History of botany (1530–1860)/Book 1/Chapter 3
Development Of the Natural System Under the influence Of the Dogma Of the Constancy Of Species.
From the year 1750 Linnaeus' terminology of the organs of plants and his binary method of naming species came into general use; the opposition which his doctrines had till then encountered by degrees died away, and if all that he taught was not universally accepted, his treatment of the art of describing plants soon became the common property of all botanists. But in course of time two very different tendencies were developed; most of the German, English, and Swedish botanists adhered strictly to Linnaeus' dictum, that the merit of a botanist was to be judged by the number of species with which he was acquainted; they accepted Linnaeus' sexual system as one that completed the science in every respect; they thought that botany had reached its culminating point in Linnaeus, and that any improvement or addition could only be made in details, by continuing to smooth over some unevennesses in the system, to collect new species and describe them. The inevitable result was that botany ceased to be a science; even the describing of plants which Linnaeus had raised to an art became once more loose and negligent in the hands of such successors; in place of the morphological examination of the parts of plants there was an endless accumulating of technical terms devoid of depth of scientific meaning, till at length a text-book of botany came to look more like a Latin dictionary than a scientific treatise. In proof of this we may appeal to Bernhardi's 'Handbuch der Botanik,' published at Erfurt in 1804, and Bernhardi was one of the best representatives of German botany of the time. How botany, especially in Germany, gradually degenerated under the influence of Linnaeus' authority into an easy-going insipid dilettantism may very well be seen from the botanical periodical, entitled 'Flora,' the first volumes of which cover the greater part of the first fifty years of the 19th century; it is scarcely conceivable how men of some cultivation could occupy themselves with such worthless matter. It would be quite lost labour to give any detailed account of this kind of scientific life, if it can be so called, this dull occupation of plant-collectors, who called themselves systematists, in entire contravention of the meaning of the word. It is true indeed that these adherents of Linnaeus did some service to botany by searching the floras of Europe and of other quarters of the globe, but they left it to others to turn to scientific account the material which they collected. But before this evil had spread very widely, a new direction to the study of systematic botany and morphology was given in France, where the sexual system had never met with great acceptance. Bernard de Jussieu and his nephew, Antoine Laurent de Jussieu, taking up Linnaeus' profounder and properly scientific efforts, made the working out of the natural system, in Linnaeus' own opinion the highest aim of botany, the task of their lives. Here more was needed than a perpetual repetition of descriptions of single plants after a fixed pattern; more exact inquiries into the organisation of plants, and especially of the parts of the fructification, must supply the foundation of larger natural groups. It was a question therefore of new inductive investigation, of real physical science, of penetrating into the secrets of organic form, whereas the botanists who confined themselves to Linnaeus' art of description made no new discoveries respecting the nature of plants. And if these men held to the dictum just quoted from Linnaeus, and therefore regarded themselves as his genuine disciples, the founders of the natural system had as good a right to the title, not because they followed his nomenclature and method of diagnosis, but because they strove after exactly that object which he had placed first in the science, the construction of the natural system; they were really the men whom he had meant when he spoke of 'methodici' and 'systematici.' The German, English, and Swedish collectors of plants adhered to the less profound, every-day, practical precepts of their master; the founders of the natural system followed the deeper traces of his knowledge. This direction proved to be the only one endowed with living power, the true possessor of the future.
The efforts of Jussieu, Joseph Gärtner, De Candolle, Robert Brown, and their successors up to Endlicher and Lindley, are not marked only by the fact that they did truly seek to exhibit the gradations of natural affinities by means of the natural system; equally characteristic of these men is their firm belief in the dogma of the constancy of species as defined by Linnaeus. Here at once was a hindrance to their efforts; the idea of natural relationship, on which the natural system exclusively rests, necessarily remained a mystery to all who believed in the constancy of species; no scientific meaning could be connected with this mysterious conception; and yet the farther the inquiry into affinities proceeded, the more clearly were all the relations brought out, which connect together species, genera, and families. Pyrame de Candolle developed with great clearness a long series of such affinities as revealed to us by comparative morphology, but how were these to be understood, so long as the dogma of the constancy of species severed every real objective connection between two related organisms? Little indeed could be made of these acknowledged affinities; still, in order to be able to speak of them and describe them, recourse was had to indefinite expressions, to which arbitrary and figurative meanings could be assigned. Where Linnaeus had spoken of a class plant or generic plant, the expression 'plan of symmetry' or 'type' was used, meaning an ideal original form, from which numerous related forms might be derived. It was left undecided, whether this ideal form ever really existed, or whether it was merely the result of intellectual abstraction; and thus the forms of thought of the old philosophy soon began to reappear. The Platonic ideas, though mere abstractions and therefore only products of the understanding, had been regarded not only by the school of Plato, but also by the so-called Realists among the schoolmen, as really existing things. The systematists obtained the idea of a type by abstraction, and the next step was easy, to ascribe with the Platonists an objective existence to this creature of thought, and to conceive of the type in the sense of a Platonic idea. This was the only view that was possible in combination with the dogma of the constancy of species, and so Elias Fries, in his 'Corpus Florarum,' 1835, in speaking of the natural system, could consistently say, 'est quoddam supranaturale,' and maintain that each division of it 'ideam quandam exponit.' So long as the constancy of species is maintained, there is no escaping from the conclusion drawn by Fries, but it is equally certain that systematic botany at the same time ceases to be a scientific account of nature. Systematists, adopting this conclusion as necessarily following from the dogma, might consider themselves as seeking to express in the natural system the plan of creation, the thought of the Creator himself; but in this way systematic botany became mixed up with theological notions, and it is easy to understand why the first feeble attempts at a theory of descent encountered such obstinate, nay, fanatical opposition from professed systematists, who looked upon the system as something above nature, a component part of their religion. And if we look back we find that these views are based on the dogma of the constancy of species, while Linnaeus' 'Philosophia Botanica' teaches us on what grounds this dogma rests, where it says, 'Novas species dari in vegetabilibus negat generatio continuata, propagatio, observationes quotidianae, cotyledones.'
In spite of all this one important advance was made by the successors of Jussieu; the larger groups of genera, the families, were defined with the certainty and precision, with which Linnaeus had fixed the boundaries of species and genera, and were supplied with characteristic marks. They succeeded also in clearly distinguishing various still larger groups founded on natural affinity, such as the Monocotyledons and Dicotyledons; the distinction between Cryptogams and Phanerogams was by degrees better appreciated, though this point could not be finally settled, so long as it was attempted to reduce the Cryptogams entirely to the scheme of the Phanerogams. The chief hindrance however to the advance of systematic botany, at least at the beginning of this period, lay in the defective morphology enshrined in Linnaeus' terminology and in his doctrine of metamorphosis. A great improvement certainly was effected in the early part of the 19th century by De Candolle's doctrine of the symmetry of plants, a doctrine which has been much undervalued, and that merely on account of its name; it is really a comparative morphology, and the first serious attempt of the kind since the time of Jung that has produced any great results; a series of the most important morphological truths, with which every botanist is now conversant, were taught for the first time in De Candolle's doctrine of symmetry in 1813. But one thing was wanting not only in Jussieu and De Candolle, but in all the systematists of this period, with the single exception of Robert Brown, and this was the history of development. The history of the morphology and systematic botany of this period shows indeed, that the comparison of mature forms leads to the recognition of many and highly important morphological facts; but as long as matured organisms only are compared, the morphological consideration of them is always disturbed by the circumstance that the organs to be compared are already adapted to definite physiological functions, and thus their true morphological character is often entirely obscured; on the other hand, the younger the organs are, the less is this difficulty experienced, and this is the real reason why the history of development is of so great service to morphology. It was then one of the characteristic features of the period we are describing, that its morphology was formed upon the study of matured forms; the history of development, or at all events of very early stages of development, could not be turned to account till after 1840, for skill in the use of the microscope, here indispensable, was not sufficiently advanced before that time to make it possible to follow the growth of organs from their first beginnings.
The establishment of natural affinities combined with the assumption of the constancy of species, the growth of comparative morphology without the history of development, lastly, the very subordinate attention still paid to the Cryptogams,—these are the special characteristics of the period which has now to be described at greater length.
Here we must once more call attention to the fact, that Linnaeus was the first to perceive that a system which was to be the expression of natural affinities could not be attained in the way pursued by Cesalpino and his immediate successors. All who have attentively studied the writings of Linnaeus which appeared after the 'Classes Plantarum' (1738) must have seen the difference between that way and the one recommended by him—a difference which is the more obvious because Linnaeus himself, like his predecessors, constructed an artificial system on predetermined principles of classification, and always employed it for practical purposes, while he published at the same time in the above-named work his fragment of a natural system, and in the preface set forth the peculiar features of the natural and artificial systems in striking contrast with one another. The first thing and the last, he says in his prefatory remarks to his fragment, which is demanded in systematic botany, is the natural method, which slighted by less learned botanists has always been highly regarded by the more sagacious, and has not yet been discovered. If, he continues, we collect the natural orders from all existing systems (up to 1738), we shall get but a small list of really allied plants, though so many systems have claimed to be natural. He had himself long laboured to discover the natural method and had found out some things that were new; but though he had not succeeded in carrying it through to a perfect work, he would continue his efforts as long as his life lasted. He makes the very important remark, that a key, that is, a priori principles of classification, cannot be given for the natural method, till all plants have been reduced to orders; that for this no a priori rule is of value, neither this nor that part of the fructification, but the simple symmetry alone (simplex symmetria) of all the parts, which is often indicated by special marks. He suggests to those who are bent on trying to find a key to the natural system, that nothing has more general value than relative position, especially in the seed, and in the seed especially the 'punctum vegetans,'—a distinct reference to Cesalpino. He says that he establishes no classes himself, but only orders; if these are once obtained, it will be easy to discover the classes. The essence of the natural system could not have been more clearly expounded in Linnaeus' time, than it is in these sentences. He established as early as 1738 sixty-five natural orders, which he at first simply numbered; but in the first edition of the 'Philosophia Botanica' in 1751, where the list is increased to sixty-seven, he gave a special name to each group; and he showed his judgment by either taking his names from really characteristic marks, or what was still better, by selecting a genus and so modifying its name as to make it serve as a general term for a whole group. Many of these designations are still in use, though the extent and content of the groups have been greatly changed. This mode of naming is an important point, because it expresses the idea, that the different genera of such a group are to some extent regarded as forms derived from the one selected to supply the name. Many of Linnaeus' orders do in fact indicate cycles of natural affinity, though single genera are not unfrequently found to occupy a false position; at all events, Linnaeus' fragment is much the most natural system proposed up to 1738, or even to 1751. It is distinguished from Kaspar Bauhin's enumeration in this, that its groups do not run into one another, but are defined by strict boundaries and fixed by names.
The Linnaean list is distinctly marked by the endeavour to make first the Monocotyledons, then the Dicotyledons, and finally the Cryptogams follow one another; that the old division into trees and herbs already rejected by Jung and Bachmann, but still maintained by Tournefort and Ray, disappears in Linnaeus' natural system will be taken for granted after what has been already said of it, and from this time forward this ancient mistake is banished for ever.
In Bernard de Jussieu's arrangement of 1759 we find some improvements in the naming, the grouping, and the succession, but at the same time some striking offences against natural affinity. He published no theoretical remarks on the system, but gave expression to his views on relations of affinity in the vegetable kingdom in laying out the plants in the royal garden of Trianon, and in the garden-catalogue. His nephew published his uncle's enumeration in the year 1789 in his 'Genera Plantarum,' affixing the date of 1759 given above. The difference between it and the Linnaean fragment does not seem sufficiently marked to make it necessary to reproduce it here. It should be noticed however that Jussieu begins with the Cryptogams, passes through the Monocotyledons to the Dicotyledons, and ends with the Conifers. Adanson's claims of priority over Bernard de Jussieu (see the 'Histoire de la Botanique' de Michel Adanson, Paris, 1864, p. 36) may be passed over as unimportant. The natural system was not advanced by Adanson to any noticeable extent; how little he saw into its real nature and into the true method of research in this department of botany is sufficiently shown by the fact, that he framed no less than sixty-five different artificial systems founded on single marks, supposing that natural affinities would come out of themselves as an ultimate product,—an effort all the more superfluous, because a consideration of the systems proposed since Cesalpino's time would have been enough to show the uselessness of such a proceeding.
The first great advance in the natural system is due to Antoine Laurent De Jussieu (1748–1836). After all that has been said no further proof is needed that he was no more the discoverer or founder of the natural system than his uncle before him. His real merit consists in this, that he was the first who assigned characters to the smaller groups, which we should now call families, but which he called orders. It is not uninteresting to note here how Bauhin first provided the species with characters, and named the genera but did not characterise them, how Tournefort next defined the limits of the genera, how Linnaeus grouped the genera together, and simply named these groups without assigning to them characteristic marks, and how finally Antoine Laurent de Jussieu supplied characters to the families which were now fairly recognised. Thus botanists learnt by degrees to abstract the common marks from like forms; the groups thus constituted were being constantly enlarged, and an inductive process was thus completed which proceeded from the individual to the more general.
It might appear that the merit of Antoine de Jussieu is rated too low, when we praise him chiefly and simply for providing the families with characters; but this praise will not seem small to those who know the difficulty of such a task; very careful and long-continued researches were necessary to discover what marks are the common property of a natural group. Jussieu's numerous monographs show with what earnestness he addressed himself to the task; and it must be added, that he was not content simply to adopt the families established by Linnaeus and by his uncle and the limits which they had assigned to them, but that he corrected their boundaries and in so doing established many new families, and was the first who attempted to distribute these into larger groups, which he named classes. But in this he was not successful. His attempt to exhibit the whole vegetable kingdom in all its main divisions, to unite the classes themselves into higher groups, was also unsuccessful, for these larger divisions remained evidently artificial. The three largest groups on the contrary, into which he first divides the world of plants, the Acotyledons, Monocotyledons, and Dicotyledons are natural; but they had been already partly marked out by Ray, afterwards by Linnaeus, and finally in Bernard de Jussieu's enumerations. Still it is the younger Jussieu's great and abiding merit, to have first attempted to substitute a real division of the whole vegetable kingdom into larger and gradually subordinate groups for mere enumerations of smaller co-ordinated groups,—an undertaking which Linnaeus expressly declared to be beyond his powers. If then Jussieu's system was far from giving a satisfactory insight into the affinities of the great divisions of the vegetable kingdom, yet it opened out many important points of view, from which they could afterwards be discovered, and it certainly became the foundation for all further advance in the natural method of classification; for this reason it is necessary to give a view of it in the following table:—
A. L. de Jussieu's System of 1789.
This table shows that Jussieu did not oppose the Cryptogams, which he calls Acotyledones, to the whole body of Phanerogams, as Ray did under the name of Imperfectae; he rather regards the Acotyledones as a class co-ordinate with the Monocotyledones and Dicotyledones; but this mistake or similar mistaken views run through all systematic botany up to 1840; the morphology founded by Nägeli and by Hofmeister's embryological investigations first showed that the Cryptogams separate into several divisions, which co-ordinate with the Monocotyledons and Dicotyledons. At the same time the use of the word Acotyledones for Linnaeus' Cryptogams shows that Jussieu overrated the systematic value of the cotyledons, because, as is seen from the introduction to his 'Genera Plantarum,' he was quite in the dark on the subject of the great difference between the spores of Cryptogamic plants and the seeds of Phanerogams. His conception of the organs of generation was essentially that of Linnaeus; hence he judged of the Cryptogams according to the scheme of the Phanerogams, and, not perceiving their peculiarities, he virtually characterised them by negative marks.
If we notice in the above table how the Phanerogams are separated into classes, it strikes us that the triple division into hypogynous, perigynous, and epigynous is repeated no less than four times; this shows that Jussieu had mistaken ideas of the value of these marks for classification, whereas the recurrence of them so often should of itself have suggested a doubt on this point. To judge of his system more exactly we must here give his series of the families, which he had already raised to the number of a hundred.
Jussieu's division of the Cryptogams and Monocotyledons offers much that is satisfactory, if we put the position of the Naiades out of sight. The grouping of the Dicotyledons on the contrary is to a great extent unsuccessful, chiefly owing to the too great importance which he attached to the insertion of the parts of the flowers, that is, to the hypogynous, perigynous, and epigynous arrangement. It is in this grouping of families into classes that the weak side of the system lies; it is utterly artificial, and the task of his successors has been to arrange the families of the Phanerogams, which were most of them well-established, and especially those of the Dicotyledons, in larger natural groups. But this could not be effected, till morphology opened new points of view for systematic botany; Jussieu, as has been already remarked, accepted Linnaeus' views of the morphology of the organs of fructification in Phanerogams, though he introduced many improvements in details. He laid greater stress on the number and relative positions of the different parts of the flower; attention to their insertion on the flowering axis, which he designated as hypogynous, perigynous, and epigynous, would have been a great step in advance, if he had not overrated its systematic value. The morphology of the fruit is very superficial in Jussieu; even the designation of dry indehiscent fruits as naked seeds recurs in his definitions, though as it happens this misconception does not cause any great disturbance. How inexact was his investigation of the organs of fructification, when they were somewhat small and obscure, is best shown by the fact that the Naiades, which are made to include Hippuris, Chara, and Callitriche, appear among the Acotyledons, and that Lemna and the Cycads are placed with the Ferns.
Jussieu explained the dictum, 'Natura non facit saltus,' to mean that the whole body of plants in its natural arrangement must exhibit a lineal series ascending from the most imperfect to the highest forms; but he does not say whether Linnaeus' comparison of the natural system to a geographical map, the countries in which answer to orders and classes, is also admissible.
His theoretical observations on the value to be given to certain marks in a systematic point of view are not attractive, and for the most part not very correct; he speaks as though some marks must have a more extensive, others a less extensive value; the perception of the fact, so far as it is true, rests entirely upon induction; that is, after the natural affinities have been already recognised to a certain extent, it becomes apparent that certain marks remain constant in larger or smaller groups; the systematist can now go on to try whether such constant marks occur in other plants also, which he had hitherto assigned to other groups, and thus put it to the test whether those marks may not be accompanied by others, which would serve to establish the affinities; that Jussieu did so proceed in defining his families admits of no doubt, but he was not himself thoroughly conscious of the fact; at all events, he did not extend this mode of proceeding, the seeking after leading marks, to the establishing of larger groups or classes, for these he founded on predetermined principles.
Jussieu's labours as a systematist were not confined to the publication of his 'Genera Plantarum'; on the contrary, his most fruitful researches began after 1802, and were continued to the year 1820, and their results appeared in a long series of monographs on different families in the Mémoires du Museum. He felt with De Candolle, Robert Brown, and later systematists, that the perfecting of the natural system depended mainly on the careful establishing and defining of families. His efforts received a new impulse from the work of a German writer, whose first volume had appeared in 1788, a year therefore before the 'Genera Plantarum,' a second following it in 1791, and a supplementary volume in 1805.
This work was JOSEPH GÄRTNER's 'De fructibus et seminibus plantarum,' in which the fruits and seeds of more than a thousand species are described and carefully figured. But almost more important than these numerous descriptions, though they offered rich material to the professed systematists, were the introductions to the first two volumes, and especially to those of 1788. They contain valuable reflections on sexuality in plants, a subject which had remained in the condition in which it was left by Camerarius (1694) till it was greatly developed by Koelreuter after 1761, and had since then been little studied, and an account of the morphology of fruits and seeds, the knowledge of which had gone back rather than advanced since the days of Malpighi and Grew. Gärtner was well qualified for this work by his unparalleled knowledge of the forms of fruits, and still more by the character of his mind. Free from Linnaeus' scholastic bias, he addressed himself to the examination of the most difficult organs of plants with as great freedom from prepossessions as exact acquaintance with the writings of others; he gives us the impression of a modern man of science more than any other botanist of the 18th century, with the exception of Koelreuter. He knew how to communicate with clearness of language and perspicuity of arrangement whatever he gathered of general importance from each investigation. Though it is easy to see that the founding of the natural system was ever before his mind as the final object of his protracted labours, he was in no eager haste to reach it; he contented himself with arranging his fruits, saying expressly that the natural system would never be founded by these means alone, though the exact knowledge of fruits and seeds supplied the most important means for decision. Thus his great work was at once an inexhaustible mine of single well-ascertained facts, and a guide to the morphology of the organs of fructification and to its application to systematic botany. The imperfections, which are to be found even in this work, are due to the circumstances of the time; in spite of Schmiedel's and Hedwig's researches into the Mosses there was still the old obscurity with regard to the organs of propagation in the Cryptogams, and this rendered a right definition of the ideas, seed and fruit, extremely difficult. But Gärtner made one great step in advance on this very point when he showed that the spores of the Cryptogams were essentially different from the seeds of Phanerogams, with which they had been hitherto compared, because they contain no embryo; he called them therefore not seeds, but gemmae. The second great hindrance to a true conception of certain characters in fruits and seeds on the part of Gärtner was the entire ignorance of the history of development which then reigned; yet even here we see an advance, if only a small one, made by him in his repeatedly going back to the young state for a more correct idea of the organs.
Above all, Gärtner put an end to the blunder of regarding dry indehiscent fruits as naked seeds, by rightly defining the pericarp as in all cases the ripened wall of the ovary, and by considering its strong or weak construction, its dry or pulpy condition, as a secondary matter. It is obvious that the whole theory of the flower was thus placed upon a better basis, since dry indehiscent fruits may come from inferior or superior ovaries. But Gärtner's theory of the seed is one of his most valuable contributions to the science. After careful consideration of the seed-envelopes, he submitted the inner portion (nucleus) enclosed by them to a searching comparative examination; he correctly distinguished the endosperm from the cotyledons, and described the variations in its form and position. This was the more needful, since Linnaeus had denied the existence of an 'albumen' in plants, which Grew had already recognised and so named; to Linnaeus it appeared to be of no use to the seed. Though Gärtner speaks of the cotyledons as uniting with the embryo to form the nucleus of the seed, yet his account shows that he regarded them as outgrowths of the embryo itself. The uncertainty which still existed in the interpretation of the parts of the seed is shown even in Gärtner by his curious notion of a 'vitellus,' which in fact takes in everything that he was unable to explain aright inside the seed; for instance, he makes the scutellum in grasses, and even the cotyledonary bodies of Zamia a vitellus, and applies the same name to the whole contents of the spores of Seaweeds, Mosses, and Ferns. In spite of the striking defects connected with this mistaken notion in his theory of the seed, his views far surpass in clearness and consistency all that had hitherto been taught on the subject. His giving the term embryo to that part of the seed which is capable of development was also an advance in respect of logic and morphology, in spite of his mistake in not admitting the cotyledons which are attached to the embryo into the conception; this, however, could easily be corrected at a later time. What Gärtner now named the embryo, had been up to his time called the 'corculum seminis,' especially by Linnaeus and Jussieu; it was evidently thought that Cesalpino's phraseology was thus retained; but he, as we have seen, understood by the words 'cor seminis' the spot where the cotyledons spring from the germ, which spot he wrongly took for the meeting-point of root and stem and the seat of the soul of the plant. And so at last after two hundred years the word disappeared from use, which might have reminded the botanist of Cesalpino's views respecting the soul of plants. A work such as Gärtner's could scarcely find a fruitful soil in Germany, where some thirty years before even Koelreuter's brilliant investigations had met with little sympathy, and Conrad Sprengel's remarkable enquiries into the relations of the structure of the flower to the insect-world in 1793 failed to be understood; Gärtner complains in the second part, published in 1791, that not two hundred copies of the first volume were sold in three years. But the work, which forms an epoch in the history of botany, was better received in France, where the Academy placed it as second in the list of the productions which in later times had been most profitable to science; there lived the man who was able to measure the whole value of such a work—Antoine Laurent de Jussieu. But even in Germany, where plant-describing was comfortably flourishing, there were not altogether wanting men who knew how to estimate both the services of Gärtner and the importance of the natural system. First among these was August Johann Georg Karl Batsch, Professor in Jena from 1761 to 1802, who published in the latter year a 'Tabula affinitatum regni vegetabilis,' with characters of the groups and families. Kurt Sprengel, who was born in 1766, and died as Professor of Botany in Halle in 1833, contributed still more to the spread of clearer views respecting the real character of the natural system and the task of scientific botany generally by numerous works, and especially by his 'Geschichte der Botanik,' which appeared in 1817 and 1818. But even this highly gifted and accomplished man agreed with the Linnaean botanists in attributing an excessive value to the describing of plants, as is shown in his history, where to exalt the merits of the old botanists he gives figures of the plants first described by them.
Meanwhile the meritorious efforts of these men were not in themselves capable of directly advancing the natural system, or of greatly increasing the number of its adherents in Germany, nor did it find general acceptance in that country till it had made considerable progress in the hands of the two foremost botanists of the time, De Candolle and Robert Brown.
Augustin Pyrame De Candolle (1778-1841) belongs to the number of those distinguished investigators of nature, who at the end of the last and the beginning of our own century made their native city Geneva a brilliant centre of natural science. De Candolle was the contemporary and fellow-countryman of Vaucher, Theodore de Saussure, and Senebier. Physics and physiology especially were being successfully cultivated at that time in Geneva, and Pyrame de Candolle was attracted to these studies; among his youthful efforts are some important investigations into the effect of light on vegetation, and the contributions which he made to vegetable physiology in his great work on that subject will be noticed in a later portion of this history. De Candolle turned his attention to all parts of theoretical and applied botany, but his importance for the history of the science lies chiefly in the direction of morphology and systematic botany, and it is this which we will now proceed to describe.
The amount and compass of De Candolle's labours as a systematic and descriptive botanist exceed those of any writer before or after him. He wrote a series of comprehensive monographs of large families of plants, and published a new edition of De Lamarck's large 'Flore Française' substantially altered and enlarged; and in addition to these and many similar works and treatises on the geographical distribution of plants, he set on foot the grandest work of descriptive botany that is as yet in existence, the 'Prodromus Systematis Naturalis,' in which all known plants were to be arranged according to his natural system and described at length, a work not yet fully completed, and in which many other descriptive botanists of the last century participated, but none to so large an extent as De Candolle, who alone completed more than a hundred families. It is not possible to give an account in few words of the service rendered to botany by such labours as these; they form the real empirical basis of general botany, and the better and more carefully this is laid, the greater the security obtained for the foundations of the whole science.
But a still higher merit perhaps can be claimed for De Candolle, inasmuch as he not only like Jussieu elaborated the system and its fundamental principles in his descriptive works, but developed the theory, the laws of natural classification, with a clearness and depth such as no one before him had displayed. To this purpose he applied morphological researches, which in profundity and wealth of thought and in the fruitfulness of their results for the whole domain of systematic botany far surpassed all that Linnaeus and Jussieu had accomplished, and show us that while engaged in his splendid labours in descriptive botany he had caught during his ten years' residence in Paris the true spirit of modern investigation of nature, as it had been developed by the French naturalists of the end of the previous century. Scarcely a trace is to be found in De Candolle of the scholasticism of Cesalpino and Linnaeus, which occasionally makes its appearance even in Jussieu. For instance, he dealt with morphology as essentially the doctrine of the symmetry of form in plants, that is, he found the basis of morphological examination in the relative position and numbers of the organs, disregarding their physico-physiological properties as of no account from the morphological point of view. He was therefore the first who recognised the remarkable discordance between the morphological characters of organs, which are of value for systematic purposes, and their physiological adaptations to the conditions of life, though it must at the same time be acknowledged, that he did not consistently carry out this principle, but committed grave offences against it in laying down his own system. It is a point of the highest interest in De Candolle's morphological speculations, that he was the first who endeavoured to refer certain relations of number and form to definite causes, and thus to distinguish what is primary and important in the symmetry of plants from merely secondary variations, as is seen in his doctrine of the abortion and adherence of organs. In these distinctions De Candolle laid the foundation of morphological views, which, though now modified to some extent, do still contain the chief elements of morphology and the natural system; but his morphological speculations were confined to the domain of the Phanerogams, and chiefly advanced the theory of the flower; a morphology of the Cryptogams was as little to be thought of in the condition of microscopy before 1820, as the application of the history of development to the establishment of morphological theories.
De Candolle published his morphology or doctrine of symmetry and his theory of classification together in a book which appeared first in 1813, with the title, 'Théorie Élémentaire de la botanique ou exposition des principes de la classification naturelle et de l'art de décrire et d'étudier les végétaux,' and again in 1819 in an improved and enlarged edition. The second edition will be the one referred to in the further account of his views. The second chapter of the second book concerns us most at present. After alluding to the fact, that anatomy and physiology are concerned with the structure of the individual organ only so far as the power to fulfil its proper function depends on the structure, he points out that the physiological point of view is no longer sufficient when we are engaged in comparing the organs of different plants. Though it is true that the function of the organs is the most important for the life and permanence of the individual, yet we find these functions modified in the case of homologous organs in different plants; for the natural classification we must take into consideration only the entire system of organisation, that is, the symmetry of the organs. All organisms of a kingdom, he continues, have the same functions with slight modifications; the immense amount of variation in systematically different species depends therefore only on the way in which the general symmetry of structure varies. This symmetry of the parts, the discovery of which is the great object in the investigation of nature, is nothing more than the sum total (l'ensemble) of the positional relations of the parts. Whenever these relations (disposition) are regulated according to the same plan, the organisms exhibit a certain general resemblance to one another, independently of the form of the organs in detail; when this general resemblance is perceived, without any attempt to give any account of it in the detail, we have what has been called habitual relationship; but it is the task of the doctrine of symmetry to resolve this likeness of habit into its elements, and to explain its causes. Without this study of symmetry it may easily happen that two different kinds of symmetry may be supposed to be alike, because they seem outwardly alike to our senses, just as forms of crystals of different systems may be confounded together for want of careful examination; the chief thing is to know the plan of symmetry in every class of plants, and the study of this is the foundation of every theory of natural affinities. But success in this study depends on the certainty with which organs are distinguished, and the distinguishing them must be independent of changes of form, size, and function. He then shows that the difficulties in the morphological comparison of organs, or, as we should now say, in the establishing the homology, are due to three causes; abortion, degeneration, and adherence (adherence). These three causes, by which the original symmetry of a class is changed and may even be utterly obscured, are then fully illustrated by examples.
In respect to abortion he distinguishes that which is produced by internal causes from that which is due to accidental and external ones; he refers especially to the abortion of two loculaments in the fruit of the horse-chestnut and the oak, to the suppression of the terminal bud in some shrubs by the adjoining axillary buds, and to the fact that all organs of plants may become abortive in a similar manner; for instance, the sexual organs disappear entirely in the disk-flowers of Viburnum Opulus, and one of the two sexes in the flower of Lychnis dioica. He goes on to answer the question, how it is possible to discover the symmetry in such cases; one method he finds supplied by monstrosities, among which there are even some that may be regarded as a return to the original symmetry, the cases known as peloria. Analogy or 'induction' is, he says, less certain, but of much more extensive application; this is founded exclusively on the knowledge of the relative position of organs. Armed with this, we find that the flower of Albuca, which corresponds to a flower of Liliaceae in everything except in having only three stamens, is to be considered one of the Liliaceae, because it has three filaments placed between the three stamens exactly in the position of the three other stamens in the Liliaceae; it must be concluded therefore that they are abortive stamens. Similar conclusions from analogy must be carried from species to species, from organ to organ, and the great systematists have in fact done so. In certain cases abortion is produced by defect, in others by excess of nourishment, of which he gives examples. An important sentence occurs in this place; everything in nature, he says, leads us to believe that all organisms in their inner nature are regular, and that different forms of abortion differently combined are the cause of all irregularity; from this point of view the smallest irregularities are important, because they lead us to expect greater ones in nearly allied plants; and wherever in a given system of organisation there are inequalities between organs of the same name, the inequality will possibly reach a maximum, that is, end by annihilating the smallest part. Thus in the Labiatae with two stamens, it is the two which in other cases also are the smaller, which are here completely aborted. When in Crassulaceae there are twice as many stamens as petals, those that alternate with the petals are larger and earlier developed, and we may therefore expect that those which are opposite the petals may become abortive; and therefore we may place a genus like Sedum, in which the latter are sometimes wanting, with Crassulaceae; but we could not do so, if we found only the stamens that are superposed upon the petals. It occurs sometimes, he continues, that an organ is prevented from fulfilling its function by partial abortion. In this case it may assume another function, as the abortive leaves of the vetch and the abortive inflorescences of the vine are employed as tendrils. In other cases the abortive organ appears to be quite useless, as for instance many rudimentary leaves. All such useless organs, says De Candolle, exist only in consequence of the primitive symmetry of all organs. Finally the abortion may be so complete that no trace of the organ remains, of which case there are however two kinds, one where the organ is at first perceptible and afterwards quite disappears, as in the abortive loculaments in the fruit of the oak; in other instances no trace is to be seen from the first of the abortive organs, as happens with the fifth stamen of Antirrhinum.
All that has here been said might be alleged word for word in proof of the theory of descent, but our author is an adherent of the dogma of the constancy of species; what from his point of view he really means by abortion is difficult to say, for the object which is aborted is wanting. If species are constant, and therefore of absolutely distinct origin, we must not speak of abortion; we can only say that an organ which is present or large in one species is small or wanting in another. In introducing the idea of abortion De Candolle at once goes beyond the dogma of the constancy of species, without being clear in his own mind with regard to this important step. His proceeding shows that facts lead even a defender of constancy against his will to theories which run counter to that dogma. This is confirmed by his perception of the correlation of growth, which is connected with abortion; he points to the fact that owing to the disappearance of sexual organs in the disk-flowers of Viburnum Opulus the corollas become larger, as do the bracts of the abortive flowers of Salvia Horminum; similarly he regards the disappearance of the seeds in Ananas, Banana, and the Bread-fruit tree as the cause of the enlargement of the pericarps; it does not escape him, that the fertile flowerstalks in Rhus Cotinus remain naked, while an elegant pubescence forms on the barren ones; the leaf-like expansion of the leaf-stalks of Acacia heterophylla, which do not develop their laminae, he refers also to this correlation of growth. He finds the most remarkable example of the kind in the doubling of flowers, where according to his view the disappearance of the anthers is a condition of the corolline expansion of the filaments; in the same way sometimes the carpel is changed into a petal through the disappearance of the stigma. Though in many of these cases it is quite possible to conceive of the relations of cause and effect in the reverse way, yet De Candolle's principle of correlation will be equally applicable.
The second cause by which the symmetry may be obliterated, namely degeneration, asserts itself in the formation of thorns, of threadlike prolongations of membranous expansions, and in the production of fleshy parts, or of parts with dry membranes.
The third kind of departure from the symmetrical plan is the adherence of parts, the theory of which he grounds first and chiefly on the phenomena of grafting, and then passes to more difficult cases. The close packing of the ovaries in some species of honeysuckle, is, he says, the primary cause of their adherence. This therefore does not depend on the plan of symmetry, but upon an accident, which however is constant in its appearance, owing to the specific constitution of such plants. In connection with the phenomena of adherence he next considers the question whether a structure composed of several parts, as for instance a compound ovary, should be considered as originally simple and afterwards divided into parts, or whether the converse is the true account, and he says that we must examine each particular case and decide which is the correct conception. Thus it may be shown that the perfoliate leaves of honeysuckles, as well as the involucres of many Umbelliferae, and monosepalous calyces and monopetalous corollas are due to adherence, and he proceeds to prove that ovaries with several loculaments and several parts have in like manner been formed by adherence of two or more carpellary leaves, and concludes by pointing out the systematic importance of such considerations. Further on he takes occasion to speak of the significance of the relative number of the parts of the flower, on which head he says much that is good, but does not thoroughly investigate the matter; it was not till a later time that Schimper's doctrine of phyllotaxis made it possible to express these relations of number and position more precisely. He concludes his rules for the application of his morphology to the determination of relations of affinity with the declaration, that the whole art of natural classification consists in discerning the plan of symmetry, and in making abstraction of all the deviations from it which he has described, much in the same way as the mineralogist seeks to discover the fundamental forms of crystals from the many derivative forms. It is obvious that all this teaching was a great step in advance upon the right path, that De Candolle has here given utterance for the first time to an important principle of morphology and systematic botany; nevertheless he did not succeed in always consistently carrying out his own principle; he was true to himself only in the determination of small groups of relationship; in framing the largest divisions of the vegetable kingdom he entirely lost sight of the rule which he had himself laid down, that the morphological character of organs and the extent to which it can be turned to account for systematic purposes is entirely independent of their physiological character, and that the most important physiological characters are just those which are of quite subordinate importance in the determination of affinities. In spite of this strange inconsistency, to De Candolle belongs the merit of being the first to point emphatically to the distinction between morphological and physiological marks, and to bring clearly to light the discordance between morphological affinity and physiological habit; but in this discordance lurks a problem, which could only be solved forty years later by Darwin's theory of selection. A genuine inductive process alone could reveal these remarkable relations between the morphological and physiological characters of organs. But it is at the same time true that De Candolle could not have made this discovery, if his predecessors had not already established a large number of affinities. It was while he was engaged in an exact comparison of forms already recognised as undoubtedly related to one another, that that which he called the plan of symmetry, and which was afterwards named a type, revealed itself to him; and as he examined it more closely, and compared it with peculiarities of habit in different plants formed on the same plan, he discovered certain causes, by means of which the deviations were to be explained; these were abortion, degeneration, and adherence. By attending to these he succeeded in discovering affinities that had been hitherto doubtful or unknown; this was at all events the true inductive way of advancing the system, and whatever the earlier systematists had effected that was really valuable had been effected virtually in the same way, only they never arrived at a clear understanding of their own mode of proceeding; they had followed unconsciously the method which De Candolle clearly understood and consciously pursued.
The majority of De Candolle's successors were far from fully appreciating the entire significance of his theory, its importance as a matter of method and principle; on the contrary in the search for affinities they continued to surrender themselves to a blind feeling rather than to a clearly recognised method, and the same must be said unhappily of De Candolle himself, when he was dealing with the establishment of the large divisions of the vegetable kingdom. With equal surprise we find him in the book before us, in which he has developed the true method in systematic botany, expressing the opinion that the most important physiological characters must be employed for the primary divisions of the system, and this idea is not improved by the fact that he ascribes to the organs physiological characters which they do not really possess; thus he regards the vessels as the most important organs of nutrition, which they are not in fact, and upon this double error he builds his primary division of the whole vegetable kingdom into vascular and cellular plants, and then by a third mistake believes that this division coincides with the division of plants into those which have and those which have not cotyledons. The already established division into Monocotyledons and Dicotyledons, which rests upon a leading and purely morphological mark, is spoilt by De Candolle through his following Desfontaines in ascribing to the Dicotyledons a different mode of growth in thickness from that of the Monocotyledons, and characterising the one as exogenous, the other as endogenous. But this notion is utterly incorrect, as von Mohl showed twelve years later; and if it were correct, it would still be unimportant in a systematic point of view, because it appeals to a mark which is morphologically of quite subordinate importance. The worst consequence of these mistakes was, that the Vascular Cryptogams were introduced into the same class with the Monocotyledons, a decided step backwards, if we compare De Candolle's system with that of Jussieu. In spite of these grave defects in the primary divisions of the whole vegetable kingdom De Candolle's system deserved the fame which it acquired and long maintained; it had this advantage over Jussieu's system that in the class of Dicotyledons, the largest division of the whole kingdom, larger sub-divisions appeared, and these served to unite families that were in many points essentially related; the Dicotyledons were in fact divided first of all into two artificial groups according to the presence of two floral envelopes or one; the first and much the larger of these was again broken up into a series of subordinate groups, which pointed in many ways to natural affinities. That these groups, which have only quite recently been materially altered, did to a very considerable extent take account of natural affinities, is due to the fact that De Candolle in framing them really followed his own rules, whereas the superior divisions, which are artificial, owe their existence to his disregard of them.
De Candolle declared emphatically against the old notion, that the vegetable system answers to a linear series,—a notion which sprang from a misunderstanding of the saying, 'Natura non facit saltus,'—and demonstrated its impossibility by examples; but he allowed himself to be too much influenced by the idea which had been thrown out by Linnaeus, and taken up by Giseke, Batsch, Bernardin de St. Pierre, L'Heritier, Du Petit-Thouars and others, that the vegetable kingdom might be compared as respects its grouping to a geographical map, in which the quarters of the globe answer to the classes, the kingdoms to the families, and so on. If the theory of descent is to a certain degree compatible with the idea of a linear sequence from the most imperfect to the highest forms of plants, it is quite incompatible with the above comparison; and systematic investigation, led astray from the right path, is in danger of ascribing the importance of real affinities to mere resemblances of habit, incidental analogies, by which a group of plants appears to be connected with five or six others. In exhibiting his system on paper De Candolle allowed the use of the linear sequence as a convenience, for here it was not, he said, a matter of any importance, since the true task of the science is to study the relations of symmetry in each family and the mutual relations of families to one another; yet in a linear presentation of the system for didactic purposes the sequence ought not to begin with the most simple plants, for these are the least known, but with the most highly developed. Thus De Candolle was the means of removing from the system the last trace of anything which harmonised with an ascending and uninterrupted development of forms. Resting on the doctrine of the constancy of species, and assuming that every group of relationship is founded on a plan of symmetry round which individual forms are grouped as crystals round their parent form, De Candolle was quite consistent in his views. The mode of representation came to prevail in the vegetable kingdom which De Candolle's contemporary, Cuvier, an equally sturdy defender of the dogma of constancy, had introduced in the animal kingdom as the type-theory. Thus the most splendid results obtained by induction were united in the case of De Candolle with the barren dogma of the constancy of species, which, as Lange wittily remarks, comes direct from Noah's ark, to form an intimate mixture of truth and error; nor did De Candolle's many adherents succeed in unravelling the coil, though they removed the chief errors from his system and introduced many improvements.
To these remarks may be appended a table of the main divisions of De Candolle's system of 1819, which so far as it is presented in linear arrangement he calls expressly an artificial system.
I. Vascular plants or plants with cotyledons.
- 1. Exogens or Dicotyledons.
- A. With calyx and corolla:
- Thalamiflorals (polypetalous hypogynous),
- Calyciflorals (polypetalous perigynous),
- Corolliflorals (gamopetalous).
- B. Monochlamydeous plants (with a single floral envelope).
- A. With calyx and corolla:
- 2. Endogens or Monocotyledons.
- A. Phanerogams (true Monocotyledons),
- B. Cryptogams (vascular Cryptogams including Naiadeae).
II. Cellular plants or Acotyledons.
- A. With leaves (Muscineae),
- B. Without leaves (Thallophytes).
The number of families, with Linnaeus 67, with A. L. de Jussieu 100, was increased by De Candolle to 161.
If the principles of comparative morphology laid down by De Candolle were at first prevented from being rapidly disseminated in Germany by the philosophical tendencies then reigning among its botanists, and especially by the obscurities of Goethe's doctrine of metamorphosis, yet these principles and his views also on the natural system won their way by degrees to acknowledgment and acceptance; and after the year 1830 the study of the system was prosecuted by the botanists of Germany, as well as by those of England and France, as the proper object of the science. We may even say that the impulse given by De Candolle worked more powerfully from that time forward in Germany than in France. It may be said too of De Candolle's contemporary, the Englishman Robert Brown (1773-1858), whose chief labours fall in the period between 1820 and 1840, that he, like De Candolle, was better appreciated during that time in Germany than in any other country. Robert Brown, who spent the five years from 1801 to 1805 in Australia, studied the flora of that quarter of the world, and discussed in numerous essays the botanical results of various journeys made by other naturalists in polar regions and in the tropics. In this way he found opportunity to leaven the ideas, which through Humboldt's influence had become predominant respecting the geography of plants, with the spirit of the natural system; he also made the morphology and systematic position of a number of families the subject of critical investigation.
Robert Brown's literary efforts were limited to these monographs; he nowhere attempted to give a connected account of the principles which he follows in them, an exposition of his morphology or a theory of classification, nor did he frame a new system. The results of his studies which were really fruitful and served to advance the science are to be found in the more general remarks, which he managed to insert quite incidentally in his monographs. In this way he succeeded in clearing up the morphology of the flower and with it the systematic position of some difficult families of plants, such as the Grasses, Orchids, Asclepiads, the newly-discovered Rafflesiaceae and others, and to throw new light at the same time on wider portions of the system; in his considerations on the structure and affinities of the most remarkable plants, which had been collected in Africa by different travellers in the years immediately following 1820, he discussed difficult and remarkable morphological relations in the structure of the flower. He referred especially in this essay (1826) to the relations between the numbers of the stamens and carpels, and those of the floral envelopes in the Monocotyledons and Dicotyledons, and showed how these typical, or as he calls them in De Candolle's phraseology, symmetrical relations were changed by abortion, while he entered at the same time into a more exact determination of the position of the aborted and of the perfect organs, in order to discover new relations of affinity. His most valuable work in this direction is a paper on a genus Kingia, discovered in New Holland in 1825; the structure of the seeds in this genus led him to seek more accurate knowledge of the unfertilised ovule in the Phanerogams generally, and especially in the Cycads and Conifers. In spite of the labours of Gärtner and the later researches of Treviranus, there was still considerable obscurity attaching to the theory of the seed, for no one had yet succeeded in referring the position of the embryo in the ripe seed to a general law. For this it was necessary to submit the ovule before fertilisation to careful examination, and Robert Brown carried out this first step to a history of development with great success; he was the first to distinguish the integuments and the nucleus in the ovule, and the embryo-sac in the nucleus, parts which Malpighi and Grew had indeed observed but had not brought out with perfect clearness. The micropyle and the hilum of the seed had not yet been properly distinguished, but had been to some extent even confounded with one another. Robert Brown showed that the hilum answers to the point of attachment of the ovule, while the micropyle is a canal formed by the integuments of the ovule and leading to the summit of the nucleus; that in anatropous ovules the micropyle lies beside the hilum, in orthotropous ovules opposite to it; that the embryo in the embryo-sac (amnion) is always formed at the spot which lies nearest the micropyle, and that the radicle of the embryo is always turned towards the micropyle,—facts which at once established the general rule by which to determine the position of the embryo in the seed and in the fruit. He also gave the first correct explanation of the endosperm as a nourishing substance formed inside the embryo-sac after fertilisation, and more than this, he was the first to distinguish the perisperm as a substance formed outside the embryo-sac in the tissue of the nucleus.
In this way Robert Brown established morphological relations in the organisation of the seed of the Monocotyledons and Dicotyledons, which count among the most important principles of classification in these classes; he was still more happy in being the first to detect the peculiar structure of the flower of Conifers and Cycads, as compared with that of other flowering plants; it was he who perceived that what had been hitherto called a female flower in these plants was really a naked ovule, a view which Trew of Nüremberg had, it is true, suggested in the year 1767. He also called attention to the agreement in structure of the male and female organs in these families. Thus one of the most remarkable facts in vegetation, the gymnospermy of the Conifers and Cycads, was for the first time established, and this led afterwards through Hofmeister's investigations to the important result, that the Gymnosperms, which had been up to that time classed with Dicotyledons, are to be regarded as co-ordinate with Dicotyledons and Monocotyledons, forming a third class through which remarkable homologies were brought to light in the propagation of the higher Cryptogams and the formation of seeds in Phanerogams. No more important discovery was ever made in the domain of comparative morphology and systematic botany. The first steps towards this result, which was clearly brought out by Hofmeister twenty-five years later, were secured by Robert Brown's researches, and he was incidentally led to these researches by some difficulties in the construction of the seed of an Australian genus. He discussed in a similar manner, if not always with such important results, a great variety of questions in morphology and systematic botany; even purely physiological problems were raised by him in this peculiar way, and especially the question how the fertilising matter of the pollen-grains is conveyed to the ovule; he had already concluded from the position of the embryo that it is conveyed through the micropyle and not through the raphe and the hilum, as was then supposed, and he was the first also to follow the passage of the pollen-tubes in the ovary of Orchids up to the ovules; but this is a point which will be more properly considered in the history of the sexual theory.
The peculiar character of the natural system as compared with every artificial arrangement is brought out into higher relief by Robert Brown than by Jussieu and De Candolle, and he succeeded better than any of his predecessors in separating purely morphological and systematically valuable relations of organisation from the physiological adaptations of organs. While the majority of systematists surrendered themselves to the guidance of a blind feeling in the discovery of affinities, their correct determinations being the accidental result of instinct and unconscious operations of the understanding, Brown endeavoured to give an account to himself in every case of the reasons why he took this or that view of the relationships which he determined; from what was already established and indubitable he gathered the value of certain marks, in order to obtain rules for the determination of unknown relationships. In this way he discovered also, that marks, which are of great value for classification within the limits of certain groups of affinity, may possibly prove to be valueless in other divisions. Thus Robert Brown in his numerous monographs supplied the model, by which others might be guided in further applying and completing the method of the natural system; and in this respect he was met by the botanists of Germany in the spirit of the best good-will and most profound appreciation, as is shown by the fact that a collection of his botanical works, translated by different German botanists, was edited in five volumes by Nees von Esenbeck as early as the period between 1825 and 1834. The natural system established itself in Germany through the labours of Brown and De Candolle; and the more correct appreciation of it as compared with the sexual system of Linnaeus was promoted by a work of Carl Fuhlrott which appeared in 1829, in which the systems of Jussieu and De Candolle are compared with those of Agardh, Batsch, and Linnaeus, and the superiority of the natural system is clearly set forth. A still greater effect in this direction was produced by the appearance in 1830 of the 'Ordines naturales plantarum' of Bartling, an independent contribution to this department of botany, and a distinct advance upon what had hitherto been effected. The contemporary monographs of Roeper on the Euphorbiaceae and Balsamineae and his treatise 'De organis plantarum' (1828), are an able, independent, and logical application of the principles of the morphology of the flower laid down by De Candolle and Brown to the elucidation of morphological and systematic conceptions. But the new methods of investigation introduced by De Candolle and Robert Brown had to encounter in Germany, and to some extent in France also, not only the antiquated views of Linnaeus, but, what was still worse, the erroneous notions of the nature-philosophy founded by Schelling. The misty tenets of this philosophy could scarcely find a more fruitful soil than the natural system with its mysterious affinities, and Goethe's doctrine of metamorphosis contributed not a little to increase the confusion. These historical phenomena will be further considered in the following chapter; at present we are more concerned to show how the professed systematists pursued the path opened by De Candolle and Brown. And here it must be noticed that from about the year 1830, in Germany especially, morphological enquiry became separated as a special subject from systematic botany; it became more and more the fashion to treat the latter as independent of morphology, and thus to forsake the source of deeper insight which comparative and genetic morphology alone can open to the systematist; morphology on the other hand took a new flight, and as it thus developed itself apart from pure systematic botany, its progress must be described by itself in a later portion of this history.
If advance in systematic botany depended on the number of systems that were proposed from 1825 to 1845, that period must be looked upon as its golden age; no less than twenty-four systems made their appearance during these twenty years, without counting those which were inspired by the views of the nature-philosophy. There was great and spreading growth, but no corresponding depth; no really new points of view were opened for classification, and as regards the true principles of the natural system there were symptoms of evident decline rather than of advance, as will be shown below. Improvements were effected certainly in the details of the system, since botanists generally adhered to the principles laid down by De Candolle, Jussieu, and Brown. Families were cleared up and better defined, and groups of families were proposed which assumed more and more the appearance of natural cycles of relationship. The class more especially treated was the extensive one of the Dicotyledons, in which the families, continually growing more and more numerous, were in Jussieu's arrangement a chaos, but had been united into larger groups in a somewhat artificial manner by De Candolle. Here we see once more how the formation of the system rises step by step from the particular to the more general; at an earlier period genera were constructed out of species, and families out of genera, and during the years from 1820 to 1845 the families were united into more comprehensive groups; but these orders or classes were not yet grouped together in such a manner as to ensure the separation of the largest divisions of the vegetable kingdom in a natural manner. The great class of Dicotyledons is not even yet so arranged that the smaller aggregates of families connect satisfactorily one with another. Nevertheless a considerable advance was made by the establishment of a large number of smaller groups of families, and Bartling and Endlicher were especially successful in founding such groups and supplying them with names and characters.
If on the other hand we turn to the primary divisions of the vegetable kingdom, we find that certain large and natural groups came to be most generally recognised and placed in the front rank in every scheme; such were the groups of the Thallophytes, Muscineae, Vascular Cryptogams, Gymnosperms, Dicotyledons and Monocotyledons. But the co-ordination of these great divisions of the whole vegetable kingdom was far from being rightly understood. It was usage rather than anything else, which gradually put them forward as primary types; in the systems themselves some received too great, others too little prominence, or other groups of doubtful character were admitted alongside of them. Bartling, for instance, whose system up to 1850 or even longer may rank as one of the most natural, adheres to De Candolle's division of the vegetable kingdom into cellular and vascular plants, and rightly divides the former into two main groups, Thallophytes and Muscineae (Homonemeae and Heteronemeae), while he separates the latter into Vascular Cryptogams and Phanerogams; but the Phanerogams are divided into Monocotyledons and Dicotyledons, which again are distributed into four groups, one of these being characterised by the presence of a vitellus, that is, of an endosperm surrounded by a perisperm,—a thoroughly artificial division. The three other divisions are named apetalous, monopetalous, and polypetalous, but the Coniferae and Cycadeae are placed in the apetalous division. Less satisfactory is the primary division into Thallophytes and Cormophytes proposed by Endlicher, the latter separating into the divisions Acrobrya (Muscineae, Vascular Cryptogams, and Cycads), Amphibrya (Monocotyledons), and Acramphibrya (Dicotyledons and Conifers); the names of the three latter groups, the first of which is utterly unnatural, are founded on erroneous assumptions respecting growth in length and thickness, which Endlicher borrowed from Unger. While Endlicher's great work has continued down to our own time to be indispensable to the botanist as a book of reference on account of the fulness of its descriptions of families and genera, the system projected by Brongniart in 1843 has acquired a sort of official authority in France. The whole vegetable kingdom is here distributed into two divisions, Cryptogams and Phanerogams, and the former are incorrectly characterised as asexual, the latter as having distinction of sex. The Phanerogams, divided into Monocotyledons and Dicotyledons, are distributed into groups in a manner that is not satisfactory; but the system has one merit, that it keeps the Gymnosperms together in one body; and if they are incorrectly classed with the Dicotyledons, it was still a sign of progress, that Robert Brown's discovery of gymnospermy was to some extent practically recognised. The system devised by John Lindley attained to about the same importance in England as attached to those of Bartling and Endlicher in Germany, and that of Brongniart in France. After various earlier attempts he proposed a system in 1845, in which, as in Brongniart's arrangement, the Cryptogams are characterised as asexual or flowerless plants, the Phanerogams as sexual or flowering plants; the former are divided into Thallogens and Acrogens, the Phanerogams into five classes; (i) Rhizogens (Rafflesiaceae, Cytineae, Balanophorae); (2) Endogens (parallel-nerved Monocotyledons); (3) Dictyogens (net-veined Monocotyledons); (4) Gymnogens (Gymnosperms); (5) Exogens (Dicotyledons). This classification is one of the most unfortunate that were ever attempted; the systematic value of the Rhizogens is much overrated on account of their striking habit; the Monocotyledons are separated into two classes on the strength of an unimportant mark. The characters assigned to all these groups are on the whole thoroughly faulty. These systems have been selected for notice from among many others, because they attained an extended notoriety and importance from the circumstance that their authors, Brongniart excepted, made them the occasions of comprehensive descriptions of the whole vegetable kingdom, and again because it would be superfluous for our present purpose to bestow a closer consideration on the systems of less eminent men. Whoever desires further information on the matter will find it in the introduction to Lindley's 'Vegetable Kingdom' of 1853.
If we consider the principles and points of view adopted in these systems, one thing especially strikes us, that, except in the case of Bartling, physiologico-anatomical marks were employed along with morphological ones to characterise the primary divisions; their authors fell into the mistake committed by De Candolle, and unfortunately these very marks rested in part or wholly on misapprehensions, as in Endlicher's division into Acrobrya, etc., and Lindley's classes of Rhizogens and Dictyogens. It was still more unfortunate that individual systematists obstinately refused to accept well authenticated facts, which it is true had not been discovered by systematists, but were nevertheless of the highest value for the system. It is scarcely credible that Lindley in 1845, and again in 1853, maintained the distinction between endogenous and exogenous growth in stems, though Hugo von Mohl had in 1831 produced decisive proof that this distinction laid down by Desfontaines and adopted by De Candolle had no real existence. The same was the case with the characters of the Cryptogams, in which the mark of having no sexual organs was repeatedly adopted as running through the whole class, although various instances of sexuality in Cryptogams were known before 1845; Schmidel had described the sexual organs of the Liverworts about the middle of the previous century, Hedwig those of the Mosses in 1782, and Vaucher in 1803 had suggested that the conjugation of Spirogyra among the Algae should be regarded as a sexual act; the systematists in fact did not know what to make of these intimations.
It was again a misfortune that the systematists in their labours often neglected to distinguish between the search for marks and the use to be made of them; the examination of all possible marks should lead to the establishing the systematic importance of certain fixed marks or their value for classification. When research has done its work, then it is sufficient in exhibiting the system to put forward only the prominent marks; and frequently a single one suffices to unite a natural group. Such a leading mark is like the standard of a regiment; its significance is not great in itself, but it serves the great practical purpose of indicating a whole group of marks which are connected with it. It was a still greater misfortune that scarcely any systematist after De Candolle endeavoured to form a clear conception in his own mind of the principles on which the natural system must be elaborated, and to set them forth in a connected form as the theory of the system. The student had to accept the arrangement offered him as a fact simply without understanding it, and the systematists themselves usually followed only a blind feeling in the framing of their groups, and never unfolded the grounds of their proceeding with logical distinctness. In this respect John Lindley forms an honourable exception, inasmuch as he did, on several occasions after 1830, give full expositions of his views on the principles of natural classification, and like De Candolle endeavoured to develop a theory of the system. But he deserves credit only for the endeavour, for the principles themselves which he laid down are not only to a great extent incorrect, but they are opposed to his own and to every other natural system. We find this opposition between theory and practice much more strongly marked in Lindley than in De Candolle; the cases only are so far different, that De Candolle laid down correct principles for the determination of affinities, but in some cases did not follow them, whereas Lindley deduced quite incorrect rules of system from existing and long-established natural affinities. The consideration of all the systems framed up to the year 1853 shows clearly that the characters of truly natural groups are to be found only in morphological marks; yet Lindley enunciates the principle that a mark, or, as he incorrectly says, an organ, is more important for classification in proportion as it possesses a higher physiological value for the preservation and propagation of the individual. If this were true, nothing would be easier than to frame a natural system of plants; it would suffice to divide plants first of all into those without and those with chlorophyll, for the presence of chlorophyll is more essential than that of any other substance to the nourishment of plants, and its physiological importance is therefore preeminent; in that case of course such Orchideae as have no chlorophyll, the Orobancheae, Cuscuta, Rafflesia, etc., would form one class with the Fungi, and all other plants the other. It is very important for the existence of a plant whether its organisation is adapted to its growing in water, or on dry land, or underground, and if we took Lindley at his word, he would be obliged to bring the Algae, Rhizocarps, Vallisnerias, water Ranunculuses, Lemna, etc., into one group. It is very important for the existence of a plant whether it grows upright of itself, or climbs upwards by the aid of tendrils or of a twining stem or otherwise, and accordingly we might on Lindley's principle collect certain ferns, the vine, the passion-flower, many of the pea kind, etc., into one order. It is obvious that Lindley's main axiom of systematic botany appears in this way utterly unreasonable; yet by this principle he judges of the systematic value of anatomical characters, those of the embryo and endosperm, of the corolla and the stamens, everywhere laying stress on their physiological importance, which in these parts has really little systematic value. This mode of proceeding on the part of Lindley, compared with his own system, which with all its grave faults is still always a morphologically natural system, proves that like many other systematists, he did not literally and habitually follow the rules he himself laid down, for if he had, something very different from a natural system must have been the result. The success which was really obtained in the determination of affinities was due chiefly to a correctness of feeling, formed and continually being perfected by constant consideration of the forms of plants. It was still therefore virtually the same association of ideas as in de l'Obel and Bauhin, operating to a great extent unconsciously, by which natural affinities were by degrees brought to light; and men like Lindley, of pre-eminent importance as systematists, were, as the above examples show, never clear about the very rules by which they worked. And yet in this way the natural system was greatly advanced in the space of fifty years. The number of affinities actually recognised increased with wonderful rapidity, as appears from a comparison of the systems of Bartling, Endlicher, Brongniart, and Lindley, with those of De Candolle and Jussieu. Nothing shows the value of the systems thus produced before 1850 as classifications of the vegetable kingdom more forcibly than the fact that a clear and methodical thinker like Darwin was able to draw from them the chief supports of the theory of descent. For it is quite certain that Darwin has not framed his theory in opposition to morphology and system, and drawn it from any hitherto unknown principles; on the contrary, he has deduced his most important and most incontestable propositions directly from the facts of morphology and of the natural system, as it had been developed up to his time. He is always pointing expressly to the fact that the natural system in the form in which it has come to him, which he accepts in the main as the true one, is not built upon the physiological, but upon the morphological value of organs; it may, he says, be laid down as a rule, that the less any portion of the organisation is bound up with special habits of life, the more important it is for classification. Like Robert Brown and De Candolle, he insists upon the high importance for purposes of classification of aborted and physiologically useless organs; he points to cases in which very distant affinities are brought to light by numerous transition-forms or intermediate stages, of which the class of the Crustaceae offers a specially striking example in the animal kingdom, while certain series of forms of Thallophytes, the Muscineae, the Aroideae and others, may be adduced as instances of the same kind in the vegetable world; in such cases the most distant members of a series of affinities have sometimes no one common mark, which they do not share with all other plants of a much larger division. From these and other similar statements of Darwin we see plainly, that he actually did gather from existing natural systems of plants and animals the rules by which systematists had worked, but which they themselves observed only more or less unconsciously, and never with a full and clear recognition of them. He says quite rightly, when the investigators of nature are practically engaged with their task, they do not trouble themselves about the physiological value of the characters which they employ for the limiting a group or the establishment of a single species. Darwin clearly perceived and consistently kept in view the discordance between the systematic affinity of organisms and their adaptation to the conditions of life, which De Candolle had already but imperfectly recognised. The clear perception of this discordance was in fact the one thing needed to mark the true character of the natural system, and to make the theory of descent appear as the only possible explanation of it. The fact which morphologists and systematists had painfully brought to light, but had not sufficiently recognised in its full importance, that two entirely different principles are united in the nature of every individual organism, that on the one hand the number, the arrangement, and the history of the development of the organs of a species point to corresponding relations in many other species, while on the other hand the manner of life and the consequent adaptation of the same organs may be quite different in these allied species. This fact admits of no explanation but the one given by the theory of descent; it is therefore the historical cause and the strongest logical support of that theory, and the theory itself is directly deduced from the results which the efforts of the systematists have established. That the majority of systematists did at first distinctly declare against the theory of descent can surprise no one who observes that they were so little able to give an account of their own mode of procedure, as appears in so striking a manner from Lindley's theoretical speculations. One consequence of this want of clearness in combination with the dogma of the constancy of species has been already mentioned in the introduction; namely, the notion professedly adopted by Lindley, Elias Fries, and others, that an idea lies at the foundation of every group of affinities, that the natural system is a representation of the plan of creation. But the question, how such a plan of creation could explain the strange fact that the physiological adaptations of organs to the conditions of life have nothing at all to do with their systematic connection, was quietly disregarded; and in fact the notion, founded on Platonic and Aristotelian philosophy, of a plan of creation and of ideal forms underlying systematic groups, could not explain this discordance between morphological and physiological characters. It would be easy to maintain the view of the systematists, that the natural system represents a plan of creation, if physiological and morphological characters went always truly hand in hand, if the adaptation of the organs to the conditions of life in the species were perfect; but facts show that the adaptation is in the best of cases comparatively imperfect, and that it is in all cases brought about by the accommodation to new requirements of organs which originally served to other functions.
- Bernard de Jussieu, born at Lyons in 1699, and at first a practicing physician there, was by Vaillant's intervention called to Paris, and alter Vaillant's death became Professor and Demonstrator at the Royal Garden. He and Peissonel were among the first who declared against the vegetable nature of the Corals. It is expressly stated in his Éloge ('Histoire de l'Académie Royale des Sciences,' Paris, 1777) that he founded his natural families on the Linnaean fragment. He died in 1777.
- A. L. de Jussieu, born at Lyons, came to Paris to his uncle Bernard in 1765. In 1790 he was a member of the Municipality, and till 1792 Superintendent of Hospitals. When the Annales du Museum were founded in 1802, he resumed his botanical pursuits. In 1826 his son Adrien took his place at the Museum. See his life by Brougniart in the 'Annales des Sciences Naturelles,' vii (1837).
- Joseph Gärtner was born at Calw in Würtenberg in 1732, and died in 1791. He commenced his studies in Göttingen in 1751, where he was a pupil of Haller. He travelled into Italy, France, Holland, and England in order to make the acquaintance of famous naturalists, and worked also at physics and zoology. In 1760 he was Professor of Anatomy in Tübingen, and in 1768 became Professor of Botany at St. Petersburg; but finding himself unable to bear the climate, he returned to Calw in 1770, and gave himself up entirely to his book, 'De fructibus et seminibus plantarum,' which he had already commenced. Banks and Thunberg, one of whom had returned from a voyage round the world, the other from Japan, handed over to him the collections of fruits which they had made. His persistent study, partly with the microscope, brought him near to blindness. There is an interesting life of Gärtner by Chaumeton in the 'Biographie Universelle.'
- Augustin Pyrame de Candolle sprang from a Provençal family, which had fled from religious persecution to Geneva, where it was and is still held in great estimation. He associated as a boy with Vaucher, and on his first visit to Paris in 1796 with Desfontaines and Dolomieu, and after his return to Geneva was a friend of Senebier. The elder Saussure, and afterwards Biot, whom he assisted in an investigation in physics, endeavoured to attach him to that study. He spent the years from 1798 to 1808 in Paris, where he lived in close intercourse with the naturalists of that city. Numerous smaller monographs, and the publication of his work on succulent plants and of a new edition of De Lamarck's 'Flore Française,' occupied this earlier period of his life. From 1808 to 1816 he was Professor of Botany at Montpellier. During this time he made many botanical journeys in all parts of France and the neighbouring countries, and wrote many monographs, his essays on the geography of plants, and his most important work, the 'Théorie élémentaire.' From 1816 till his death in 1841 he resided once more in Geneva, which had freed itself in 1813 from the enforced connection with France established in 1798. Here De Candolle found time to take part in political and social questions, in addition to an almost incredible amount of botanical labour. (Notice sur la vie et les ouvrages de A. P. De Candolle par De la Rive, Geneve, 1845.)
- Robert Brown was the son of a Protestant minister of Montrose, and studied medicine first at Aberdeen and afterwards in Edinburgh; he then became a surgeon in the army, and was at first stationed in the north of Ireland. When the Admiralty despatched a scientific expedition to Australia under Captain Flinders in 1801, he was appointed naturalist to the expedition on the recommendation of Sir Joseph Banks, F. Bauer being associated with him as botanical draughtsman, Good as gardener, Westall as landscape-painter; one of the midshipmen of the vessel was John Franklin. In consequence of the unseaworthiness of the ship Flinders left Australia, intending to return with a better one, but was shipwrecked on the voyage and detained by the French at Port Louis as a prisoner of war till 1810. The naturalists of the expedition remained in Australia till 1805, when Brown returned to England with 4000 for the most part new species of plants. Sir J. Banks appointed him his librarian and keeper of his collections in 1810; he was also Librarian to the Linnaean Society of London. In 1823 he received the bequest of Banks' library and collections, which were to be transferred after his death to the British Museum; but by his own wish they were deposited there at once, and he himself received the appointment of Custodian of the Museum and remained in that position till his death. At Humboldt's suggestion Sir Robert Peel's Ministry granted him a yearly pension of £200. His merits were universally acknowledged, and Humboldt even named him 'botanicorum facile princeps.'
- Stephen Ladislaus Endlicher was born at Pressburg in 1805, and abandoning the study of theology became Scriptor in the Imperial Library at Vienna in 1828, and in 1836 Custos of the botanical department of the Imperial Collection of Natural History. Having graduated at the University in 1840, he became Professor of Botany and Director of the Botanic Garden. His library and herbarium, valued at 24,000 thalers, he presented to the State, and with his private means founded the Annalen des Wiener-Museums, purchased botanical collections and expensive botanical books, and published his own works and works of other writers. His official salary was small, and having exhausted his resources in these various expenses, he put an end to his own life in March 1849. Endlicher was not only one of the most eminent systematists of his day, but a philologist also, and a good linguist. He wrote among other things a Chinese grammar. See 'Linnaea,' vol. xxxiii (1864 and 1865), p. 583.
- John Lindley, Professor of Botany in the University of London, was born at Chatton near Norwich in 1799, and died in London in 1865.
- Auguste de Saint Hilaire was born at Orleans in 1779, and died there in 1853; he was Professor at Paris, and in 1840 published his 'Leçons de Botanique comprenant principalement la Morphologie Vegétale,' etc. This work contains a somewhat diffuse account of P. de Candolle's doctrine of symmetry, together with Goethe's theory of metamorphosis and Schimper's doctrine of phyllotaxis, and his own views also on classification founded on the comparative morphology of the day. It is marked by fewer errors than will be found in Lindley's theoretical writings, but it is less profound, and touches only incidentally on fundamental questions; at the same time it possesses historical interest as giving a lucid description of the state of morphology before 1840.