg-cell is formed in the embryo-sac before fertilisation, and that it is this which is excited to further development by the appearance of the pollen-tube, and produces the embryo. Hofmeister had observed the organisation of the ovule, the nature of the embryo-sac and of the pollen-grain, and the formation of the embryo from the fertilised egg-cell step by step and cell by cell, and his account of these processes was aided by the light which Nägeli's theory of the cell, and his reference of all processes of development to the processes of cell-formation, had thrown upon the history of development. He went on to apply the same method to the study of the embryology of the Muscineae and the Vascular Cryptogams, and followed the development of the sexual organs cell by cell in a large number of species; he observed the origination of the egg-cell which was to be subsequently fertilised, and the formation of spermatozoids, and above all he showed the divisions which take place in the fertilised egg-cell, and the relation of its segments to the further growth of the sexual product in course of formation. The whole course of development in the Muscineae and Vascular Cryptogams displayed a return twice repeated to the single cell as the starting-point in each case of a new phase of development; the true relation between the asexually produced spore and its germ-product on the one side, and the sexually generated embryo on the other, and their significance in the history of development, were brought out clearly by Hofmeister's investigation, while the exactness of his method rendered lengthy discussions on the subject unnecessary. With these embryological processes, especially those of the Rhizocarps and Selaginellae, in which the presence of two kinds of spores was now for the first time correctly interpreted, Hofmeister compared the embryology of the Conifers, and by their aid that of the Angiosperms also.
The results of the investigations published in the 'Vergleichende Untersuchungen' in 1849 and 1851 were magnificent beyond all that has been achieved before or since in the domain of descriptive botany; the merit of the many valuable particulars, shedding new light on the most diverse problems of the cell-theory and of morphology, was lost in the splendour of the total result, which the perspicuity of each separate description revealed to the reader before he came to the conclusion of the work, and there a few words in plain and simple style gave a summary of the whole. Briefly to describe this result in all its importance for botanical science is a difficult task; the idea of what is meant by the development of a plant was suddenly and completely changed; the intimate connection between such different organisms as the Liverworts, the Mosses, the Ferns, the Equisetaceae, the Rhizocarps, the Selaginellae, the Conifers, the Monocotyledons, and Dicotyledons could now be surveyed in all its relations with a distinctness never before attained. Alternation of generations, lately shown to exist though in quite different forms in the animal kingdom, was proved to be the highest law of development, and to reign according to a simple scheme throughout the whole long series of these extremely different plants. It appeared most clearly in the Ferns and Mosses, though at the same time with a certain difference in each; in the Ferns and allied Cryptogams a small inconspicuous body grows out of the asexually produced spore, and immediately produces the sexual organs; from the fertilisation of these organs proceeds the root-bearing and leafy stem of the Fern, which in its turn again produces only asexual spores. In the Muscineae, on the other hand, a much differentiated and usually long-lived plant is developed from the spore, and this plant proceeds again after some time to form sexual organs, the product of which is the so-called Moss-plant. The first generation that arose from the spore, the sexual, is in the Muscineae the vegetative plant, while in the Ferns and their allies the whole fulness of vital activity and of morphological differentiation is unfolded in the second generation which is asexually produced. Here all was at once clear and obvious; but Hofmeister's researches also showed that the same scheme of development holds good in the Rhizocarps and Selaginellae where two kinds of spores are formed; and it appeared plainly from their case that the recognition of the true relation between the production of spores and sexual organs is the guide to the morphological interpretation. When the processes in the large female spore of the most perfect of the Cryptogams was known, the formation of the seeds in the Conifers was at once understood; the embryo-sac in these answered to this large spore, while the endosperm represented the prothallium, and the pollengrain the microspore; the last trace of alternation of generations, so obvious in the Ferns and Mosses, was seen in the formation of the seed in the Phanerogams. The changes, which the alternation of generations passes through from the Muscineae upwards to the Phanerogams, were, if possible, still more surprising than the alternation of generations itself. The reader of Hofmeister's 'Vergleichende Untersuchungen' was presented with a picture of genetic affinity between Cryptogams and Phanerogams, which could not be reconciled with the then reigning belief in the constancy of species. He was invited to recognise a connection of development which made the most different things appear to be closely united together, the simplest Moss with Palms, Conifers, and angiospermous trees, and which was incompatible with the theory of original types. The assumption that every natural group represents an idea was here quite out of place; the notion entertained up to that time of what was really meant by the natural system had to be entirely altered; it could as little pass for a body of Platonic ideas as for a mere framework of conceptions. But the effect of the work was great in respect to the system also; the Cryptogams were now the most important objects in the study of morphology; the Muscineae were the standard by which the lower Cryptogams must be tried, the Ferns were the measure for the Phanerogams. Embryology was the thread which guided the observer through the labyrinth of comparative and genetic morphology; metamorphosis now received its true meaning, when every organ could be referred back to its parent-form, the staminal and carpellary leaves of the Phanerogams, for example, to the spore-bearing leaves of the Vascular Cryptogams. That which Häckel, after the appearance of Darwin's book, called the phylogenetic method, Hofmeister had long before actually carried out, and with magnificent success. When Darwin's theory was given to the world eight years after Hofmeister's investigations, the relations of affinity between the great divisions of the vegetable kingdom were so well established and so patent, that the theory of descent had only to accept what genetic morphology had actually brought to view.
So gorgeous a picture as Hofmeister had designed of the genetic connection of the vegetable kingdom, except the Thallophytes, could not possibly be completely perfect and correct in all its separate features; there were still many gaps to fill up and particular observations to correct. Hofmeister himself continued his labours; the remarkable genera Isoetes and Botrychium were in the following years more carefully studied by himself, the fertilisation and embryology of the Equisetaceae by himself and Milde, and those of Ophioglossum by Mettenius, and all were fitted into their place in the system. To the present day it is always a profitable task to submit the different forms of the Muscineae, the Vascular Cryptogams, and the Gymnosperms to exact investigation in order to ascertain all the details in the process of development in these plants, the formation of the embryo, the succession of cells at the apex, the first appearance and further growth of the lateral organs; and the more careful the observation, the more clearly even to its farthest results does the correctness of the alternation of generations asserted by Hofmeister everywhere appear. It does not fall within the limit of this history to pursue the subject further, and to show how the doctrine of alternation of generations and the knowledge of the morphology of the Cryptogams were further advanced by later and distinguished researches, such as those of Cramer on the Equisetaceae, of Pringsheim on Salvinia (1862), of Nägeli and Leitgeb on the formation of roots in the Cryptogams, of Hanstein on the germination of the Rhizocarps, and of others.
The method of investigation which starts from the first steps towards the formation of the embryo before and after fertilisation, and follows the advancing segmentation and growth through all the stages of development up to the final completion of the embryo-plant, has led since 1850 in the case of the Muscineae, Vascular Cryptogams, and Phanerogams to great certainty in the morphological explanation of the organs, while the determination of affinities has ceased to be arbitrary and insecure; the way was now known which would lead to the desired end, whenever it was sought to establish the affinities of a genus of Cryptogams or of the larger groups of Phanerogams; the day of ingenious guessing and trying was over; the only plan was patient investigation, and this always yielded a result of lasting value.
The case was quite different with the Thallophytes still in 1850; what was certainly known about them only showed how uncertain the rest was; the Algae, Fungi, and Lichens presented a chaotic mass of obscure forms in contrast with the well-ordered knowledge of the Muscineae and Vascular plants. In the Mosses and Ferns the series of developments within the limits of the species was so set forth in its several stages, that all the important points in the advancing growth were clearly ascertained, while the alternation of generations at once sharply distinguished and connected together the chief sections in the development; on the other hand the development of the Algae and Fungi seemed to break up into a disorderly and motley throng of forms that appeared and disappeared, and it seemed scarcely possible to discover their regular genetic connection. Here the important point was to determine which of the known forms belonged to one and the same cycle of development, for these plants go back at the most various stages of development to the segregation of single cells, which are the beginning of a new development either repeating or carrying on the old one. The beginnings of the most different species of Algae lay mixed up together in the same drop of water, those of quite different Fungi grew together and even upon one another on the same substratum; in the Lichens, Fungus and Alga were united together. Such was the case with the small and microscopic species; the large Seaweeds, the Mushrooms, and the large Lichens were easier to distinguish specifically, but less if possible was known of their development than of that of the microscopic Thallophytes.
Nevertheless the knowledge of individual forms in these organisms had been considerably extended before 1850. Collectors and amateurs, intent only on determining what is immediately presented to the eye and making little enquiry into origin and affinities, were indefatigable in adding to their collections, and made catalogues and proposed various systems founded on external marks taken at pleasure. The names of species were counted by thousands, their characters filled thick volumes and the figures large folios; the abundance of forms in the Thallophytes proved to be so great that many botanists devoted their whole attention to them, many collected and described only the Algae, others only the Fungi and Lichens. It is true that a deeper insight into the connection of these forms of life with one another and with other plants was not to be obtained in this way; still an empirical basis was formed for a knowledge of the Cryptogams, such as had been established for the Phanerogams by the herbals of the 17th century. All forms open to observation were named and arranged in one way or another; and there was no difficulty in understanding what form was meant, when names, or tables and figures, were cited from the various books. Of such works, those of Agardh, Harvey, and Kützing on the Algae, those of Nees von Esenbeck, Elias Fries, Léveillé, and Berkeley on the Fungi, and especially Corda's elaborate work on the latter plants are the most valuable.
The views entertained on the subject of the development and propagation of the lower Cryptogams down to the year 1850 were very uncertain and fluctuating. In some Algae, Fungi, and Lichens certain organs of multiplication and propagation were known, in others they were quite unknown; some forms appeared in places and under circumstances which seemed to necessitate the assumption of spontaneous generation; in 1827 Meyen declared that the small Algae, known as 'Priestley's matter,' which are formed in stagnant water and even in closed vessels, are produced by free generation, and Kützing endeavoured to show this by experiment in 1833; some Fungi were regarded as diseased growths from other organisms, many were supposed to spring up spontaneously, though they might be capable at the same time of propagating themselves by spores; this view was shared by even the best botanists with regard to the most simple Fungi up to 1850. But the systematic investigation of the Algae and Fungi was as little hindered by the notion of spontaneous generation after 1850 as that of Phanerogams had been in the 17th century by the same notion; it was however at first affected by the view put forth by Hornschuch in 1821 and by Kützing in 1833, that the simplest of all Alga-cells (Protococcus and Palmella), once produced spontaneously, could develop according to circumstances into a variety of Algae, and even of Lichens and Mosses; as some observers even now consider Penicillium and Micrococcus to be the starting-points of very different Fungi. There was a difficulty also in drawing the boundary-line between the lower animals and plants; the difficulty was solved by classing all objects capable of independent movement with animals; thus whole families of Algae (the Volvocineae, Bacillariaceae, and others) were claimed by the zoologists, and when the swarmspores of a genuine Alga were seen for the first time in the act of escaping, the phenomenon was described as the changing of the plant into an animal. Trentepohl in 1807, and Unger in 1830, explained in this way the escape of the zoospores of Vaucheria. The remarkable thing is, not that such views were entertained, but that the majority of botanists combined with them a belief in the constancy of species. But this dogma rendered good service to the science in this instance, for the botanists, who at a later time applied themselves to the systematic examination of the Algae and Fungi, confided in the constancy of the processes of development in each species, which they expected would assert itself in these forms as in the Mosses and higher plants.
With much that was obscure and doubtful, the result of occasional observation accompanied by uncritical interpretation, the literature of the subject had contained for some time a certain number of single well-established facts of real importance, which were well adapted to serve as starting-points for earnest and exact investigation. Among the Algae the genera Spirogyra and Vaucheria especially had supplied remarkable phenomena; Joseph Gärtner observed the formation of zygospores in Spirogyra in 1788, Hedwig saw in the mode of their production at least a suggestion of sexuality (1798), and Vaucher, in his 'Histoire de Conferves d'eau douce,' which appeared in 1803 and was far in advance of its time, called conjugation distinctly a sexual process; the optical means at his disposal did not enable him to observe the fertilisation in Vaucheria (Ectosperma), which was named after him, though he described the sexual organs accurately; the movement also of the zoospores in this genus escaped him, and Trentepohl first observed their escape and swarming in 1807. Vaucher had also observed the formation of new nets in the old cells of Hydrodictyon, and Areschoug repeated the observation in 1842, when he saw the swarming of young cells in the old ones. Bischoff, as early as 1828, saw the spermatozoids of Chara, though without understanding them. Observations on conjugating Algae were multiplied; Ehrenberg in 1834 saw corresponding phenomena in Closterium, and Morren described them more exactly in 1836. The formation of swarmspores in fresh-water and salt-water Algae was frequently observed between 1820 and 1830, and in his 'Neues System,' iii, which appeared in 1839, Meyen gave a summary of all that was known up to that time of the propagation of the Algae. But a new aspect was given to the knowledge of the Algae by those researches of Nägeli between the years 1844 and 1849, which have been already mentioned, and which are the first since Vaucher's time that can be regarded as systematic. Nägeli studied especially the laws of cell-division in sexual multiplication and growth, but he considered the Florideae to be the only Algae that were sexually differentiated, and distinguished the rest as being without sexuality. Braun in his 'Verjüngung' (1850) made numerous contributions to the biology of the fresh-water Algae, affording many and most interesting glimpses into a connection still little understood between these forms; and in 1852 he gave an account of the history of growth in the Characeae, a work conceived in Nägeli's spirit and a model of scientific research, in which the mode of derivation of every cell from the apical cell of the stem was shown, the sexual organs were minutely examined, and the relation established between the direction of the 'streaming' of the cell-contents and the morphology of the organs. Gustav Thuret had already made the zoospores of the Algae the subject of detailed examination. Such was the condition of affairs with respect to the Algae about the year 1850, when Hofmeister made the formation of the embryo in the Phanerogams, the Vascular Cryptogams, and the Muscineae the central point of investigation in morphology and systematic botany. He made it clear that a perfect insight into the whole cycle of development in the plant and into its affinities can only be obtained, if we succeed in making its sexual propagation, the first commencement of the embryo, the starting-point of the investigation. It was natural to expect as happy results from the embryology of the Algae, as had been obtained in the case of the higher plants; it was important therefore, that the observer should no longer rest satisfied with a knowledge of the sexual multiplication of the Algae; he must enquire into their asexual propagation, and by its aid discover the complete history of their development. Former observations suggested the probability that here too sexual propagation is the prevailing rule; but it was easy to foresee that it would be a task of great labour to make out a connected history of development, a task of which the collectors who liked to call themselves systematists had never formed a conception; but Nägeli's and Hofmeister's researches had made botanists familiar with the highest demands of this kind, and the men who were to gain new conquests for genuine science were already engaged in the work in 1850. A splendid result appeared in 1853, in Thuret's account of the fertilisation of the genus Fucus; this was a simple process as a matter of embryology; but the sexual act was so clear, and even open to experimental treatment, that it threw light at once upon other cases more difficult to observe. Then followed discoveries of sexual processes in rapid succession; Pringsheim solved the old enigma in Vaucheria in 1855, and between 1856 and 1858 in the Oedogonieae, Saprolegnieae and Coleochaetae; in 1855 Cohn observed the sexual formation of spores in Sphaeroplea. Pringsheim however was not content with carefully observing the sexual act; he gave detailed descriptions of growth in the same families in its progress cell by cell, of the formation of the sexual organs, and the development of the sexual product. The asexual propagations which are intercalated into the vegetation and embryology were shown in their true connection. Processes were recognised which often recalled the alternation of generations in the Muscineae; it was shown that very different forms of sexuality and of general development occur in the Algae, and these led to the formation of systematic groups, quite different from those founded on the superficial observation of collectors. It soon appeared in the Algae, and later in the Fungi and Lichens, that special investigation must lay new foundations for the system. From the confused mass of forms not before understood, Pringsheim brought out a series of characteristic groups, which, thoroughly examined and skilfully described in words and by figures, stood out as islands in the chaotic sea of still unexamined forms, and threw light in many ways on all around them. In like manner the morphology of the Conjugatae was thoroughly examined by De Bary before 1860; fragments of the history of development in the Algae were added by Thuret, and he and Bornet cleared up the remarkable embryology of the Florideae in 1867, while Pringsheim established the pairing of the swarm-spores in the Volvocineae in 1869. The Algae offer at present a greater variety in the processes of development than any other class of plants; sexual and asexual propagation and growth work one into the other in a way which opens entirely new glimpses into the nature of the vegetable world.
The old conceptions of the nature of plants had been greatly modified by Hofmeister's discovery of the alternation of generations, and the reduction to it of the formation of the seed in Phanerogams; in like manner the first beginnings of plant-life, the simplest forms of Algae, exhibit phenomena, which compel us to revise our fundamental conceptions of morphology, if we are ever to be able to give a systematic view of the whole vegetable kingdom.
The methodical examination of the Fungi after 1850 led to similar but still more comprehensive results. From earliest times the Fungi had been objects of wonder and superstition; what Hieronymus Bock said of them has been told in the first chapter; this was repeated by Kaspar Bauhin, and similar notions existed till late into our own century; about the middle of the 17th century Otto Von Münchausen thought that mushrooms were the habitations of Polypes, and Linnaeus assented to that view. What the nature-philosophers, as Nees von Esenbeck for instance, had to say on the nature of Fungi, need not be reproduced here. Still some useful observations had been accumulating for some time on this subject; as early as 1729 Micheli had collected the spores of numerous Fungi, had sown them and obtained not only mycelia but also sporophores (fructifications), and Gleditsch confirmed these observations in 1753; Jacob Christian Schaeffer about the year 1762 published very good figures of all the Fungi of Bavaria and the Palatinate, and collected the spores of many species. Yet Rudolphi and Link at the beginning of the present century ventured to deny the germination of the spores of Fungi; Persoon in 1818 thought that some Fungi grow from spores, others from spontaneous generation. A decided improvement appears after 1820 in the views of botanists with respect to Fungi, and to this Ehrenberg's elaborate essay, 'De Mycetogenesi,' published in that year in the Leopoldina, contributed greatly. In that work he collected together all that was then known on the nature and propagation of the Fungi, and communicated observations of his own on spores and their germination; he gave figures also of the course of the hyphae in large sporophores and in other parts, but his most important service was a description of the first observed case of sexuality in a Mould, namely, the conjugation of the branches of Syzygites. In the same year Nees von Esenbeck sowed Mucor stolonifer on bread, and obtained ripe sporangia in three days (Flora, 1820, p. 528); Dutrochet proved in 1834 (Mém. ii. p. 173) that the larger Fungi are only the sporophores of a filiform branching plant, which spreads usually under ground or in the interstices of organic substances, and had been till that time regarded as a peculiar form of Fungus under the name of Byssus. Soon after, Trog (Flora, 1837, p. 609) carried these observations further; he distinguished the mycelium from the sporophore, and pointed out that the former is often perennial and is the first product of the germinating spores. He made an attempt to examine the morphology of the larger sporophores, and showed that it was possible to collect the spores of mushrooms on paper, and that those of Peziza and Helvella are forcibly ejected in little clouds of dust; he also produced new proofs of Gleditsch's statement, that the spores of Fungi are disseminated everywhere by the air. Schmitz published in 'Linnaea,' between the years 1842 and 1845 excellent observations on the growth and mode of life of several of the larger Fungi. It was not unnecessary at that time to make it clearly understood that the spores of Fungi reproduce their species exactly.
But the lower, the small and simple Fungi, those especially which are parasitic on plants and animals, were the most attractive objects in the whole field of mycology. Here were difficulties in abundance, here were the darkest enigmas with which botany has ever had to deal, here was new ground to be slowly won by extreme scientific circumspection and foresight. In these forms, as in the Algae, the first thing to be done was to make out the complete history of development in a few species; but it was much more difficult in the Fungi than in the Algae to discover what properly belonged to one cycle of development, and to separate it from casual phases of development of other associated Fungi. The merit of first breaking ground in this direction belongs to the brothers Tulasne, who published before 1850 the first more exact researches into the Smuts and Rusts; these were followed by a long series of excellent works on different forms of Fungi, especially the subterranean, whose mode of life and anatomy were described and illustrated by splendid figures; but their account of the development of Ergot of rye (1853), their further investigations into the formation of the spores and the germination of Cystopus, Puccinia, Tilletia, and Ustilago, and their discovery of the sexual organs in Peronospora before 1861, were of greater theoretical importance. The 'Selecta Fungorum Carpologia,' which appeared in three volumes from 1861 to 1865 with fine figures, some of which represented the process of development, contributed greatly to the reformation of mycology. Meanwhile, Cessati had published investigations into the Muscardine-fungus of the silkworm-caterpillar, and Cohn into a remarkable Mould, the Pilobolus.
But mycology owes its present form to none more than to Anton De Bary, whose writings, the fruit of twenty years' labour, it would take too much space to enumerate one by one. With a correct understanding of the only means which can lead to sure results in this difficult branch of study, De Bary made it his first endeavour to perfect the methods of observation, and not only sought for the stages of development of the lower Fungi in their natural places of growth, but cultivated them himself with all possible precautions, and thus obtained complete and uninterrupted series of developments. By these means he succeeded in proving that parasitic Fungi make their way into the inside of healthy plants and animals, and that this is the explanation of the remarkable fact, that Fungi live in the apparently uninjured tissue of other organisms, a fact which formerly had led to the supposition that such Fungi owe their origin to spontaneous generation, or to the living contents of the cells of their entertainers. Pringsheim had already observed these occurrences in 1858 in the case of an unusually simple water-fungus (Pythium). De Bary showed that the intrusive parasite vegetates inside the plant or animal which is its host, and afterwards sends out its organs of propagation into the open air, and that at a given time the organism attacked by the fungus sickens or dies. These investigations were not only of high scientific interest to the biologist, but they produced a series of results of the greatest importance to agriculture and forestry, and even to medicine.
With the Fungi, even more than with the Algae, the chief difficulty in making out a complete series of developments in the history of each species arose from the frequent intercalation of the asexual mode of multiplication into the course of its development, and in the further peculiarity, that the several stages of development in some cases could only be completed on different substrata. One of the most important tasks was to find the sexual organs, the existence of which was rendered probable by various analogies, and after De Bary had observed the sexual organs in the Peronosporeae in 1861, he succeeded in 1863 in proving for the first time that the whole fruit-body of an Ascomycete is itself the product of a sexual act, which takes place on the threads of the mycelium.
The literature of mycology based on De Bary's methods of observation and its actual results has been enriched by others also in various directions since 1860; in the case of the Fungi, as in that of the Algae, it is not possible yet to see to what results investigation will ultimately lead; but it is one of the fairest fruits of strictly inductive method, that it has succeeded in smoothing this thorny and indeed perilous route, where the enquirer is constantly in danger of being misled, and in satisfying the severest demands of science. Conclusions have been already reached that are important for morphology and systematic botany, and among these the establishment of the nature of the large sporophores, and of processes similar to the alternation of generations in the higher Cryptogams should be especially mentioned. But the most important result remains to be told; it is, that the two classes of Algae and Fungi, hitherto kept strictly separate, must obviously be now united, and an entirely new classification adopted, in which Algae and Fungi recur as forms differing only in habit in various divisions founded on their morphology.
A few words must be given here to the Lichens. They are the division of the Thallophytes, whose true nature was last recognised, and that only in modern times; till after 1850 scarcely more was known of their organisation than Wallroth had discovered in 1825, namely, that green cells, known as gonidia are scattered through the fungus-like hyphal tissue of the thallus. After Mohl's investigations in 1833, it was known that free spores were formed in the tubes of the fructifications (apothecia), and that a dust collected from the thallus and consisting of a mixture of gonidia and hyphae was in a condition to propagate the species. The genetic relation between the chlorophyll-containing gonidia and the fungus-like hyphae long continued to be obscure, till at last, after 1868, it was shown that the gonidia are true Algae, and the hyphal tissue a genuine Fungus, and that therefore the Lichens are not a class co-ordinating with the Algae and Fungi, but a division of Ascomycetes, which have this peculiarity, that they spin their threads round the plants on which they feed, and take them up into their tissue. De Bary suggested this explanation, but it was Schwendener who adopted it without reserve and openly declared it, as much to the surprise as the annoyance of Lichenologists. It may be foreseen that their opposition will yield to the weight of facts, which already leave no doubt in the minds of the unprejudiced.
Thus researches in the domain of the Thallophytes have led during the last twenty years to a complete revolution in the views entertained with respect to the nature of these organisms, and enriched botany with a series of surprising achievements; and the movement there is still far from having come to an end. But we must regard it as one of the great results for the whole science that through the examination of the lower and higher Cryptogams, morphology and systematic botany have been rescued from many ancient prejudices, that the survey has become freer, the methods of investigation surer, the questions more clearly seen and put in more definite form.
- See A. Bayer, 'Leben und Wirken F. Unger's,' Gratz (1872), p. 52.
- See Darwin's repudiation of this statement on p. 421 of Ed. 6 of the 'Origin of Species.'
- Casimir Christoph Schmidel was born in 1718 and died in 1792; he was Professor of Medicine in Erlangen, and was the first who described the sexual organs in various Liverworts.
- Lantzius Beninga, born in East Friesland in 1815, was a professor in Göttingen, and died in 1871.
- Gottlieb Wilhelm Bischoff was born at Dürkheim on the Hardt in 1797, and died as Professor of Botany at Heidelberg in 1854. He wrote various manuals and text-books which are careful and industrious compilations, but being entirely conceived in the spirit of the times preceding Schleiden they are now obsolete; his investigations however into the Hepaticae, Characeae, and Vascular Cryptogams, illustrated by very beautiful drawings from his own hand, are still of value; and the same may be said of his 'Handbuch der botanischen Terminologie und Systemkunde' on account of its numerous figures.
- Karl Adolf Agardh (1785-1859) was until 1835 Professor in Lund, afterwards Bishop of Wermland and Dalsland. Jacob Georg Agardh, born in 1813, was Professor in Lund. William Henry Harvey (1811-1866) was Professor of Botany in Dublin. Friedrich Traugott Kützing, born in 1807, was Professor in the Polytechnic School of Nordhausen.
- C. G. Nees von Esenbeck published his 'System der Pilze und Schwämme' in 1816; Th. F. L. Nees von Esenbeck, in conjunction with A. Henty, a 'System der Pilze' in 1837. The first (1776-1858) was for a long time President of the Leopoldina, Professor of Botany in Breslati, and one of the chief representatives of the nature-philosophy. Elias Fries, born in 1794, became Professor of Botany in Upsala in 1835; he died in 1878. Léveillé (1796-1870) was a physician in Paris. August Joseph Corda was born at Reichenberg in Bohemia in 1809, and became custodian of the National Museum in Prague in 1835; he undertook a journey to Texas in 1848, from which he never returned, having probably perished by shipwreck in 1849. Weitenweber, in the 'Abhandlungen der Böhmischen Gesellschaft der Wissenschaft,' Bd. 7, Prag, 1852, gives a full account of this eminent mycologist. Corda was the first who thoroughly applied the microscope to copying and describing every form of Fungus that was known to him, and especially the minuter ones. His 'Icones Fungorum hucusque cognitorum' (1837-1854) are still an indispensable manual in the study of the subject.
- Jean Pierre Étienne Vaucher, the instructor and friend of P. de Candolle, was a minister and professor in Geneva.
- Trentepohl's communication is to be found in the 'Botanische Bermerkungen und Berichtigungen' of A. W. Roth, Leipsic, 1807.
- Pier' Antonio Micheli, born at Florence in 1679, was Director of the Botanic Garden there, and died in 1737. Johann Jacob Dillen (Dillenius), born in Darmstadt in 1687, was Professor of Botany in Oxford, and died in 1747. These two botanists were the first who submitted the Mosses and the lower Cryptogams to scientific examination, and endeavoured to prove the presence of sexual organs in these plants.
- Jacob Christian Schaeffer, born in 1718, was Superintendent in Regensburg; he died in 1790.
- See Sachs, 'Lehrbuch der Botanik,' ed. 4 (1874), p. 245.
- Fr. Wilh. Wallroth, born in the Harz in 1792, was district physician at Nordhausen. He died in 1857. See 'Flora' for 1857, p. 336.