Transcriber’s Notes
Obvious typographical errors have been silently corrected. Variations in hyphenation and accents have been standardised but all other spelling and punctuation remains unchanged.
The Errata have been implemented.
The cover was prepared by the transcriber and is placed in the public domain.
HISTORY OF BOTANY
SACHS
London
HENRY FROWDE
Oxford University Press Warehouse
Amen Corner, E.C.
HISTORY OF BOTANY
(1530-1860)
BY
JULIUS VON SACHS
PROFESSOR OF BOTANY IN THE UNIVERSITY OF WÜRZBURG
AUTHORISED TRANSLATION
BY
HENRY E. F. GARNSEY, M.A.
Fellow of Magdalen College, Oxford
REVISED BY
ISAAC BAYLEY BALFOUR, M.A., M.D., F.R.S.
Professor of Botany in the University
And Keeper of the Royal Botanic Garden, Edinburgh
Oxford
AT THE CLARENDON PRESS
1890
[All rights reserved]
Oxford
PRINTED AT THE CLARENDON PRESS
BY HORACE HART, PRINTER TO THE UNIVERSITY
[PREFACE.]
Botanical Science is made up of three distinct branches of knowledge, Classification founded on Morphology, Phytotomy, and Vegetable Physiology. All these strive towards a common end, a perfect understanding of the vegetable kingdom, but they differ entirely from one another in their methods of research, and therefore presuppose essentially different intellectual endowments. That this is the case is abundantly shown by the history of the science, from which we learn that up to quite recent times morphology and classification have developed in almost entire independence of the other two branches. Phytotomy has indeed always maintained a certain connection with physiology, but where principles peculiar to each of them, fundamental questions, had to be dealt with, there they also went their way in almost entire independence of one another. It is only in the present day that a deeper conception of the problems of vegetable life has led to a closer union between the three. I have sought to do justice to this historical fact by treating the parts of my subject separately; but in this case, if the present work was to be kept within suitable limits, it became necessary to devote a strictly limited space only to each of the three historical delineations. It is obvious that the weightiest and most important matter only could find a place in so narrow a frame, but this I do not exactly regard as a misfortune, and in the interests of the reader it is rather an advantage; for, in accordance with the objects of the ‘General History of the Sciences,’ this History of Botany is not intended for professional persons only, but for a wider circle of readers, and to these perhaps even the details presented in it may here and there seem wearisome.
The style of the narrative might have been freer, and greater space might have been allotted to reflections on the inner connection of the whole subject, if I had had before me better preliminary studies in the history of botany; but as things are, I have found myself especially occupied in ascertaining questions of historical fact, in distinguishing true merit from undeserved reputation, in searching out the first beginnings of fruitful thoughts and observing their development, and in more than one case in producing lengthy refutations of wide-spread errors. These things could not be done within the allotted space without a certain dryness of style and manner, and I have often been obliged to content myself with passing allusions where detailed explanation might have been desired.
As regards the choice of topics, I have given prominence to discoveries of facts only when they could be shown to have promoted the development of the science; on the other hand, I have made it my chief object to discover the first dawning of scientific ideas and to follow them as they developed into comprehensive theories, for in this lies, to my mind, the true history of a science. But the task of the historian of Botany, as thus conceived, is a very difficult one, for it is only with great labour that he succeeds in picking the real thread of scientific thought out of an incredible chaos of empirical material. It has always been the chief hindrance to a more rapid advance in botany, that the majority of writers simply collected facts, or if they attempted to apply them to theoretical purposes, did so very imperfectly. I have therefore singled out those men as the true heroes of our story who not only established new facts, but gave birth to fruitful thoughts and made a speculative use of empirical material. From this point of view I have taken ideas only incidentally thrown out for nothing more than they were originally; for scientific merit belongs only to the man who clearly recognises the theoretical importance of an idea, and endeavours to make use of it for the promotion of his science. For this reason I ascribe little value, for instance, to certain utterances of earlier writers, whom it is the fashion at present to put forward as the first founders of the theory of descent; for it is an indubitable fact that the theory of descent had no scientific value before the appearance of Darwin’s book in 1859, and that it was Darwin who gave it that value. Here, as in other cases, it appears to me only true and just to abstain from assigning to earlier writers merits to which probably, if they were alive, they would themselves lay no claim.
J. SACHS.
Würzburg, July 22, 1875.
[THE AUTHOR’S PREFACE]
To the English translation of the History of Botany of Julius von Sachs.
I am gratefully sensible of the honourable distinction implied in the determination of the Delegates of the Clarendon Press to have my History of Botany translated into the world-wide language of the British Empire. Fourteen years have elapsed since the first appearance of the work in Germany, from fifteen to eighteen years since it was composed,—a period of time usually long enough in our age of rapid progress for a scientific work to become obsolete. But if the preparation of an English translation shows that competent judges do not regard the book as obsolete, I should be inclined to refer this to two causes. First of all, no other work of a similar kind has appeared, as far as I know, since 1875, so that mine may still be considered to be, in spite of its age, the latest history of Botany; secondly, it has been my endeavour to ascertain the historical facts by careful and critical study of the older botanical literature in the original works, at the cost indeed of some years of working-power and of considerable detriment to my health, and facts never lose their value,—a truth which England especially has always recognised.
But the present work is not a simple enumeration of the names of botanists and of their writings, no mere list of the dates of botanical discoveries and theories; such was not at all my plan when I designed it. On the contrary I purposed to present to the reader a picture of the way in which the first beginnings of scientific study of the vegetable world in the sixteenth century made their appearance in alliance with the culture prevailing at the time, and how gradually by the intellectual efforts of gifted men, who at first did not even bear the name of botanists, an ever deepening insight was obtained into the relationship of all plants one to another, into their outer form and inner organisation, and into the vital phenomena or physiological processes dependent on these conditions.
For the attainment of this end it was above all things necessary for me to form a clear judgment respecting the influence of the views and principles enunciated by the different authors on the further development of botanical science. This is to the historian of science the central point round which all beside should be disposed, and without which the entire work breaks up into a collection of unmeaning details, and it is one which demands knowledge of the subject, and capacity and impartiality of judgment. On questions connected with times long gone by the decision of the experts has in most cases been already given, though I myself found to my surprise that older authors had for centuries been regarded as the founders of views which they had distinctly repudiated as absurd, showing how necessary it is that the works of our predecessors should from time to time be carefully read and compared together. But in the majority of cases there is no dispute at the present day respecting the historical value, that is the operative influence on posterity, of works written three hundred or even one hundred years ago.
But it is a very different matter when the author of a book like mine ventures, as I have done for sufficient reasons but at the same time with regret, to sit in judgment on the works of men of research and experts, who belong to our own time and who exert a lively influence on their generation. In this case the author can no longer appeal to the consentient opinion of his contemporaries; he finds them divided into parties, and involuntarily belongs to a party himself. But it is a still more weighty consideration that he may subsequently change his own point of view, and may arrive at a more profound insight into the value of the works which he has criticised; continued study and maturer years may teach him that he overestimated some things fifteen or twenty years ago and perhaps undervalued others, and facts, once assumed to be well established, may now be acknowledged to be incorrect.
Thus it has happened in my own case also in some but not in many instances, in which I have had to express an opinion respecting the character of works which appeared after 1860, and which to some extent influenced my judgment on the years immediately preceding them. But this was from fifteen to eighteen years ago when I was working at my History. It might perhaps be expected that I should remove all such expressions of opinion from the work before it is translated. In some few cases, in which this could be effected by simply drawing the pen through a few lines, I have so done; but it appeared to me that to alter with anxious care every sentence which I should put into a different form at the present day would serve no good purpose, for I came to the conclusion that my book itself may be regarded as a historical fact, and that the kindly and indulgent reader may even be glad to know what one, who has lived wholly in the science and taken an interest in everything in it old and new, thought from fifteen to eighteen years ago of the then reigning theories, representing as he did the view of the majority of his fellow-botanists.
However, these remarks relate only to two famous writers on the subjects with which this History is concerned. If the work had been brought to a close with the year 1850 instead of 1860, I should hardly have found it necessary to give them so prominent a position in it. Their names are Charles Darwin and Karl Nägeli. I would desire that whoever reads what I have written on Charles Darwin in the present work should consider that it contains a large infusion of youthful enthusiasm still remaining from the year 1859, when the ‘Origin of Species’ delivered us from the unlucky dogma of constancy. Darwin’s later writings have not inspired me with the like feeling. So it has been with regard to Nägeli. He, like Hugo von Mohl, was one of the first among German botanists who introduced into the study that strict method of thought which had long prevailed in physics, chemistry, and astronomy; but the researches of the last ten or twelve years have unfortunately shown that Nägeli’s method has been applied to facts which, as facts, were inaccurately observed. Darwin collected innumerable facts from the literature in support of an idea, Nägeli applied his strict logic to observations which were in part untrustworthy. The services which each of these men rendered to the science are still acknowledged; but my estimate of their importance for its advance would differ materially at the present moment from that contained in my History of Botany. At the same time I rejoice in being able to say that I may sometimes have overrated the merits of distinguished men, but have never knowingly underestimated them.
Dr. J. von SACHS,
Foreign Fellow of the Royal Society.
Würzburg, March 24, 1889.
NOTE BY THE TRANSLATOR.
No History of Botany in English has ever been published, and it is to supply in some measure this want, long felt by English-speaking students, that this translation of Professor Sachs’ masterly sketch has been prepared.
H. E. F. G.
[CONTENTS.]
| [FIRST BOOK]. | |
| History of Morphology and Classification. | |
| 1530-1860. | |
| PAGE | |
| Introduction | [3] |
| [CHAPTER I]. | |
| The Botanists of Germany and the Netherlands from Brunfels to Caspar Bauhin, 1530-1623 | [13] |
| [CHAPTER II]. | |
| Artificial Systems and Terminology of Organs from Cesalpino to Linnaeus, 1583-1760 | [37] |
| [CHAPTER III]. | |
| Development of the Natural System under the Influence of the Dogma of the Constancy of Species, 1759-1850 | [108] |
| [CHAPTER IV]. | |
| Morphology under the Influence of the Doctrine of Metamorphosis and of the Spiral Theory, 1790-1850 | [155] |
| [CHAPTER V]. | |
| Morphology and Systematic Botany under the Influence of the History of Development and the knowledge of the Cryptogams, 1840-1860 | 182 |
| [SECOND BOOK]. | |
| History of Vegetable Anatomy. | |
| 1671-1860. | |
| Introduction | [219] |
| [CHAPTER I]. | |
| Phytotomy founded by Malpighi and Grew, 1671-1682 | [229] |
| [CHAPTER II]. | |
| Phytotomy in the Eighteenth Century | [246] |
| [CHAPTER III]. | |
| Examination of the Matured Framework of Cell-Membrane in Plants, 1800-1840 | [256] |
| [CHAPTER IV]. | |
| History of Development of the Cell, Formation of Tissues, Molecular Structure of Organised Forms, 1840-1860 | [311] |
| [THIRD BOOK]. | |
| History of Vegetable Physiology. | |
| 1583-1860. | |
| Introduction | [359] |
| [CHAPTER I]. | |
| History of the Sexual Theory | |
| 1. From Aristotle to R. J. Camerarius | [376] |
| 2. Establishment of the Doctrine of Sexuality in Plants by R. J.Camerarius, 1691-1694 | [385] |
| 3. Dissemination of the New Doctrine; its Adherents and Opponents, 1700-1760 | [390] |
| 4. The Theory of Evolution and Epigenesis | [402] |
| 5. Further Development of the Sexual Theory by J. G. Koelreuter and Konrad Sprengel, 1761-1793 | [406] |
| 6. New opponents of Sexuality and their refutation by Experiments, 1785-1849 | [422] |
| 7. Microscopic Investigation into the Processes of Fertilisation in the Phanerogams, the Pollen-Tube and Eggs, 1830-1850 | [431] |
| 8. Discovery of Sexuality in the Cryptogams, 1837-1860 | [436] |
| [CHAPTER II]. | |
| History of the Theory of Nutrition of Plants, 1583-1860 | [445] |
| 1. Cesalpino, 1583 | [450] |
| 2. First Inductive Experiments and Opening of New Points of View in the History of the Theory of the Nutrition of Plants, to 1730 | [453] |
| 3. Fruitless Attempts to Explain the Movement of the Sap in Plants, 1730-1780 | [482] |
| 4. The Modern Theory of Nutrition Founded by Ingen-Houss and Theodore de Saussure, 1779-1804 | [491] |
| 5. Vital Force. Respiration and Heat of Plants. Endosmose, 1804-1840 | [504] |
| 6. Settlement of the Question of Food-Material of Plants, 1840-1860 | [524] |
| [CHAPTER III]. | |
| History of Phytodynamics | |
| From end of 17th century to about 1860 | [535] |
| [Index] | [565] |
[ERRATA.]
- Page 18, line 3 from bottom, for Chini read Ghini
- Page 20, line 7, for Schmiedel read Schmidel
- Page 160, line 2 from bottom, for many read some
- Page 160, note, for Robert read Louis Marie Aubert
- Page 201, line 11, for asexually read sexually.
[FIRST BOOK]
HISTORY OF MORPHOLOGY AND CLASSIFICATION
(1530-1860)
[INTRODUCTION.]
The authors of the oldest herbals of the 16th century, Brunfels, Fuchs, Bock, Mattioli and others, regarded plants mainly as the vehicles of medicinal virtues; to them plants were the ingredients in compound medicines, and were therefore by preference termed ‘simplicia,’ simple constituents of medicaments. Their chief object was to discover the plants employed by the physicians of antiquity, the knowledge of which had been lost in later times. The corrupt texts of Theophrastus, Dioscorides, Pliny and Galen had been in many respects improved and illustrated by the critical labours of the Italian commentators of the 15th and of the early part of the 16th century; but there was one imperfection which no criticism could remove,—the highly unsatisfactory descriptions of the old authors or the entire absence of descriptions. It was moreover at first assumed that the plants described by the Greek physicians must grow wild in Germany also, and generally in the rest of Europe; each author identified a different native plant with some one mentioned by Dioscorides or Theophrastus or others, and thus there arose as early as the 16th century a confusion of nomenclature which it was scarcely possible to clear away. As compared with the efforts of the philological commentators, who knew little of plants from their own observation, a great advance was made by the first German composers of herbals, who went straight to nature, described the wild plants growing around them and had figures of them carefully executed in wood. Thus was made the first beginning of a really scientific examination of plants, though the aims pursued were not yet truly scientific, for no questions were proposed as to the nature of plants, their organisation or mutual relations; the only point of interest was the knowledge of individual forms and of their medicinal virtues.
The descriptions were at first extremely inartistic and unmethodical; but the effort to make them as exact and clear as was possible led from time to time to perceptions of truth, that came unsought and lay far removed from the object originally in view. It was remarked that many of the plants which Dioscorides had described in his Materia Medica do not grow wild in Germany, France, Spain, and England, and that conversely very many plants grow in these countries, which were evidently unknown to the ancient writers; it became apparent at the same time that many plants have points of resemblance to one another, which have nothing to do with their medicinal powers or with their importance to agriculture and the arts. In the effort to promote the knowledge of plants for practical purposes by careful description of individual forms, the impression forced itself on the mind of the observer, that there are various natural groups of plants which have a distinct resemblance to one another in form and in other characteristics. It was seen that there were other natural alliances in the vegetable world, beside the three great divisions of trees, shrubs, and herbs adopted by Aristotle and Theophrastus. The first perception of natural groups is to be found in Bock, and later herbals show that the natural connection between such plants as occur together in the groups of Fungi, Mosses, Ferns, Coniferae, Umbelliferae, Compositae, Labiatae, Papilionaceae was distinctly felt, though it was by no means clearly understood how this connection was actually expressed; the fact of natural affinity presented itself unsought as an incidental and indefinite impression, to which no great value was at first attached. The recognition of these groups required no antecedent philosophic reflection or conscious attempt to classify the objects in the vegetable world; they present themselves to the unprejudiced eye as naturally as do the groups of mammals, birds, reptiles, fishes and worms in the animal kingdom. The real resemblance of the organisms in such groups is unconsciously accepted by the mind through the association of ideas, and it is not till this involuntary mental act, which in itself requires no effort of the understanding, is accomplished, that any necessity is felt for obtaining a clearer idea of the phenomenon, and the sense of this necessity is the first step to intentional systematic enquiry. The series of botanical works published in Germany and the Netherlands from 1530 to 1623, from Brunfels to Kaspar Bauhin, shows very plainly how this perception of a grouping by affinity in the vegetable kingdom grew more and more distinct; but it also shows how these men merely followed an instinctive feeling in the matter, and made no enquiry into the cause of the relationship which they perceived.
Nevertheless a great step in advance was thus taken; all the foreign matter introduced into the description of plants by medical superstition and practical considerations was seen to be of secondary importance, and was indeed altogether thrown aside by Kaspar Bauhin; the fact of natural affinity, the vivifying principle of all botanical research, came to the front in its place, and awakened the desire to distinguish more exactly whatever was different, and to bring together more carefully all that was like in kind. Thus the idea of natural affinity in plants is not a discovery of any single botanist, but is a product, and to some extent an incidental product, of the practice of describing plants.
But before the exhibition of the natural affinity gave birth to the first efforts at classification on the part of de l’Obel (Lobelius) and afterwards of Kaspar Bauhin, the Italian botanist Cesalpino (1583) had already attempted a system of the vegetable kingdom on a very different plan. He was led to distribute all vegetable forms into definite groups not by the fact of natural affinity, which impressed itself on the minds of the botanists of Germany and the Netherlands through involuntary association of ideas, but by philosophical reflection. Trained in the philosophy which flourished in Italy in the 16th century, deeply imbued with the doctrines of Aristotle, and practised in all subtleties of the schools, Cesalpino was not the man to surrender himself quietly to the influence of nature on the unconscious powers of the mind; on the contrary, he sought from the first to bring all that he learnt from the writings of others and from his own acute observation of the forms of plants into subjection to his own understanding. Hence he approached the task of the scientific botanist in an entirely different way from that of de l’Obel and Kaspar Bauhin. It was by philosophical reflections on the nature of the plant and on the substantial and accidental value of its parts, according to Aristotelian conceptions, that he was led to distribute the vegetable kingdom into groups and sub-groups founded on definite marks.
This difference in the origin of the systematic efforts of Cesalpino on the one hand and of de l’Obel and Bauhin on the other is unmistakably apparent; the Germans were instinctively led by the resemblances to the conception of natural groups, Cesalpino on the contrary framed his groups on the sharp distinctions which resulted from the application of predetermined marks; all the faults in Bauhin’s system are due to incorrect judgment of resemblances, those of Cesalpino to incorrectness in distinguishing.
But the main point of difference lies in the fact, that the system is presented by de l’Obel and Bauhin without any statement of the principles on which it rests; in their account of it the association of ideas is left to perfect itself in the mind of the reader, as it grew up before in the authors themselves. De l’Obel and Bauhin are like artists, who convey their own impressions to others not by words and descriptions, but by pictorial representations; Cesalpino, on the other hand, addresses himself at once to the understanding of his reader and shows him on philosophic grounds that there must be a classification, and states the principles of this classification; it was on philosophic grounds also that he made the characters of the seed and the fruit the basis of his arrangement, while the German botanists, paying little attention to the organs of fructification, were chiefly influenced by the general impression produced by the plant, by its habit as the phrase now is.
The historians of botany have overlooked the real state of the case as here presented, or have not described it with sufficient emphasis; due attention has not been paid to the fact, that systematic botany, as it began to develope in the 17th century, contained within itself from the first two opposing elements; on the one hand the fact of a natural affinity indistinctly felt, which was brought out by the botanists of Germany and the Netherlands, and on the other the desire, to which Cesalpino first gave expression, of arriving by the path of clear perception at a classification of the vegetable kingdom which should satisfy the understanding. These two elements of systematic investigation were entirely incommensurable; it was not possible by the use of arbitrary principles of classification which satisfied the understanding to do justice at the same time to the instinctive feeling for natural affinity which would not be argued away. This incommensurability between natural affinity and a priori grounds of classification is everywhere expressed in the systems embracing the whole vegetable kingdom, which were proposed up to 1736, and which including those of Cesalpino and Linnaeus were not less in number than fifteen. It is the custom to describe these systems, of which those of Cesalpino, Morison, Ray, Bachmann (Rivinus), and Tournefort are the most important, by the one word ‘artificial’[1]; but it was by no means the intention of those men to propose classifications of the vegetable kingdom which should be merely artificial, and do no more than offer an arrangement adapted for ready reference. It is true that the botanists of the 17th century and Linnaeus himself often spoke of facility of use as a great object to be kept in view in constructing a system; but every one who brought out a new system did so really because he believed that his own was a better expression of natural affinities than those of his predecessors. If some like Ray and Morison were more influenced by the wish to exhibit natural affinities by means of a system, and others as Tournefort and Magnol thought more of framing a perspicuous and handy arrangement of plants, yet it is plain from the objections which every succeeding systematist makes to his predecessors, that the exhibition of natural affinities was more or less clearly in the minds of all as the main object of the system; only they all employed the same wrong means for securing this end, for they fancied that natural affinities could be brought out by the use of a few easily recognised marks, whose value for systematic purposes had been arbitrarily determined. This opposition between means and end runs through all systematic botany from Cesalpino in 1583 to Linnaeus in 1736.
But a new departure dates from Linnaeus himself, since he was the first who clearly perceived the existence of this discord. He was the first who said distinctly, that there is a natural system of plants, which could not be established by the use of predetermined marks, as had been previously attempted, and that even the rules for framing it were still undiscovered. In his Fragments of the date of 1738, he gave a list of sixty-five groups or orders, which he regarded provisionally as cycles of natural affinity, but he did not venture to give their characteristic marks. These groups, though better separated and more naturally arranged than those of Kaspar Bauhin, were like his founded solely on a refined feeling for the relative resemblances and graduated differences that were observed in comparing plants with one another, and this is no less true of the enumeration of natural families attempted by Bernard de Jussieu in 1759. To such of these small groups of related forms as had not been already named both Linnaeus and Jussieu gave names, which they took not from certain marks, but from the name of a genus in each group. But this mode of naming plainly expresses the idea which from that time forward prevailed in systematic botany, that there is a common type lying at the foundation of each natural group, from which all its forms though specifically distinct can be derived, as the forms of a crystal may all be derived from one fundamental form,—an idea which was also expressed by Pyrame de Candolle in 1819.
But botanists could not rest content with merely naming natural groups; it was necessary to translate the indistinct feeling, which had suggested the groups of Linnaeus and Bernard de Jussieu, into the language of science by assigning clearly recognised marks; and this was from this time forward the task of systematists from Antoine Laurent de Jussieu and de Candolle to Endlicher and Lindley. But it cannot be denied, that later systematists repeatedly committed the fault of splitting up natural groups of affinity by artificial divisions and of bringing together the unlike, as Cesalpino and the botanists of the 17th century had done before them, though continued practice was always leading to a more perfect exhibition of natural affinities.
But while natural relationship was thus becoming more and more the guiding idea in the minds of systematists, and the experience of centuries was enforcing the lesson, that predetermined grounds of classification could not do justice to natural affinities, the fact of affinity became itself more unintelligible and mysterious. It seemed impossible to give a clear and precise definition of the conception, the exhibition of which was felt to be the proper object of all efforts to discover the natural system, and which continued to be known by the name of affinity. A sense of this mystery is expressed in the sentence of Linnaeus: ‘It is not the character (the marks used to characterise the genus) which makes the genus, but the genus which makes the character;’ but the very man, who first distinctly recognised this difficulty in the natural system, helped to increase it by his doctrine of the constancy of species. This doctrine appears in Linnaeus in an unobtrusive form, rather as resulting from daily experience and liable to be modified by further investigation; but it became with his successors an article of faith, a dogma, which no botanist could even doubt without losing his scientific reputation; and thus during more than a hundred years the belief, that every organic form owes its existence to a separate act of creation and is therefore absolutely distinct from all other forms, subsisted side by side with the fact of experience, that there is an intimate tie of relationship between these forms, which can only be imperfectly indicated by definite marks. Every systematist knew that this relationship was something more than mere resemblance perceivable by the senses, while thinking men saw the contradiction between the assumption of an absolute difference of origin in species (for that is what is meant by their constancy) and the fact of their affinity. Linnaeus in his later years made some strange attempts to explain away this contradiction; his successors adopted a way of their own; various scholastic notions from the 16th century still survived among the systematists, especially after Linnaeus had assumed the lead among them, and it was thought that the dogma of the constancy of species might find especially in Plato’s misinterpreted doctrine of ideas a philosophical justification, which was the more acceptable because it harmonised well with the tenets of the Church. If, as Elias Fries said in 1835, there is ‘quoddam supranaturale’ in the natural system, namely the affinity of organisms, so much the better for the system; in the opinion of the same writer each division of the system expresses an idea (‘singula sphaera (sectio) ideam quandam exponit’), and all these ideas might easily be explained in their ideal connection as representing the plan of creation. If observation and theoretical considerations occasionally suggested objections to such views, these objections were usually little regarded, and in fact reflections of this kind on the real meaning of the natural system did not often make their appearance; the most intelligent men turned away with an uncomfortable feeling from these doubts and difficulties, and preferred to devote their time and powers to the discovery of affinities in individual forms. At the same time it was well understood that the question was one which lay at the foundation of the science. At a later period the researches of Nägeli and others in morphology resulted in discoveries of the greatest importance to systematic botany, and disclosed facts which were necessarily fatal to the hypothesis, that every group in the system represents an idea in the Platonic sense; such for instance were the remarkable embryological relations, which Hofmeister discovered in 1851, between Angiosperms, Gymnosperms, Vascular Cryptogams and Muscineae; nor was it easy to reconcile the fact, that the physiologico-biological peculiarities on the one hand and the morphological and systematic characters on the other are commonly quite independent of one another, with the plan of creation as conceived by the systematists. Thus an opposition between true scientific research and the theoretical views of the systematists became more and more apparent, and no one who paid attention to both could avoid a painful feeling of uncertainty with respect to this portion of the science. This feeling was due to the dogma of the constancy of species, and to the consequent impossibility of giving a scientific definition of the idea of affinity.
This state of things finally ceased with the appearance of Darwin’s first and best book on the origin of species in 1859; from a multitude of facts, some new, but most of them long well-known, he showed that the constancy of species was no longer an open question; that the doctrine was no result of exact observation, but an article of faith opposed to observation. The establishment of this truth was followed almost as a matter of course by the true conception of that which had been hitherto figuratively called affinity; the degrees of affinity expressed in the natural system indicated the different degrees of derivation of the varying progeny of common parents; out of affinity taken in a figurative sense arose a real blood-relationship, and the natural system became a table of the pedigree of the vegetable kingdom. Here was the solution of the ancient problem.
Darwin’s theory has this special interest in the history of the science, that it established clearness in the place of obscurity, a scientific principle in place of a scholastic mode of thought, in the domain of systematic botany and morphology. Yet Darwin did not effect this change in opposition to the historical development of our science or independently of it; on the contrary his great merit is that he has correctly appreciated the problems long existing in systematic botany and morphology from the point of view of modern research, and has solved them.
That the constancy of species is incompatible with the idea of affinity, that the morphological (genetic) nature of organs does not proceed on parallel lines with their physiological and functional significance, are facts which were known in botany and zoology before the time of Darwin; but he was the first to show, that variation and natural selection in the struggle for existence solve these problems, and enable us to conceive of these facts as the necessary effects of known causes; it is at the same time explained, why the natural affinity first recognised by de l’Obel and Kaspar Bauhin cannot be exhibited by the use of predetermined principles of classification, as was attempted by Cesalpino.
[CHAPTER I.]
The Botanists of Germany and the Netherlands from Brunfels to Kaspar Bauhin[2].
1530-1623.
When those who are accustomed to modern botanical literature take up for the first time the works of Otto Brunfels (1530), Leonhard Fuchs (1542), Hieronymus Bock (Tragus), or of the later authors Rembert Dodoens (Dodonäus), Charles de l’Écluse (Carolus Clusius), Matthias de l’Obel (Lobelius, 1576), or even those of Kaspar Bauhin from the beginning of the 17th century, they are surprised not only by the strange form, the curious and unfamiliar accessories from which what is really useful must be laboriously extracted, but still more by the extraordinary poverty of thought which characterises these composers of usually very thick folios. If however instead of travelling backwards from the present time they pursue the opposite direction; if they have previously occupied themselves with the botanical views of Aristotle and the comprehensive botanical works of his disciple Theophrastus of Eresus, with Pliny’s Natural History and the medical science of Dioscorides; if they have made themselves acquainted with the botanical literature of the middle ages and noted how it continually grows less and less valuable, and have proceeded through the works of Albertus Magnus, as prolix as they are deficient in ideas, to the ‘Hortus Sanitatis’ (Garden of Health), the popular work on natural history before and after 1500, and similar productions, then certainly they receive a very different and almost imposing impression even from the first herbals, those of Brunfels, Bock, and Fuchs. These books will appear to them almost modern in comparison with the last-named productions of medieval superstition, nor will they fail to perceive that a new epoch of natural science commenced with these men, and above all that they laid the foundations of modern botany. They give us, it is true, nothing but separate descriptions of the wild and cultivated plants of Germany, and these for the most part of common occurrence, arranged by Fuchs alphabetically, by Bock grouped under the heads of herbs, shrubs, and trees, and following one another under each head in the most motley order; it is true that these descriptions are so naive and inartistic as hardly to offer points of comparison with modern scientifically correct diagnoses; but the great point is, that they are taken from the plants as they lay before the writers, who had often seen and carefully examined them. Woodcuts are added to supply any defects in the description, and to give a clear idea of the plant intended by the name; and these figures, which always give the whole plant and were drawn immediately from nature by the hands of practised artists, are so true to nature that a botanist’s eye at once recognises in every case the object meant to be represented. These figures and descriptions (the latter are wanting in Brunfels[3], 1530) would have rendered a great service to the science, even if they had not been as good as they are; for botanical literature had sunk so low, that not only were the figures embellished with fabulous additions, as in the ‘Hortus Sanitatis,’ and sometimes drawn purely from fancy, but the meagre descriptions of quite common plants were not taken from nature, but borrowed from earlier authorities and eked out with superstitious fictions. The powers of independent judgment were oppressed and stunted in the middle ages, till at last the very activity of the senses, resting as it does to a great extent on unconscious operations of the understanding, became weak and sickly; natural objects presented themselves to the eye even of those who made them their study in grotesquely distorted forms; every sensuous impression was corrupted and deformed by the influence of a superstitious fancy. In comparison with these perversions the artless descriptions of Bock appear suitable and true, and are refreshing from their immediate contact with nature; while in the more learned Fuchs criticism of other writers is already seen united with actual examination of natural objects. Great was the gain when men began once more to look at plants with open eyes, to take pleasure in their variety and beauty. It was not necessary for a while that they should speculate on the nature of plants, or the cause of plant-life; time enough for that when sufficient practice had been gained in the perception of their resemblances and differences.
The German fathers of botany connected their labours with the botanical literature of classical antiquity only so far as they sought to recognise in the plants of their own country those named by Theophrastus, Dioscorides, Pliny and Galen. The attempt to do this indeed led to many mistakes, for the descriptions of the ancient botanists were very imperfect and often quite unserviceable for the recognition of the plants described. In this point therefore the compilers of herbals found no models worthy of imitation in the old writers. But in seeking to recover a knowledge of the medicinal plants of the Greek physicians[4], they were compelled to compare together a great variety of native plants, and thus to exercise and perfect the faculty of apprehending differences of form. This mode of proceeding, arising out of medical requirements, directed the attention entirely to the individual form, which was also the chief thing required in the interest of pure science, and much more was thus gained than if these men had only followed the philosophical writings of Aristotle[5] and Theophrastus[6]. The Greek authors built their views on the philosophy of botany on very weak foundations; scarcely a plant was known to them exactly in all its parts; they derived much of their knowledge from the accounts of others, often from dealers in herbs. From this scanty material and from various popular superstitions had Aristotle formed his views on the nature of plants, and if Theophrastus possessed more experimental knowledge, he still saw facts in the light of his master’s philosophical doctrines. If we succeed in the present day in extracting much that is accurate from the writings of Aristotle and Theophrastus, it was nevertheless well that the first compilers of herbals ceased to pay attention to them, and occupied themselves with accumulating descriptions of individual plants worked out by themselves with all possible exactness. History shows that in this way a new science arose in the course of a few years, while the philosophical botany of Aristotle and Theophrastus has led to no important result. Moreover we shall see how even in the hands of a philosophically gifted and scholarly man like Cesalpino the teaching of Aristotle had only a mischievous effect on the study of plants.
If the compilers of herbals did not aim at deducing general conclusions from their observations, yet the continually accumulating descriptions of individual forms gradually gave rise of themselves to perceptions of an abstract and more comprehensive character. The feeling for resemblance and difference of form especially was developed, and finally the idea of natural relationship; and though this idea was as yet by no means worked out with scientific precision, it was nevertheless, even in the indistinct form in which it appears in de l’Obel in 1576 and more clearly in Kaspar Bauhin in 1623, a result of the highest value, and one of which neither learned antiquity nor the middle ages had ever caught a glimpse. The perception of a natural affinity among plants could only be obtained from exact description a thousand times repeated, never from the abstractions of the Aristotelian school, which rested essentially on superficial observation. It appears then that the scientific value of the herbals of the 16th century lay mostly in the description of such plants as every botanist found in a somewhat limited portion of his native land, and considered worth his notice; at the same time the later compilers endeavoured to give a universal character to each herbal by admitting plants which had not been actually seen by the writer; each as far as possible gathered from his predecessors all that they had seen, and added what he had himself seen that was new; but in contrast with the previous centuries the peculiar merit of each new herbal was held to depend not on what the compiler had borrowed from his predecessors, but on what he had added from his own observation. Hence every one was anxious to introduce into his work as many plants unknown till that time or unnoticed as he possibly could, and the number of descriptions of individual forms mounted rapidly up; in Fuchs in 1542 we find about five hundred species described and figured, but in 1623 the number of species as enumerated by Kaspar Bauhin had risen to six thousand. As the botanists were spread over a large part of Germany, Fuchs in Bavaria and afterwards at Tübingen, Bock on the middle Rhine, Konrad Gesner at Zürich, Dodoens and de l’Obel in the Netherlands, a territory of considerable extent was thus examined; it was enlarged by the contributions which travellers brought or transmitted to the botanists, and de l’Écluse especially traversed a large part of Germany and Hungary and even of Spain, and eagerly collected and described the plants of those countries. During this period also the number of known plants was increased from Italy, partly by the exertions of Italian botanists, such as Mattioli, and partly by travelling Germans. The first flora of the Thüringer-Wald was written by Thal, but not published till after his death in 1588. Botanical gardens even, though in more modest form than in our day, were already helping in the 16th century to add to the knowledge of plants; the first were formed in Italy, as at Padua in 1545, at Pisa in 1547, at Bologna in 1567 under Aldrovandi, afterwards under Cesalpino. Soon similar collections of living plants were made in the north; in 1577 a botanic garden was founded at Leyden, over which de l’Éluse long presided, in 1593 at Heidelberg and at Montpellier; in the course of the next century the number of these gardens was considerably increased.
The preserving of dried plants, the formation of the collections which we now call herbaria, dates from the 16th century; at that time however the word herbarium meant a book of plants. In this matter also the Italians led the way. According to Ernst Meyer, Luca Ghini seems to have been the first who made use of dried plants for scientific purposes, and his two pupils Aldrovandi and Cesalpino are said to have formed the first herbaria in our sense of the word; one of the first collections of the kind, perhaps of the date of 1559, was the herbarium formed by Ratzenberger, which was discovered in the museum at Cassel a few years since and described by Kessler.
These are matters somewhat external to our immediate subject, but they show how lively an interest was taken in botany in the latter half of the sixteenth century; this is still more shown by the great number of books of plants, published with numerous and expensive plates and in some cases going through several editions. But the artistic and scientific value of the drawings, which were appended to the descriptions and in later herbals were reckoned by thousands, did not keep equal pace with their number; Fuchs’ splendid figures remained unapproached, and gradually, as the distance from Dürer’s time increased, the wood-cuts grew smaller and poorer[7], and sometimes even quite indistinct. The art of describing on the contrary continually improved; the descriptions became fuller, and gradually a certain method appeared in assigning marks and in estimating their value; critical remarks on the identity or non-identity of species, the separation of forms previously considered to be alike, and similar matters occur more frequently. The descriptions in de l’Éluse may in fact claim to be called scientific; in Kaspar Bauhin they appear in the form of terse and methodical diagnoses.
The most remarkable thing to us in these descriptions from Fuchs and Bock to Bauhin is the striking neglect of the flowers and fruit. The earliest descriptions, especially those of Bock, endeavour to depict the form of the plant in words, to render directly the impression on the senses; special attention was paid to the shape of the leaves, the nature of the ramification, the character of the roots, the size and colour of the flowers. Konrad Gesner[8] was the only one who bestowed a closer attention on the flowers and parts of the fruits; he figured them repeatedly, and recognised their great value for the determination of affinity, as we learn from his expressions in his letters; but the much occupied and much harassed man died before he could complete the work on plants which he had long been preparing, and when in the 18th century Schmidel published Gesner’s figures, which meanwhile had passed through various hands, the work too long delayed remained useless to a science which had already outstripped it.
It will be gathered from the above remarks, that we find in these authors no approach to a system of morphology founded on a comparative examination of the parts of plants, and therefore no regular technical language. Still the more learned among them felt the necessity of connecting the words they used in describing a plant with a fixed sense, of defining their conceptions; and though their first efforts in this direction were weak, they deserve notice, because they show more than anything else how great has been the advance in the study of nature from the 16th century to the present day.
The first attempt to establish a botanical terminology is to be found as early as 1542 in the ‘Historia Stirpium’ of Leonhard Fuchs[9]. Four pages at the beginning of the work are thus occupied. A considerable number of words are explained in alphabetical order—the mode of arrangement which he followed also in describing his plants. It is difficult to give a clear idea of this the first botanic terminology by selected examples; yet the attempt must be made, because it is in this way only that we learn to see from what feeble beginnings the later scientific terminology and morphology has been developed. Thus we read: ‘Acinus’ denotes not merely, as many believe, the grains inside the grape, but the whole fruit, which consists of juice, of a fleshy portion with the stones (‘vinaceis’), and of the outer skin. Galen is quoted as authority for the following explanation: ‘Alae’ are said to be the hollows (angles) between the stem and its branches (the leaves), from which new sprouts (‘proles’) proceed. ‘Asparragi,’ the germs of herbs which appear before the leaves and the first edible shoots are developed. ‘Baccae’ are smaller ‘foetus’ of herbs, shrubs, and trees, which appear separate and isolated on the plant, as for example laurel-berries (‘partus lauri’), and differ from acini, inasmuch as these are more crowded together. ‘Internodium’ is that which lies between the articulations or knees. ‘Racemus’ is used for the bunch of grapes, but does not belong to the vine only, but also to the ivy and other herbs and shrubs which bear clusters of any kind. The majority of such explanations of names concern the forms of the stem and the branches, but the most remarkable thing about the whole list is, that it does not include the words flower and root; yet under the word ‘julus’ occurs the statement, that it is that which in the hazel ‘compactili callo racematim cohaeret,’ and may be described as a long worm borne on a special pendent stalk and coming before the fruit. Though the word flower is not explained, yet some parts of the flower are mentioned; thus it is said, ‘stamina sunt, qui in medio calycis erumpunt apices, sic dicta quod veluti filamenta intimo floris sinu prosiliant.’ The explanation of the word fruit may be added: ‘Fructus, quod carne et semine compactum est; frequenter tamen pro co, quod involucro perinde quasi carne et semine coactum est, accipi solet.’
Progress in this direction was slow but still recognisable. In the last edition of the ‘Pemptades’ of Dodoens[10] of the year 1616, a folio volume of 872 pages, only one page and a third are devoted to the explanation of the parts of plants; but the selection of the words explained and the substance of the explanations hit the essential points better than in Fuchs. We find for instance: Root (‘radix, ῥίζα’) is the name given in the tree and in every other plant to the lower part, by which it penetrates into the earth and cleaves to it, and by which it draws its nourishment. This part, unlike the leaves which are usually deciduous, is common to all plants, a few only excepted which live and grow without roots, such as Cassytha, Viscum, and the plant called ‘Hyphear,’ Fungi, Mosses, and Fuci, all which are however usually reckoned among φῦτα. ‘Caudex’ is in trees and shrubs that which springs from the root and rises above the ground, and by which the nourishment is carried upwards; the same part is called in herbs caulis or cauliculus. Leaf (‘folium’) is in every plant that which clothes and adorns it, and without which trees and other plants appear naked. The definition of a flower would lose in a translation: ‘flos, ἄνθος, arborem et herbarum gaudium dicitur, futurique fructus spes est; unaquaeque etenim stirps pro natura sua post florem partus ac fructus gignit.’ The parts of the flower are with him the calyx (‘calyx’), in which the blossom is at first enclosed and with which the ‘foetus’ is soon surrounded, stamens (‘stamina’) which arise like threads from the depth of the blossom and from the calyx, and ‘apices’ (anthers), certain thickish appendages on the summit of the stamens. ‘Julus’ (catkin) is that which hangs down round and long in place of the flower, as in the walnut, hazel, mulberry, beech, and other trees. ‘Fructus’ is that in which the seed is formed, but frequently it is itself the seed, as where the latter is not enclosed in anything else and is formed naked. We must not be led by these words to think of our Gymnosperms, but must understand that here, as with all botanists till the time of A. L. de Jussieu and Joseph Gärtner (1788), naked seeds mean dry indehiscent fruits.
De l’Obel, from whom especially we might have looked for similar explanations, has given none.
The absence of more profound comparative examination of the parts of plants, as shown in the examples of terminology here adduced, may serve as an additional support of the assertion, that natural affinity was not inferred from exact comparison of the form of organs, but was the result of a feeling arising from the likeness of habit directly apprehended by the senses, that is by the collective impression produced by the whole plant.
Passing to the consideration of the attempts in systematic botany made by the Germans in this period, the chief thing to notice is, that the division into the main groups of trees, shrubs, undershrubs, and herbs was the one generally adopted; these groups were borrowed from antiquity and were maintained even by the special systematists, from Cesalpino to the beginning of the 18th century; nor was any change made in principle when these four groups were reduced to three or two (trees and herbs). It was moreover considered to be self-evident that trees were the most perfect plants. Hence when relationship is spoken of in subsequent remarks, it must be understood that this holds good only within the groups just mentioned. The classifications of the German and Dutch botanists not only sprang from the describing of individual plants, but they were originally in a certain sense identical with it. In undertaking to describe individual forms, the first task was to separate those which closely resembled one another, for the resemblance of systematically-allied plants is often so great, that to distinguish them specifically requires consideration and careful comparison. The resemblance is more obvious than the difference. There are moreover many plants which are entirely distinct from one another in their inner nature, but which appear strikingly alike if we regard the impression produced immediately on the senses, and the converse of this statement is equally true. Hence the attempt to circumscribe and fix individual forms in the act of describing was at once found to involve difficulties, the solution of which leads directly to the conception of some kind of arrangement. A comparison of the herbals of Fuchs and Bock up to Kaspar Bauhin shows very plainly how these difficulties were gradually overcome, how the describing of single species led necessarily, and without the intention of the describer, to considerations of a distinctly systematic character. Where the species in a group of forms, which we now designate as a genus or family, closely resemble each other in habit, there arose of itself the instinctive feeling that such forms belong to one another. This feeling asserted itself in words when, as was done from the first, a number of such forms were without conscious reflection designated by the same name; thus, to mention one of many examples, we find Bock applying the name Wolfsmilk, Euphorbia, not to one species of the genus, but to several, which he then distinguishes by epithets (common, least, cypress, sweet). The customary mode of expression in the herbals is very instructive on this point; there are, they say, two or more of this or that plant which have not been hitherto distinguished. But this feeling of connection and similarity of kind was produced not only by forms that were closely allied, but also by such as belong to extensive groups of the system; thus the words moss, lichen, fungus, alga, fern, had long served to include a great number of distinct forms, though the separation of these groups had nowhere in truth been carried out with logical precision.
These remarks are important as serving to show in the most decisive manner the incorrectness of the assertion, that the study of organisms sprang from the recognition of individual species; that it is this which is directly given, and that without it no advance in the science is possible. The historical fact rather is, that descriptive botany began often, perhaps most often, not with species but with genera and families, that very often at first whole groups of forms were conceived of as unities, which had to be divided later and of set purpose into separate forms; and up to the present day one part of the task of the systematist is to undertake the splitting up of forms previously regarded as identical. The notion that the species is the object originally presented to the observer, and that certain species were afterwards united into genera, is one that was invented in post-Linnaean times under the dominion of the dogma of the constancy of species; it happened so sometimes, but just as often the genus was the object first presented, and the task of the describer was to resolve it into a number of species. In the 16th century the conception neither of genus or species had yet been defined; for the botanists of that period genera and species had the same objective reality. But, in the process of continually making the descriptions of individual plants more exact, forms once separated were united, and those before assumed to be identical were separated, till it gradually became apparent that both operations must be pursued with system and method. It cannot therefore be exactly said that somebody first established the species, another the genus, and a third person again the larger groups. It is more correct to say that the botanists of the 16th century carried out this process of separation up to a certain point without intending it, and in the effort to give the greatest possible preciseness to their descriptions of individual forms. It lay therefore in the nature of the case, that those groups which we call genera and species should first be cleared up, and we find in fact at the end of this period in Kaspar Bauhin the genera already distinguished by names, if not by characters; the species by names and characters. Together with these smaller groups, many more comprehensive ones, which we now designate families, were also marked off and supplied with names, which are still in use. The 16th century established the groups and names of Coniferae, Umbelliferae, Verticillatae (Labiatae), Capillares (Ferns), and others. It is true that the determination of the limits of these groups by distinct marks was not yet attempted, but the plants belonging to these groups were again and again treated of in special chapters or ranged in due succession one after another. But as long as this was done to some extent without design, and the real meaning of this relationship was not yet recognised, other considerations of very various kinds influenced the composition of the books and disturbed the natural arrangement. The feeling for natural affinity supplants all other considerations in de l’Obel first, and after him much more completely in Kaspar Bauhin.
Enough perhaps has now been said to render the main result of the botanical efforts of the period, which we are considering, intelligible to the reader; but a clear view of the method of describing plants at that time, and of the way in which systematic botany came into being, can only be shown by examples; and if we proceed to give some here, it is with the purpose with which figures copied as exactly as possible from nature are added to treatises on natural history, because a real understanding is only to be gained in this way. The botanical literature of the 16th century is so different from that of the 19th, that a very indistinct idea of it could be obtained from a statement of results expressed in modern terms.
Fuchs, Historia Stirpium, 1542.
The common plant now known as Convolvulus arvensis is there called Helxine cissampelos, and is described in the following manner:
‘Nomina.—Ἑλξινὴ κισσάμπελος Graecis, Helxine cissampelus et Convolvulus Latinis nominatur. Vulgus herbariorum et officinae Volubilem mediam et vitealem appellant, Germani Mittelwinden oder Weingartenwinden. Recte autem Cissampelos dicitur, in vineis enim potissimum nascitur et folio hederaceo. Convolvulus vero quod crebra revolutione vicinos frutices et herbas implicet.
Forma.—Folia habet hederae similia, minora tamen, ramulos exiguos circumplectentes quodcumque contigerint. Folia denique ejus scansili ordine alterna subeunt. Flores primum candidos lilii effigie, dein in puniceum vergentes, profert. Semen angulosum in folliculis acinorum specie.
Locus.—In vineis nascitur, unde etiam ei appellatio cissampeli, ut diximus, indita est.
Tempus.—Aestate, potissimum autem Julio et Augusto mensibus, floret.‘
Hieronymus Bock[11], at page 299 of his ‘Herbal,’ published at Strassburg in 1560, describes the same plant and Convolvulus sepium as follows:
‘Of the white wind-bell.
‘Two common wind-plants grow everywhere in our land with white bell-flowers. The larger prefers to dwell by hedges, and creeps over itself, twists and twines, etc. The little wind- or bell-flower (Convolvulus arvensis) is like the large one with its roots, round stems, leaves and bell-flowers, in all things smaller, thinner, and shorter. Some flowers on this plant are quite white, some of a beautiful flesh colour, painted with reddish brown streaks. It grows in dry meadows, in herb-and onion-gardens, and does harm therein, because with its creeping and twining it oppresses other garden herbs, and is also bad to exstirpate, because the thin white rootlets make their way deep downwards, spread very widely, and are continually putting forth new and young clusters like hops.’
Then follows a long paragraph on the names, that is, a critical review of the opinions of different writers on the question, which of Dioscorides’ or Pliny’s names should be applied to the plant described. ‘I must think,’ says Bock, ‘that this flower is a wild sort, Scammonia Dioscoridis (but harmless), which herb Dioscorides also calls colophonia, dactylion, apopleumenon, sanilum, and colophonium,’ and so on. Then follows a chapter on its virtue and effect externally and internally.
As regards the arrangement of the 567 species described by Bock, he divides his book into three parts, the first and second containing the smaller herbs, the third the shrubs and trees. In each part closely allied plants are generally described in larger or smaller numbers one immediately after another, though the compiler is all the time under the influence of very various considerations, and follows no general principle. For instance, our Convolvulus stands in the midst of a number of other very different plants, which either climb as the ivy, or twine with tendrils as Smilax; then follows Lysimachia Nummularia, which simply runs along the ground, then the hop, Solanum Dulcamara, Clematis, Bryonia, Lonicera, and different Cucurbitaceae; immediately after come the Burdocks, Teasels, and Thistles, and these are followed by some Umbelliferae. The whole work is conceived in a similar spirit; the feeling for relationship is clearly to be traced within very narrow circles, but it finds imperfect expression and is frequently disturbed by reference to biological habit; this appears especially in the beginning of the third part, which treats of shrubs generally, shrubs which form hedges, and trees, ‘as they grow in our German land’; the first chapter is on the fungi which grow on trees, the second on some mosses, and these are followed immediately by the mistletoe. Then come the heather and some smaller shrubs, and finally larger and the largest trees. The chapter on Fungi under the section ‘Of names’ contains a statement of views on the nature of fungi, such as are often repeated even into the 17th century: ‘Mushrooms are neither herbs nor roots, neither flowers nor seeds, but merely the superfluous moisture of the earth and trees, of rotten wood and other rotten things. From such moisture grow all tubera and fungi. This is plain from the fact that all the above-mentioned mushrooms, those especially which are used for eating, grow most when it will thunder or rain, as Aquinas Ponta says. For this reason the ancients paid peculiar regard to them, and were of opinion that tubera, since they come up from no seed, have some connection with the sky; Porphyrius speaks also in this manner, and says that fungi and tubera are called children of the gods, because they are born without seeds and not as other kinds.’
We pass over Valerius Cordus, Conrad Gesner, Mattioli[12], and some other unimportant writers, and turn to Dodoens, de l’Écluse, and Dalechamps, in whom a marked tendency to orderly arrangement appears, though the principle of arrangement in all three lies essentially in points external and accidental, and above all in the relations of the plant-world to mankind. Within the divisions thus artificially formed a constantly increasing attention is paid to natural affinities, but at the same time allied forms are separated without scruple in deference to the artificial principle of classification. It can also be plainly seen, that these writers think more of giving some order to their matter than of discovering the arrangement that will be in conformity with nature. It is impossible to give the reader a good idea of these classifications in our scientific language; it would be necessary to transcribe them. For brevity’s sake we will here quote de l’Écluse only[13], the best of the three writers named above. In his ‘Rariorum plantarum historia,’ which appeared as early as 1576, but which lies before the writer of these pages in the edition of 1601, the first book treats of trees, shrubs, and undershrubs; the second of bulbous plants; the third of sweet-smelling flowers; the fourth of those without smell; the fifth of poisonous, narcotic, and acrid plants; the sixth of those that have a milky juice, and of Umbelliferae, Ferns, Grasses, Leguminosae, and some Cryptogams.
A similar arrangement is found in Dalechamps[14]; that of Dodoens in his ‘Pemptades’ is more perplexed and unnatural; but the design in both of them is evidently much the same as that of de l’Écluse. This design is best seen from the introductory observations to each book; de l’Écluse, for instance, says at page 127, ‘Having treated of the history of trees, shrubs, and undershrubs, and put these together in the preceding book, we will now in this second book describe such plants as have a bulbous or tuberous root, many of which attract and delight the eyes of all persons in an extraordinary degree by the elegance and variety of their flowers, and which therefore ought not to have the lowest place assigned to them among garland-plants (‘inter coronarias’). We will begin with the plants of the lily kind, on account of their size and the beauty of their flowers, etc. etc.‘ The introductions to the several books of the ‘Pemptades’ of Dodoens are more learned and more diffuse. It is plain that the composers of these works had no thought of arranging their matter on the principles of a true natural system, but were only anxious to give some kind of order to their descriptions of individual plants. Hence their divisions do not appear under the names of classes and subdivisions (‘genera majora et minora,’ as they would have been called at that time), but they are sections of the whole work kept as symmetrical as was possible. If we would discover in these works whatever may really lay claim to systematic value, we must not rely on the sections as they are typographically distinguished, but must observe within each of them the order in which the plants are given, and then it becomes apparent that within the frame once established forms naturally allied are, as far as may be, grouped together. For instance, we find in the second book of de l’Écluse’s work first of all a long list of true Liliaceae and Asphodeleae, Melanthaceae, and Irideae described in unbroken succession; then comes Calamus, and then without any explanation a number of the Ranunculaceae, among which the genera Ranunculus and Anemone are very well distinguished; but then follows the genus Cyclamen with several species, and next a number of Orchideae, in the middle of which appear Orobanche and Corydalis, followed by Helleborus niger, Veratrum album, Polygonatum, and others. So it is in the other sections, though in general the species of a genus stand together, and even the genera of a family are not unfrequently united; but with all this there are no proper breaks, because other considerations are perpetually disturbing the feeling for natural relationship. The descriptions of de l’Écluse are generally commended, and they deserve to be commended for their fulness of detail and their attention to the structure of the flowers, though he, like de l’Obel and Dodoens, describes the leaves more minutely than any other part of the plant.
With de l’Obell[15], as has been already observed, the feeling for natural affinity declares itself for the first time so decidedly as to outweigh if not entirely to set aside all other considerations. The fact is disclosed to us in the preface to his ‘Stirpium adversaria nova’ of 1576, where these words occur: ‘proinde adversariorum voce novas veteribus additas plantas et novum ordinem quadantenus innuimus. Qui ordo utique sibi similis et unus progreditur ducitque a sensui propinquioribus et magis familiaribus ad ignotiora et compositiora, modumque sive progressum similitudinis sequitur et familiaritatis, quo et universim et particulatim, quantum licuit per rerum varietatem et vastitatem, sibi responderet. Sic enim ordine, quo nihil pulchrius in coelo aut in sapientis animo, quae longe lateque disparata sunt unum quasi fiunt, magno verborum memoriae et cognitionis compendio, ut Aristoteli et Theophrasto placet.’
We must not indeed expect to find that de l’Obel really produced a natural system of plants; but his ‘Observationes’ still more than his ‘Adversaria’ attest his efforts to arrange plants according to their resemblances in form; and in these efforts he is guided not by instinct merely and the general habit, but mainly and with evident purpose by the form of the leaves; thus beginning with Grasses, which have narrow, long, and simple leaves, he proceeds to the broader-leaved Liliaceae and Orchideae; then passing on to the Dicotyledons he exhibits the main groups in fairly well limited masses. Still the Ferns appear in the middle of the Dicotyledons on account of the form of their leaves, while on the other hand, the Cruciferae, Umbelliferae, Papilionaceae and Labiatae remain but little disturbed in their continuity by secondary considerations.
The progress of botanical science in the period which we have been considering reaches its highest point in the labours of Kaspar Bauhin[16], as regards both the naming and describing of individual plants and their classification according to likeness of habit. In Bauhin all secondary considerations have disappeared; his works may be called botanical in the strict scientific meaning of the word, and they show how far it is possible to advance in a descriptive science without the aid of a general system of comparative morphology, and how far the mere perception of likeness of habit is a sufficient foundation for a natural classification of plants; it was scarcely possible to make greater advances on the path pursued by the botanists of Germany and the Netherlands.
The descriptions of species in the ‘Prodromus Theatri Botanici’ of Kaspar Bauhin (1620) notice all obvious parts of the plant with all possible brevity and in a fixed order; the form of the root, height and form of the stem, characters of the leaves, flowers, fruit, and seed are given in concise sentences seldom occupying more than twenty short lines; the description of a single species is here in fact developed into an art and becomes a diagnosis.
A still higher value must be set on the fact, that in Kaspar Bauhin the distinction between species and genus is fully and consciously carried out; every plant has with him a generic and a specific name, and this binary nomenclature, which Linnaeus is usually thought to have founded, is almost perfectly maintained by Bauhin, especially in the ‘Pinax’; it is true that a third and fourth word is not unfrequently appended to the second, the specific name, but this additional word is evidently only an auxiliary. It is remarkable on the other hand, that he has added no characters to the names of the genera; it is only from the name that we know that several species belong to one genus; we might almost believe that the characters of the genus are intended to be supplied by the strange etymological explanation appended in italics to the generic name. These fanciful etymologies maintained themselves to the end of the 17th century, when Tournefort did battle with them; they were an evil which sprang in a great measure from Aristotelian and scholastic modes of thought, and from the belief that it was possible to conceive of the nature of a thing from the original meaning of its name.
Nothing shows better the earnestness of Bauhin’s research than the fact, that he devoted the labour of forty years to his ‘Pinax,’ in order to show how each one of the species given by him was named by earlier botanists. The example already given from Fuchs shows how many names a plant had received by the middle of the 16th century; even in Dioscorides and Pliny we find a whole row of names given for a single plant, and the botanists of Fuchs’ time used their utmost endeavours to attach the names in Dioscorides and other ancient writers to particular plants found in central Europe. Dioscorides, Theophrastus, and Pliny either add no descriptions to the names of their plants, or they describe them in so unsatisfactory a manner, that it was a very difficult task for the science of that day, as it is still for us, to recognise the plants of the ancient writers; hence arose such a confusion of names that the reader of a botanical work can never be sure whether the plant of one author is the same as that of another with the same name. A description of a plant is therefore usually accompanied in the 16th century by a critical enquiry how far the name used agrees with that of other authors. Kaspar Bauhin sought to put an end to this condition of uncertainty by his ‘Pinax,’ in which he showed in the case of all species known to him what were the names given to them by the earlier writers, and he has thus enabled us to see our way through the nomenclature of the period of which we are speaking; the ‘Pinax’ is in a word the first and for that time a completely exhaustive book of synonyms, and is still indispensable for the history of individual species—no small praise to be given to a work that is more than 250 years old.
It would not have been unsuitable to the purpose of the author of the ‘Pinax,’ if he had allowed himself to give the plants in alphabetical order, but instead of this we find a careful arrangement according to natural affinities. This directly proves what is also confirmed by the ‘Prodromus,’ that Bauhin regarded such an arrangement as of the greatest importance. In this point, as in others, he goes far beyond his predecessors; he pursues the same method as de l’Obel had pursued forty years before, but he carries it out more thoroughly. At the same time he shares with his predecessors the peculiarity of not distinguishing the larger groups, which with some exceptions answer to our present families, by special names or by descriptions; it is only from the order in which the species follow one another that we can gather his views on natural relationship. It follows therefore that the natural families, so far as they are distinguishable in Bauhin’s works, have no sharp bounding lines; we might almost conclude that he purposely avoided assigning such limits, that he might be able to pass without interruption from one chain of relationship to another.
Like de l’Obel, Bauhin proceeds in his enumeration from the supposed most imperfect to the more perfect forms, beginning with the Grasses and the majority of Liliaceae and Zingiberaceae, passing on to dicotyledonous herbs, and ending with shrubs and trees.
The Cryptogams that were known to him stand in the middle of the series of dicotyledonous herbs, between the Papilionaceae and the Thistles, the Equisetaceae being reckoned among the Grasses. On the great distinction between Cryptogams and Phanerogams the views of Bauhin were evidently less clear than those of many of his predecessors; but it will not seem strange that he should place some Phanerogams, as for instance the Duckweeds, among the Cryptogams and the Salviniaceae among the Mosses, and unite the Corals, Alcionieae, and Sponges with the Seaweeds, when we consider that it was not till the middle of the 18th century that more correct views arose in respect to these forms, and that Linnaeus himself could not decide whether the Zoophytes should be excluded from the vegetable kingdom and ranked with animals. The knowledge of plants in the scientific sense of the word was till the beginning of the 19th century limited to the Phanerogams; and in speaking of principles and methods in descriptive botany before that time we must think only of the Phanerogams, or at most of the Phanerogams and the Ferns. The methodical examination of the Cryptogams belongs to quite recent botanical research. The matter is here alluded to only in connection with the fact, that it is from the works of Kaspar Bauhin, a writer of ability, in whom the first period of scientific botany culminates, that we most clearly see how great the advance has been since his time.
[CHAPTER II.]
Artificial Systems and Terminology of Organs from Cesalpino to Linnaeus.
1583-1760.
While botany was being developed in Germany and the Netherlands in the manner described in the previous chapter, and long before this process of development reached its furthest point in Kaspar Bauhin, Andrea Cesalpino in Italy was laying down the general plan, on which the further advance of descriptive botany was to proceed in the 17th and till far into the 18th century; all that was done in the 17th century in Germany, England, and France towards furthering morphology and systematic botany was done with a reference to Cesalpino’s principles, whether these were accepted and made use of, or whether it was sought to refute them. This connection with Cesalpino became gradually less close and less obvious, being concealed by new points of view and by the increase of material for observation; but Cesalpino’s ideas on the theoretical principles of systematic botany and the nature of plants appear so plainly, even in the views of Linnaeus, that no one can read both authors without lighting not unfrequently upon passages in Linnaeus’ ‘Fundamenta’ or in his ‘Philosophia Botanica,’ which remind him of Cesalpino, and even upon sentences borrowed from him. As we saw in Kaspar Bauhin the close of the course of development commenced by Fuchs and Bock, so we may regard Linnaeus as having built up and completed the edifice of doctrine founded by Cesalpino.
Cesalpino comes before us, in strong contrast with the simple-minded empiricism of the German fathers of botany, as the thinker in presence of the vegetable world. Their main task was the amassing descriptions of individual plants. Cesalpino made the material gathered by experience the subject of earnest reflection; he sought especially to obtain universals from particulars, important principles from sensuous perceptions; but as his forms of thought were entirely Aristotelian, it was inevitable that his interpretation of the facts should introduce into them much that would have to be got rid of subsequently by the inductive method. Cesalpino differs also from the German botanists in another respect; he did not rest satisfied with the general impression produced by the plants, but carefully examined the separate parts and paid attention to the small and concealed organs; he was the first who converted observation into real scientific research; and thus we find in him a remarkable union of inductive natural science and Aristotelian philosophy, a mixture which gives a peculiar character to the theoretical efforts of his successors down to Linnaeus.
Cesalpino was moreover much before his time in his mode of contemplating the vegetable kingdom, seeking always for philosophical combinations and comprehensive points of view. His work which appeared in 1583 exercised no perceptible influence on his contemporaries; a trace of such influence only may be seen in Kaspar Bauhin thirty or forty years later, while the work of the botanists who followed Bauhin down to 1670 was confined everywhere to increasing the knowledge of individual plants. With this object travels were undertaken after 1600 to all parts of the world; many new botanic gardens were added to the few which had been founded in the 16th century—as at Giessen in 1617, at Paris in 1620, at Jena in 1629, at Oxford in 1632, at Amsterdam in 1646, at Utrecht in 1650. Instead of endeavouring to embrace with their labours the whole vegetable kingdom, botanists preferred to devote themselves to the examination of single districts. This gave rise to the first local floras (the word flora, however, was first introduced by Linnaeus in the next century), and of these Germany especially soon produced a considerable number; a flora of Altorf was published by Ludwig Jungermann in 1615, of Ingolstadt by Albert Menzel in 1618, of Giessen by Jungermann in 1623, of Dantzic by Nicolaus Oelhafen in 1643, of Halle by Carl Scheffer in 1662, of the Palatinate by Frank von Frankenau in 1680, of Leipsic by Paul Ammann in 1675, of Nuremberg by J. Z. Volkamer in 1700.
But though travel, catalogues in local floras, and the cultivation of plants in botanic gardens promote knowledge of very varied kind, yet this remains scattered about among descriptions of plants, until at last a writer with powers of combination and wider and deeper glance endeavours to gain some general conclusions from them. Such attempts we first meet with late in the second half of the 17th century in Morison, Ray, Bachmann (Rivinus), Tournefort, and others, who took up Cesalpino’s principles after they had lain neglected for almost a hundred years, and indeed were almost forgotten by botanists.
In the dearth of higher scientific efforts during this period, the describing of plants and cataloguing of species prolonged a somewhat pitiful existence. This describing, a work of great usefulness in the fathers of German botany, was now become by perpetual repetition a mechanical labour; all that was to be gained in this way had already been gained by de l’Obel and Bauhin. This sterility which followed upon the fruitful beginnings of the 16th century was general; neither in Germany nor Italy, neither in France nor England, did the botanists produce anything of importance. The representatives of the science did not count among the more highly gifted or among the thinkers of their time; and so content with the minor work of collecting and cataloguing plants, and with endeavouring to know all plants as far as possible by name, they lost whatever capacity they may have possessed for more difficult operations of the mind simply by not attempting them.
There was one man indeed in Germany who studied the vegetable kingdom in the first half of the 17th century in the spirit of Cesalpino before him, but who, like Cesalpino, found no honour among contemporary botanists. This man was the well-known philosopher Joachim Jung, who invented a comparative terminology for the parts of plants, and occupied himself with critical enquiries into the theory of the system, the naming of species and other subjects, embodying their results in a long array of aphorisms. Free from the genius-stifling burden which the knowledge of individual species had become, a man possessed of varied accomplishments and a well-trained mind, Jung was better qualified than the professed botanists to see what was wanted in botany and would advance it—a phenomenon more than once repeated in the history of the science. But his results remained unknown to all except his immediate pupils, till Ray admitted them into his great work on plants in 1693, and made them the foundation of his own theoretical botany. Enriched by Ray’s good morphological remarks, Jung’s terminology passed to Linnaeus, who adopted it as he adopted every thing useful that literature offered him, improving it here and there, but impairing its spirit by his dry systematising manner.
The labours of the botanists of Germany and the Netherlands during the 17th century, which culminated in Kaspar Bauhin, were not without important influence upon the development of systematic botany which began with Cesalpino. When Cesalpino wrote the work which forms an epoch in the science, he was perhaps unacquainted with the natural classification of de l’Obel (1576); at least there is nothing in his book which shows that he had seen it; it appears even as though he had made the discovery independently, that there is an actual connection of relationship among plants expressed in their organisation as a whole; it is at any rate certain that this fact assumed from the first an entirely different expression in his system from that which it received at the hands of de l’Obel and Bauhin, inasmuch as he was not guided by an indistinct feeling for resemblances, but believed that he could establish on predetermined grounds a system of marks, by which the objective relationship must be recognised. If Cesalpino was thus in advance of the German botanists, since he endeavoured to express with clearness and on principle that which they only felt indistinctly, he was at the same time treading a dangerous path, and one which led succeeding botanists astray till the time of Linnaeus,—the path which must always lead to artificial classifications, since the natural system can never be laid down upon a priori principles of division. Through this labyrinth, in which botanists down to Linnaeus wandered fruitlessly hither and thither, there remained one guide consistently pointing to the goal to be attained, namely, the feeling for natural affinity first vividly apprehended by the German botanists, and expressed by them to some extent in their classifications. And when at last Linnaeus and Bernard de Jussieu made the first feeble attempts at a natural arrangement, it was the same indistinct perception which asserted itself in them as in de l’Obel and Bauhin, and enabled them to see that the path hitherto trodden could only lead astray.
The period in the development of descriptive botany which begins with Cesalpino and reaches to Linnaeus may accordingly be perhaps best characterised by saying, that botanists sought to do justice to natural affinities by means of artificial classifications, till at length Linnaeus clearly perceived the contradiction involved in this method of proceeding. But inasmuch as Linnaeus left it to the future to work out the natural system, and arranged the plants which he described in a confessedly artificial manner, he so far marks rather the close of a previous condition of the science than the beginning of modern botany.
These introductory observations will have supplied the reader with the thread which will guide him through the following account of the more prominent points in the history of botanical science from Cesalpino to Linnaeus.
The often-quoted work of Andrea Cesalpino[17], ‘De plantis libri XVI,’ appeared in Florence in the year 1583. If the value of the contemporary German botanists lies pre-eminently in the accumulation of descriptions of individual plants, and these, it is true, occupy fifteen books of Cesalpino’s work, it is on the contrary the introduction in the first book, a discussion of the general theory of the subject, which in his case is of much the higher importance for the history of botany. This contains in thirty pages a full and connected exposition of the whole of theoretical botany, and though based on broad and general views is at the same time extremely rich in matter conveyed in a very concise form. The different branches into which the subject has since been divided are here united into an inseparable whole; morphology, anatomy, biology, physiology, systematic botany, terminology are so closely combined, that it is difficult to explain Cesalpino’s views on any one more general question without at the same time touching on a variety of other matters. Three things more especially characterise this introductory book; first, a great number of new and delicate observations; secondly, the great importance which Cesalpino assigns to the organs of fructification as objects of morphological investigation; lastly, the way in which he philosophises in strictly Aristotelian fashion on the material thus gained from experience. If this treatment has produced a work beautiful in style and fascinating to the reader, if the whole subject is vivified by it while each separate fact gains a more general value, it is on the other hand apparent that the writer is often led astray by the well-known elements of the Aristotelian philosophy, which are opposed to the interests of scientific investigation. Mere creations of thought, the abstractions of the understanding, are treated as really existent substances, as active forces, under the name of principles; final causes appear side by side with efficient; the organs and functions of the organism exist either alicujus gratiâ or merely ob necessitatem; the whole account is controlled by a teleology, the influence of which is the more pernicious because the purposes assumed are supposed to be acknowledged and self-evident, plants and vegetation being conceived of as in every respect an imperfect imitation of the animal kingdom. It was moreover a necessary consequence of the treatment of his material adopted by Cesalpino, that his ignorance of the sexuality of plants and of the use of leaves as organs of nutrition led him to false and mischievous conclusions; this defect of knowledge would have been of less importance in a purely morphological consideration of plants, as we shall see presently in Jung; but with Cesalpino morphological and physiological considerations are so mixed up together, that a mistake in the one direction necessarily involved mistakes in the other.
These remarks on Cesalpino’s method may be illustrated by some examples tending to show how closely he attaches himself to Aristotle, and how certain Aristotelian conceptions, the origin of which has not been sufficiently regarded, passed through him into later botanical speculation. We shall recur in the History of Physiology to Cesalpino’s views on nutrition, and to his rejection of the doctrine of sexuality in plants.
‘As the nature of plants,’ so begins Cesalpino’s book, ‘possesses only that kind of soul by which they are nourished, grow, and produce their like, and they are therefore without sensation and motion in which the nature of animals consists, plants have accordingly need of a much smaller apparatus of organs than animals.’ This idea reappears again and again in the history of botany, and the anatomists and physiologists of the 18th century were never weary of dilating on the simplicity of the structure of plants and of the functions of their organs. ‘But since,’ continues Cesalpino, ‘the function of the nutritive soul consists in producing something like itself, and this like has its origin in the food for maintaining the life of the individual, or in the seed for continuing the species, perfect plants have at most two parts, which are however of the highest necessity; one part called the root by which they procure food; the other by which they bear the fruit, a kind of foetus for the continuation of the species; and this part is named the stem (‘caulis’) in smaller plants, the trunk (‘caudex’) in trees.‘
This in the main correct conception of the upright stem as the seed-bearer of the plant was also long maintained in botany. We should observe also that the production of the seed is spoken of as merely another kind of nutrition, a notion which afterwards prevented Malpighi from correctly explaining the flower and fruit, and in a modified form led Kaspar Friedrich Wolff in 1759 to a very wrong conception of the nature of the sexual function. The next sentence in Cesalpino takes us into the heart of the Aristotelian misinterpretation of the plant, according to which the root answers to the mouth or stomach, and must therefore be regarded in idea as the upper part although it is the lower in position, and the plant would have to be compared with an animal set on its head, and the upper and lower parts determined accordingly: ‘this part (the root) is the nobler (‘superior’) because it is prior in origin and sunk in the ground; for many plants live by the roots only after the stem with the ripe seeds has disappeared; the stem is of less importance (‘inferior’) although it rises above the ground; for the excreta, if there are any, are given off by means of this part; it is, therefore, with plants as with animals as regards the expressions ‘pars superior’ and ‘inferior.’ When indeed we take into consideration the mode of nourishment, we must define the upper and the lower in another way; since in plants and animals the food mounts upward (for that which nourishes is light because it is carried upwards by the heat), it was necessary to place the roots below and to make the stem go straight upwards, for in animals also the veins are rooted in the lower part of the stomach, while their main trunk ascends to the heart and the head.’ Here, in genuine Aristotelian fashion, the facts are forced into a previously constructed scheme.
Cesalpino’s discussion of the seat of the soul in plants is of special interest in connection with certain views of later botanists. ‘Whether any one part in plants can be assigned as the seat of the soul, such as the heart in animals, is a matter for consideration—for since the soul is the active principle (‘actus’) of the organic body, it can neither be ‘tota in toto’ nor ‘tota in singulis partibus,’ but entirely in some one and chief part, from which life is distributed to the other dependent parts. If the function of the root is to draw food from the earth, and of the stem to bear the seeds, and the two cannot exchange functions, so that the root should bear seeds and the shoot penetrate into the earth, there must either be two souls different in kind and separate in place, the one residing in the root, the other in the shoot, or there must be only one, which supplies both with their peculiar capabilities. But that there are not two souls of different kinds and in a different part in each plant may be argued thus; we often see a root cut off from a plant send forth a shoot, and in like manner a branch cut off send a root into the ground, as though there were a soul indivisible in its kind present in both parts. But this would seem to show that the whole soul is present in both parts, and that it is wholly in the whole plant, if there were not this objection that, as we find in many cases, the capabilities are distributed between the two parts in such a way that the shoot, though buried in the ground, never sends out roots, for example in Pinus and Abies, in which plants also the roots that are cut off perish.’ This, he thinks, proves that there is only one soul residing in root and stem, but that it is not present in all the parts; in a further discussion he seeks to discover the true seat of the soul. He points out an anatomical distinction between the shoot and the root; the root consists of the rind and an inner substance which in some cases is hard and woody, in others soft and fleshy. In the stem on the other hand there are three constituent parts; outside the rind, inside the pith, between the two a body which in trees is called the wood. This, on the whole, correct distinction between stem and root is followed by a thoroughly Aristotelian deduction.
‘Since then in all creatures’ (we must remark, that this is assuming a point which has yet to be proved in the case of the half of living creatures) ‘nature conceals the principle of life in the innermost parts, as the entrails in animals, it is reasonable to conclude that the principle of life in plants is not in the rind, but is more deeply hidden in the inner parts, that is, in the pith, which is found in the stem and not in the root. That this was the opinion of the ancients we may gather from the name, for they called this part in plants the heart (‘cor’), or brain (‘cerebrum’ or ‘matrix’), because from this part in some degree the principle of foetification (the formation of the seed) is derived.’ Here we see why the seed must, according to Cesalpino, have its origin in the pith; the idea was loyally repeated after him by Linnaeus, as we shall see hereafter. The argument, which is a long one, ends with the sentence: ‘There are then two chief parts in plants, the root and the ascending part; therefore the most suitable spot for the heart of plants seems to be in the central part, namely, where the shoot joins on to the root. There appears also at this spot a certain substance differing both from the shoot and from the root, softer and more fleshy than either, for which reason it is usually called the cerebrum; it is edible in many plants while they are young.’ We shall see below how important a part this seat of the soul of the plant, brought to light with such difficulty and with all appliances of scholasticism, is intended to play in Cesalpino’s system, and how by this a priori path he was led to the use of the position of the embryo in the seed as his principle of division. It may be remarked here that the point of union between the root and the stem, in which Cesalpino placed the seat of the plant-soul, afterwards received the name of root-neck (collet); and though the Linnaean botanists of the 19th century were unaware of what Cesalpino had proved in the 16th, and did not even believe in a soul of plants, they still entertained a superstitious respect for this part of the plant, which is really no part at all; and this, it would seem, explains the fact, that an importance scarcely intelligible without reference to history was once attributed to it, especially by some French botanists. To return once more to Cesalpino’s ‘cor,’ he is not much troubled by the circumstance that plants can be reproduced from severed portions; in true Aristotelian manner he says that although the principle of life is actually only one, yet potentially it is manifold. Ultimately a ‘cor’ is found in the axil of every leaf, by which the axillary shoot is united with the pith of the mother-shoot, and finally, in direct contradiction to the previous proof that the crown of the root is the seat of the plant-soul, it is distinctly affirmed in Chapter V that the soul of plants is in some sense diffused through all their parts.
The theoretical introduction to his excellent and copious remarks on the parts of fructification may supply another example of Cesalpino’s peripatetic method: ‘As the final cause (‘finis’) of plants consists in that propagation which is effected by the seed, while propagation from a shoot is of a more imperfect nature, in so far as plants do exist in a divided state, so the beauty of plants is best shown in the production of seed; for in the number of the parts, and the forms and varieties of the seed-vessels, the fructification shows a much greater amount of adornment than the unfolding of a shoot; this wonderful beauty proves the delight (‘delitias’) of generating nature in the bringing forth of seeds. Consequently as in animals the seed is an excretion of the most highly refined food-substance in the heart, by the vital warmth and spirit of which it is made fruitful, so also in plants it is necessary that the substance of the seeds should be secreted from the part in which the principle of the natural heat lies, and this part is the pith. For this reason, therefore, the pith of the seed (that is, the substance of the cotyledons and of the endosperm) springs from the moister and purer part of the food, while the husk which surrounds the seed for protection springs from the coarser part. It was unnecessary to separate a special fertilising substance from the rest of the matter in plants, as it is separated in animals which are thus distinguished as male and female.‘
This last remark and some lengthy deductions which follow are intended to prove, after the example of Aristotle, the absence and indeed the impossibility of sexuality in plants, and accordingly Cesalpino goes on to compare the parts of the flower, which he knew better than his contemporaries, with the envelopes of the ova in the foetus of animals, which he regards as organs of protection. Calyx, corolla, stamens, and carpels are in his view only protecting envelopes of the young seed, as the leaves are only a means of protecting the young shoots. Moreover by the word flower (‘flos’) Cesalpino understands only those parts of the flower which do not directly belong to the rudiment of the fruit, namely, the calyx, the corolla, and the stamens. This must be borne in mind if we would understand his theory of fructification, and especially his doctrine of metamorphosis. We must also note, that by the expression pericarp he understands exclusively juicy edible fruit-envelopes, though at the same time pulpy seed-envelopes inside the fruit pass with him for pericarps. The parts of his flower are the ‘folium,’ which evidently means the corolla, but in certain cases includes also the calyx; the ‘stamen,’ which is our style; and the ‘flocci,’ our stamens. We see that Cesalpino uses the same word ‘folium’ without distinction for calyx, corolla, and ordinary leaves; just as he, and Malpighi a hundred years later, unhesitatingly regarded the cotyledons as metamorphosed leaves. In fact the envelopes of the flower and the cotyledons approach so nearly to the character of leaves, that every unprejudiced eye must instinctively perceive the resemblance; and if doubts arose on this point in post-Linnaean times, it was only a consequence of the Linnaean terminology, which neglected all comparative examination.
Moreover the doctrine of metamorphosis appears in a more consistent and necessary form in Cesalpino than in the botanists of the 19th century before Darwin; it flows more immediately from his philosophical views on the nature of plants, and appears therefore up to a certain point thoroughly intelligible. We may also consider as part of this doctrine in Cesalpino the view that the substance of the seed (embryo and endosperm) arises from the pith, because the pith contains the vital principle[18], and as the pith in the shoot is surrounded for protection by the wood and the bark, so the substance of the seed is surrounded by the woody shell, and by the bark-like pericarp or by a fruit-envelope answering to a pericarp. According to Cesalpino therefore the substance of the seed with its capability of development springs from the pith, the woody shell from the wood, the pericarp from the rind of the shoot. The difficulty which arises from this interpretation, namely, that in accordance with his theory the parts of the flower also, the calyx, the corolla, and the stamens ought to spring from the outer tissues of the shoot, he puts aside with the remark (p. 19) that these parts of the flower are formed when the pericarp is still in a rudimentary state; that the pericarp is only fully developed after these parts have fallen off, and that they are so thin that there is nothing surprising in this view of the matter. We see in Cesalpino’s doctrine of metamorphosis without doubt the theory of the flower afterwards adopted by Linnaeus, though in a somewhat different form. That Linnaeus himself regarded the theory ascribed to him on the nature of the flower as the opinion of Cesalpino also, is shown in his ‘Classes Plantarum,’ where in describing Cesalpino’s system he says: ‘He regarded the flower as the interior portions of the plant, which emerge from the bursting rind; the calyx as a thicker portion of the rind of the shoot; the corolla as an inner and thinner rind; the stamens as the interior fibres of the wood, and the pistil as the pith of the plant.’ It may be observed however that this was not exactly what Cesalpino says; but it is nevertheless certain that Linnaeus’ own view as given in these words was intended to reproduce that of Cesalpino; and if it does not do this exactly, there is no essential difference in principle between the two, Linnaeus’ conception being perhaps a more logical statement of Cesalpino’s meaning. Cesalpino’s doctrine of metamorphosis appears plainly on another occasion also; he says, that we do not find envelopes, stamens, and styles in all flowers; the flowers change in some cases into another substance, as in the hazel, the edible chestnut, and all plants that bear catkins; the catkin is in place of a flower, and is a longish body arising from the seat of the fruit, and in this way fruits appear without flowers, for the styles (‘stamina’) form the longer axis of the catkin (‘in amenti longitudinem transeunt’), while the leafy parts and the stamens are changed into its scales. All this shows that the notion of a metamorphosis, of which we find intimations as early as Theophrastus, was a familiar one to Cesalpino, and it fitted in perfectly with his Aristotelian philosophy, while Goethe’s doctrine on the same subject is equally scholastic in its character, and therefore looks strange and foreign in modern science. It has already been observed that Cesalpino includes only the envelopes and stamens under the word flower, and distinguishes the rudiments of the fruit from them; therefore he says that there are plants which produce something in the shape of a catkin, without any hope of fruit, for they are entirely unfruitful; but those which bear fruit have no flowers, as Oxycedrus, Taxus, and among herbs Mercurialis, Urtica, Cannabis, in which the sterile plants are termed male, the fruitful female. Thus he distinguished the cases which we now call dioecious from the previously mentioned monoecious plants, among which he reckons the maize.
All this may serve to give the reader some idea, though a very incomplete one, of Cesalpino’s theory; to do him justice, it would be necessary to give a full account of his very numerous, accurate, and often acute observations on the position of leaves, the formation of fruit, the distribution of seeds and their position in the fruit, of his comparative observations on the parts of the fruit in different plants, and above all of his very excellent description of plants with tendrils and climbing plants, of those that are armed with thorns and the like. Though there is naturally much that is erroneous and inexact in his accounts, yet we have before us in the chapters on these subjects the first beginning of a comparative morphology, which quite casts into the shade all that Aristotle and Theophrastus have said on the subject. But the most brilliant portions of his general botany are contained in the 12th, 13th, and 14th chapters, in which he gives the outlines of his views on the systematic arrangement of plants; to prepare the way for what is to follow, he shows first that it is better to give up the four old divisions of the vegetable kingdom, and to unite the shrubs with the trees and the undershrubs with the herbs. But how these genera are to be distinguished into species is, he says, hard to conceive, for the multitude of plants is almost innumerable; there must be many intermediate genera containing the ‘ultimae species,’ but few are as yet known. He then turns to the divisions founded on the relations of plants to men. Such groups, he says, as vegetables and kinds of grain, which are put together under the name of ‘fruges’ and kitchen-herbs (‘olera’), are formed more from the use made of them than from the resemblance of form, which we require; and he shows this by good examples. The discerning of plants, he continues, is very difficult, for so long as the genera (larger groups) are undetermined, the species must necessarily be mixed up together[19]; the difficulty arises from our uncertainty as to the rules by which we should determine the resemblances of the genera. While there are two chief parts in plants, the root and the shoot, we cannot, as it seems, determine the genera and species from the likeness or unlikeness either of the one or of the other; for if we make a genus of those plants which have a round root, as the turnip, Aristolochia, Cyclamen, Arum, we separate generically things which agree together in a high degree, as rape and radish which agree with the turnip, and the long Aristolochia which agrees with the round, while at the same time we unite things most dissimilar, for the Cyclamen and the turnip are in every other respect of a quite different nature; the same is the case with divisions which rest merely on differences in the leaves and flowers.
In pursuing these reflections, which have the conception of species chiefly in view, he arrives at the following proposition: That according to the law of nature like always produces like, and that which is of the same species with itself.
All that Cesalpino says on systematic arrangement shows that he was perfectly clear in his own mind with regard to the distinction between a division on subjective grounds, and one that respects the inner nature of plants themselves, and that he accepted the latter as the only true one. He says, for instance, in the next chapter: ‘We seek out similarities and dissimilarities of form, in which the essence (‘substantia’) of plants consists, but not of things which are merely accidents of them (‘quae accidunt ipsis’).’ Medicinal virtues and other useful qualities are, he says, just such accidents. Here we see the path opened, along which all scientific arrangement must proceed, if it is to exhibit real natural affinities; but at the same time there is a warning already of the error which beset systematic botany up to Darwin’s time; if in the above sentence we substitute the word idea for that of substance, and the two expressions have much the same meaning in the Aristotelian and Platonic view of nature, we recognise the modern predarwinian doctrine, that species, genera, and families represent ‘ideam quandam’ and ‘quoddam supranaturale.’
Pursuing his deductions, Cesalpino next shows, that the most important divisions, those of woody plants and herbs, must be maintained in accordance with the most important function of vegetation, that of drawing up the food through root and shoot; this division passed from the first and later on up to the time of Jung for an unassailable dogma, to which science simply had to conform. The second great function of plants is the producing their like, and this is effected by the parts of fructification. Though these parts are only found in the more perfect forms, yet the subdivisions (‘posteriora genera’) must be derived in both trees and herbs from likeness and unlikeness in the fructification. And thus Cesalpino was led, not by induction but by the deductive path of pure Aristotelian philosophy, to the conclusion, that the principles of a natural classification are to be drawn from the organs of fructification; for which conclusion Linnaeus declared him to be the first of systematists, while he thought de l’Obel and Kaspar Bauhin, who founded their arrangements on the habit only, scarcely deserving of notice.
It appears, then, that Cesalpino obtained the subdivisions which he founded on the organs of fructification from a priori views of the comparative value of organs, such as run through all Aristotelian philosophy. Of much interesting matter in the remainder of his introduction we must mention only that he makes the highest product of plants to be the fructification, of animals sense and movement, of man the intellect; and because the latter stands in need of no special bodily instruments, there is no specific difference in men, and therefore only one species of man.
In his 14th chapter he gives in broad outline a view of the system of plants which he founded on the fructification, beginning with the least perfect; no one who knows the botanical writers of the 17th and 18th centuries will be surprised to find that Cesalpino admits the doctrine of ‘generatio spontanea’ in the case of the lower plants, and in a somewhat crude form; this came from the teaching of Aristotle, and even a hundred years later Mariotte endeavoured to set up a plausible defence of spontaneous generation on physical grounds even in highly developed plants.
‘Some plants,’ says Cesalpino, ‘have no seed; these are the most imperfect, and spring from decaying substances; they have only therefore to feed themselves and grow, and are unable to produce their like; they are a sort of intermediate existences between plants and inanimate nature. In this respect Fungi resemble Zoophytes, which are intermediate between plants and animals, and of the same nature are the Lemnae, Lichenes, and many plants which grow in the sea.’
Some on the other hand produce seed, which they form after their peculiar nature in an imperfect condition, as the mule among animals; these are of the same nature as mere monstrosities or diseased growths of other plants, and many occur in the class of grain and bear empty ears. Cesalpino is evidently speaking of the Ustilagineae, but he includes also the Orobancheae and Hypocystis, which instead of seed contain only a powder; and he adds that some of the more perfect plants are sterile, but they do not belong to this division, because the peculiarity is confined in their case to individuals.
Some plants bear a substance, a kind of wool, on the leaves, which to some extent answers to seed, because it serves to propagate the plant; such plants have neither stem, flower, nor true seed, and the Ferns are of this kind. We should notice this conclusion from Cesalpino’s morphology, that plants without true seeds have also no stem; the view that ferns have no stems continued to be held by later botanists, though the original reason for it was gradually lost; and those who in the middle of the 19th century argued still in favour of this opinion, little suspected that they were endeavouring to establish a dogma of the Aristotelian philosophy. It is a similar case to that of the crown of the root mentioned above. But other plants, continues Cesalpino, produce true seeds; and he proceeds to treat of this division first, on account of its great extent as comprising all perfect plants. Three things, he says, contribute especially to the constitution of organs, the number, position, and shape of the parts; the play of nature in the composition of fruits varies according to their differences, and hence arise the different divisions of plants. He then shows how he proposes to apply these relations to the framing of his system, but his various points of view may be omitted here, as they can be better and more shortly gathered from the table below.
Other marks to be derived from roots, stems, and leaves, may be used, he says, for forming the smaller divisions. Lastly, some marks which contribute to the constitution neither of the whole plant nor of the fruit, such as colour, smell, taste, are mere accidents and are due to cultivation, place of growth, climate, and other causes.
The first of Cesalpino’s sixteen books ends with this general view of his system. The remaining fifteen books contain about 600 pages of descriptions of individual plants arranged in fifteen classes; some of the descriptions are exceedingly minute; the trees come first, and are followed by the shrubs on account of their affinity (‘ob affinitatem’). Two things have interfered with the recognition and acceptance of this system; the omission of a general view to precede the text, and its appearance in the traditional form of books and chapters, such as we find in de l’Écluse, Dodoens, and Bauhin, instead of in classes and orders, though it is true that the headings and introductions to the several books contain the designations and general characteristics of the classes described in them. Linnaeus has done good service by giving in his ‘Classes Plantarum’ a general view of all the systems proposed before his time, among which he gives the first rank to that of Cesalpino; he has also pointed out the peculiar characteristics of each system, and has appended to the old names of the genera those with which he has himself made us familiar. This invaluable work, which is a key to the understanding of the efforts that were made in systematic botany from Cesalpino to Linnaeus himself, will often be referred to in later pages of this history; it will supply us here with a tabular view of Cesalpino’s main divisions as precisely formulated by Linnaeus, which is well worth the space it will occupy, as presenting the first plan proposed for a systematic arrangement of the vegetable kingdom, with characters for each division. For the better understanding of these diagnoses it should be remembered that the ‘cor’ (heart) is the important point in the seed with Cesalpino, and that it is the place in the embryo where the radicle and the plumule unite, as has been said in a former page; Cesalpino himself says somewhat inexactly, the place from which the cotyledons spring.
The characters of the classes are given, for brevity’s sake, in Latin.
Arboreae
(Arbores et frutices).
I. Corde ex apice seminis. Seminibus saepius solitariis (e.g. Quercus, Fagus, Ulmus, Tilia, Laurus, Prunus).
II. Corde e basi seminis, seminibus pluribus (e.g. Ficus, Cactus, Morus, Rosa, Vitis, Salix, Coniferae, etc.).
Herbaceae
(Suffrutices et herbae).
III. Solitariis seminibus. Semine in fructibus unico (e.g. Valeriana, Daphne, Urtica, Cyperus, Gramineae).
IV. Solitariis pericarpiis. Seminibus in fructu pluribus, quibus est conceptaculum carnosum, bacca aut pomum (e.g. Cucurbitaceae, Solaneae, Asparagus, Ruscus, Arum).
V. Solitariis vasculis. Seminibus in fructu pluribus quibus est conceptaculum e siccâ materiâ (e.g. various Leguminosae, Caryophylleae, Gentianeae, etc.).
VI. Binis seminibus. Semina sub singulo flosculo invicem conjuncta, ut unicum videantur ante maturitatem; cor in parte superiore, quâ flos insidet. Flores in umbellâ (Umbelliferae).
VII. Binis conceptaculis (e.g. Mercurialis, Poterium, Galium, Orobanche, Hyoscyamus, Nicotiana, Cruciferae).
VIII. Triplici principio (ovary) non bulbosae. Semina trifariam distributa; corde infra sito, radix non bulbosa (e.g. Thalictrum, Euphorbia, Convolvulus, Viola).
IX. Triplici principio bulbosae. Semina trifariam distributa; corde infra sito, radix bulbosa (Large-flowered Monocotyledons).
X. Quaternis seminibus. Semina quatuor nuda in communi sede (Boragineae and Labiatae).
XI. Pluribus seminibus, anthemides. Semina nuda plurima, cor seminis interius vergens; flos communis distributus per partes in apicibus singuli seminis (Compositae only).
XII. Pluribus seminibus, cichoraceae aut acanaceae. Semina nuda plurima, cor seminis inferius vergens, flos communis distributus per partes in apicibus singuli seminis (Compositae, Eryngium, and Scabiosa).
XIII. Pluribus seminibus, flore communi. Semina solitaria plurima, corde interius; flos communis, non distributus, inferius circa fructum (e.g. Ranunculus, Alisma, Sanicula, Geranium, Linum).
XIV. Pluribus folliculis. Semina plura in singulo folliculo (e.g. Oxalis, Gossypium, Aristolochia, Capparis, Nymphaea, Veratrum, etc.).
XV. Flore fructuque carentes (Filices, Equiseta, Musci including Corals, Fungi).
The examples appended by me to the diagnoses show that with the exception of the sixth, tenth, and fifteenth classes, no one perfectly represents a natural group of the vegetable kingdom. Most of them are a collection of heterogeneous objects, and the distinction of Dicotyledons and Monocotyledons, almost perfectly carried out by de l’Obel and Bauhin, is to a great extent effaced; the ninth class certainly contains only Monocotyledons, but not all of them. This result of great efforts on the part of a mind so well trained as Cesalpino’s is highly unsatisfactory. Not a single new group founded on natural affinities is established, which does not appear already in the herbals of Germany and the Netherlands. It is characteristic of the natural system to reveal itself to a certain extent more readily to instinctive perception than to the critical understanding. We have seen that Cesalpino intended as far as possible to give expression in his system to natural affinities, and the final result was a series of highly unnatural groups, almost every one of which is a collection of the most heterogeneous forms. The cause of this apparently so remarkable fact is this, that he believed that he could establish on predetermined grounds the marks which indicate natural affinities. The uninterrupted labour of nearly 300 years, starting again and again from the same principle or practically under its influence, has given us inductive proof that the path taken by Cesalpino is the wrong one. And if, while this path was pursued even into the middle of the 18th century, we see natural groups emerge with increasing distinctness, it is because the botanist, though on the wrong track, was still continually gaining better acquaintance with the ground over which he was wandering, and attained at length to an anticipation of the truer way.
Joachim Jung[20] was born in Lübeck in the year 1587, and died after an eventful life in 1657. He was a contemporary of Kepler, Galileo, Vesal, Bacon, Gassendi, and Descartes. After having been already a professor in Giessen, he applied himself to the study of medicine in Rostock, was in Padua in 1618 and 1619, and there, as we may confidently believe, became acquainted with the botanical doctrines of Cesalpino, who had died fifteen years before. Returning to Germany, he held various professorships during the succeeding ten years in Lübeck and Helmstädt, and became Rector of the Johanneum in Hamburg in 1629. He occupied himself with the philosophy of the day, in which he appeared as an opponent of scholasticism and of Aristotle, and also with various branches of science, mathematics, physics, mineralogy, zoology, and botany. In all these subjects he displayed high powers as a student and a teacher, and especially as a critical observer; in botany at least he was a successful investigator. He was the first in Germany, as Cesalpino had been in Italy, who combined a philosophically educated intellect with exact observation of plants.
His pupils were at first the only persons who profited by his botanical studies, for with his many occupations and a perpetual desire to make his investigations more and more complete he himself published nothing. In 1662 his pupil Martin Fogel printed the ‘Doxoscopiae Physicae Minores,’ a work of enormous compass left in manuscript at the master’s death, and another pupil, Johann Vagetius, the ‘Isagoge Phytoscopica,’ in 1678. Ray however tells us that a copy of notes on botanical subjects had already reached England in 1660. The ‘Doxoscopiae’ contains a great number of detached remarks on single plants and on their distinguishing marks, and propositions concerning the methods and principles of botanical research,—all in the form of aphorisms which he had from time to time committed to paper. The number and contents of these aphorisms show the earnest attention which he bestowed on the determination of species; he is displeased that so many botanists devote more time and labour to the discovery of new plants, than to referring them carefully and logically to their true genera by means of their specific differences. He was the first who objected to the traditional division of plants into trees and herbs, as not founded on their true nature. But how firmly this old dogma was established is well shown by the fact, that Ray at the end of the century still retained this division, though he founded his botanical theories on the ‘Isagoge’ of Jung. Jung was in advance of Cesalpino and his own contemporaries in repeatedly expressing his doubt of the existence of spontaneous generation.
The ‘Isagoge Phytoscopica,’ a system of theoretical botany, very concisely written and in the form of propositions arranged in strict logical sequence, was a more important work and had more lasting effects upon the history of botany. We must look more closely into the contents of this volume, because it contains the foundation of the terminology of the parts of plants subsequently established by Linnaeus. Since the matter of the ‘Isagoge’ is produced in Ray’s ‘Historia Plantarum’ in italics, with special mention of the source from which it is derived, it cannot be doubted that Linnaeus had made acquaintance with the teaching of Jung as a young man, in any case before 1738. It is as important as a matter of history to know that Linnaeus’ terminology is founded on Jung, as it is to learn that his most general philosophical propositions on botanical subjects are to be traced to Cesalpino. It will moreover be fully shown in the account of the doctrine of sexuality that his knowledge of that subject was derived from Rudolf Jacob Camerarius.
The first chapter of the ‘Isagoge’ discusses the distinction between plants and animals. A plant is, according to Jung, a living but not a sentient body; or it is a body attached to a fixed spot or a fixed substratum, from which it can obtain immediate nourishment, grow and propagate itself. A plant feeds when it transforms the nourishment which it takes up into the substance of its parts, in order to replace what has been dissipated by its natural heat and interior fire. A plant grows when it adds more substance than has been dissipated, and thus becomes larger and forms new parts. The growth of plants is distinguished from that of animals by the circumstance that their parts are not all growing at the same time, for leaves and shoots cease to grow as soon as they arrive at maturity; but then new leaves, shoots, and flowers are produced. A plant is said to propagate itself when it produces another specifically like itself; this is the idea in its broader acceptation. We see that here, as in Cesalpino, the idea of the species is connected with that of propagation. The second chapter, headed ‘Plantae Partitio,’ treats of the most important morphological relations in the external differentiation of plants; here Jung adheres essentially to Cesalpino’s view, that the whole body in all plants, except the lowest forms, is composed of two chief parts, the root as the organ which takes up the food, and the stem above the ground which bears the fructification. Jung, too, draws attention to the meeting-point of the two parts, Cesalpino’s ‘cor,’ but under the name of ‘fundus plantae.’
The upper part, or a portion of the plant, is either a stem, a leaf, a flower, a fruit, or a structure of secondary importance, such as hairs and thorns. His definition of the stalk and the leaf is noteworthy; the stalk, he says, is that upper part which stretches upwards in such a manner, that a back and front, a right and left side, are not distinguished in it. A leaf is that which is extended from its point of origin in height, or in length and breadth, in such a manner, that the bounding surfaces of the third dimension are different from one another, and therefore the outer and inner surfaces of the leaf are differently organised. The inner side of the leaf, which is also called the upper, is that which looks towards the stem, and is therefore concave or less convex than the other side. One conclusion he draws, which is a striking one for that time, that the compound leaf is taken for a branch by inexperienced or negligent observers, but that it may easily be determined by having an inner and an outer surface, like the simple leaf, and by falling off as a whole in autumn. He calls a plant ‘difformiter foliata,’ whose lower leaves are strikingly different from the upper, an idea which Goethe, in the fragment in Guhrauer, seems to have altogether misunderstood.
In connection with these general definitions, the different forms of the stem and of the ramification, and the varieties of leaves are pointed out and supplied with distinctive names, which are for the most part still in use. The fourth chapter treats of the division of the stem into internodes; if the stem or branch, says Jung, is regarded as a prismatic body, the articulations, that is, the spots where a branch or a leaf-stalk arises, are to be conceived of as cross-sections parallel to the base of the prism. These spots when they are protuberant are called knees or nodes, and that which lies between such spots is an internode.
It is not possible to quote all the many excellent details which follow these definitions; but some notice must be taken of Jung’s theory of the flower, which he gives at some length from the 13th to the 27th chapters. It suffers, as in Cesalpino, from his entire ignorance of the difference of sexes in plants, which is sufficient to render any satisfactory definition of the idea of a flower impossible. Like Cesalpino too he distinguishes the pistil from the flower, instead of making it a part of the flower. He regards the flower as a more delicate part of the plant, distinguished by colour or form, or by both, and connected with the young pistil. Like all botanists up to the end of the 18th century, he follows Cesalpino in including under the term fruit both the dry indehiscent fruits which were supposed to be naked seeds, and any seed-vessel. He differs from him in calling the stamens ‘stamina,’ and the style ‘stilus,’ but like Cesalpino he uses the word ‘folium’ for the corolla. He calls a flower perfect only when it has all these three parts. He afterwards describes the relations of form and number in the parts of the flower, and among other things he enunciates the first correct view of the nature of the capitulum in the Compositae, which Cesalpino quite misunderstood; and he examined inflorescences and superior and inferior flowers, which Cesalpino had already distinguished, with more care than they had previously received. In his theory of the seed he follows Cesalpino, and adds nothing to him.
There is nothing which more essentially distinguishes the theoretical botany of Jung, and marks the advance which he made upon Cesalpino’s views, than the way in which he discusses morphology in as entire independence as was possible of all physiological questions, and therefore abstains from teleological explanations. His eye is fixed on relations of form only, while his mode of treating them is essentially comparative, and embraces the whole of the vegetable kingdom that was known to him. Jung certainly learnt much from Cesalpino; but in rejecting at least the grosser aberrations of the Aristotelian philosophy and of scholasticism, he freed himself from the prepossessions of his master, and succeeded in arriving at more correct conceptions of the morphology of plants. That his mathematical gifts assisted him in this respect is easy to be gathered from his definitions as given above, which bring into relief the symmetry apparent in the forms of stems and leaves. No more profound or apt definitions were supplied till Schleiden and Nägeli introduced the history of development into the study of morphology.
While Cesalpino, Kaspar Bauhin, and Jung stand as solitary forms each in his own generation, the last thirty years of the 17th century are marked by the stirring activity of a number of contemporary botanists. While during this period physics were making rapid advances in the hands of Newton, philosophy in those of Locke and Leibnitz, and the anatomy and physiology of plants by the labours of Malpighi and Grew, systematic botany was also being developed, though by no means to the same extent or with equally profound results, by Morison, Ray, Bachmann (Rivinus), and Tournefort. The works of these men and of their less gifted adherents, following rapidly upon or partly synchronous with each other, led to an exchange of opinions and sometimes to polemical discussion, such as had not before arisen on botanical subjects; this abundance of literature, with the increased animation of its style, excited a more permanent interest, which spread beyond the narrow circle of the professional adepts. The systematists above-named endeavoured to perfect the morphology and the terminology of the parts of plants, and they found ready to their hands in the works of their predecessors a considerable store of observations and ideas, upon which they set themselves to work. A very great number of descriptions of individual plants had been accumulated since the time of Fuchs and Bock, and the fact of natural affinity had been recognised in the ‘Pinax’ of Kaspar Bauhin as the foundation of a natural system; Cesalpino had pointed to the organs of fructification as the most important for such a system, and Jung had supplied the first steps to a comparative morphology in place of a mere explanation of names. The botanists of the last thirty years of the 17th century could not fail to perceive that the series of affinities as arranged by de l’Obel and Bauhin could not be defined by predetermined marks in the way pursued by Cesalpino, nor fashioned in this way into a well-articulated system. Nevertheless they held fast in principle to Cesalpino’s mode of proceeding, though they endeavoured to amend it by obtaining their grounds of division, not as he had done, chiefly from the organisation of the seed and fruit, but from other parts of the flower; variations in the corolla, the calyx, and the general habit were employed to found systems, which were intended to exhibit natural affinities. And while the true means were thus missed, the end itself was not clearly and decidedly adhered to; a system was desired for the purpose of facilitating the acquisition of a knowledge of the greatest possible number of individual forms; the weight of the burden caused by the foolish demand that every botanist should know all described plants, was continually increasing, and naturally led to seeking some alleviation in systematic arrangement. Excessive devotion to the describing of plants stood in the way of such a profound study of the principles of systematic botany as might have led to enduring results, and even destroyed the very capacity for those difficult intellectual operations, which were absolutely necessary to build up a truly natural system on scientific foundations; the wood could not be seen for the trees. Above all the morphology founded by Jung, though acknowledged and employed, was not sufficiently developed by the labours of others to form the foundation of the system in its grander features,—a reproach which must be made against the systematists of the succeeding hundred years with few exceptions. How could the botanists of the 17th century succeed in acquiring a true conception of the larger groups indicated by natural affinity, when they still held to the old division into trees and herbs, which Jung had already set aside and which is opposed to all consistent morphology, and when they paid so little attention to the structure of the seed and the fruit, that they commonly treated dry indehiscent fruits as naked seeds, and were guilty of other and similar mistakes? But if nothing new and good in principle found its way into systematic botany, much service was rendered to it in matters of detail. The working out of various systems helped to show what marks are not admissible in fixing the limits of the natural groups; the contradiction between the method and aim of the systematists became in this empirical way continually more apparent, till at length Linnaeus was able to recognise it distinctly; and this was beyond doubt a great gain.
To attempt to give an account of all the systematists of England, France, Italy, Germany, and the Netherlands during this period would serve only to obscure the subject; all that is historically important will be brought out more clearly by mentioning those only who have really enriched systematic botany. Whoever wishes for a more complete knowledge of all the systems which made their appearance before Linnaeus will find a masterly account of them in his ‘Classes Plantarum,’ and another worth consulting in Michel Adanson’s ‘Histoire de la Botanique’ (Paris, 1864). It is sufficient for our present purpose to consider more particularly the labours of the four men whose names have recently been mentioned.
Robert Morison[21], who was born in Aberdeen in 1620 and died in London in 1683, was the first after Cesalpino and Bauhin who devoted himself to systematic botany, that is, to founding and perfecting the classification of plants. He was reproached by his contemporaries and successors with having borrowed without acknowledgment from Cesalpino; this was an exaggeration. Morison commenced his efforts as a systematist with a careful examination of Kaspar Bauhin’s ‘Pinax’; there he obtained his conceptions of natural relationship in plants; and if he afterwards founded his own system more peculiarly on the forms of the fruit, it was in a very different way from that adopted by Cesalpino. Linnaeus answers the reproach above-mentioned by the pertinent remark, that Morison departs as far from Cesalpino in this point as he is inferior to him in the purity of his method. In the year 1669 appeared a work with the characteristic title, ‘Hallucinationes Kaspari Bauhini in Pinace tum in digerendis quam denominandis plantis,’ which Haller justly calls an ‘invidiosum opus’; for as there are writers at all times who ungratefully accept all that is good and weighty in their predecessors as self-evident, while they point with malicious pleasure to every little mistake which the originator of a great idea may commit, so Morison has no word of recognition for the great and obvious merits of the ‘Pinax,’ though such a recognition was specially due from one whose design was to point out the numerous mistakes in that work on the subject of affinities. Kurt Sprengel in his ‘Geschichte,’ ii. p. 30, also suspects with reason that Jung’s manuscript, which was communicated by Hartlieb to Ray in 1661, was not unknown to Morison, and in this paper he might certainly have found much that suited his purposes. Sprengel says well, that the ‘Hallucinationes’ are a well-grounded criticism of the arrangement of plants, which the Bauhins had chosen; that the writer goes through the ‘Pinax’ page by page, and shows what plants occupy a false position, and that it is certain that Morison laid the first foundation of a better arrangement and a more correct discrimination of genera and species.
His ‘Plantarum umbelliferarum distributio nova,’ Oxford, 1672, shows considerable advance; it is the first monograph which was intended to carry out systematic principles strictly within the limits of a single large family. The very complex arrangement is founded exclusively on the external form of the fruit, which he naturally terms the seed. It is the first work in which the system is no longer veiled by the old arrangement in books and chapters, perspicuity being provided for by typographical management,—an improvement which de l’Obel, it is true, made a feeble attempt to introduce a hundred years before. Morison also endeavours to give a clear idea of the systematic relations within the family by the aid of linear arrangement, to some extent the first hint of what we now call a genealogical tree, and a proof at any rate of the lively conception which he had formed of affinity, not drawn indeed only ‘ex libro naturae,’ as the title of his book states, but in principle from Bauhin. Morison’s inability to appreciate the merits of his predecessors, and to believe that when he made a step in advance the way had ever been trodden before, may be seen in this work also. One of its merits is, that it contains for the first time careful representations of separate parts of plants, executed in copper plate[22]. In 1680 appeared the first volumes of his ‘Historia plantarum universalis Oxoniensis,’ the third portion of which was published after his death by Bobart in 1699,—a collection of most of the plants then known and a large number of new ones with descriptions; the systematic arrangement in this work is to be seen in Linnaeus’ ‘Classes Plantarum.’ If Morison in his criticism of Bauhin displayed considerable acuteness within narrow circles of affinity, his universal system on the contrary shows extremely small feeling for affinities on the large scale; the most different forms are brought together even in the smaller divisions; the last class of his Bacciferae, for example, contains genera like Solanum, Paris, Podophyllum, Sambucus, Convallaria, Cyclamen, a result which is the more surprising as Morison does not, like Cesalpino, confine himself to single fixed marks, but has regard also to the habit. On the whole his arrangement as an expression of natural affinities must be ranked after those of de l’Obel and Bauhin.
Morison’s merit lay in truth less in the quality of what he did, than in the fact that he was the first to renew the cultivation of systematic botany on a comprehensive scale. The number of his adherents was always small; in Germany Paul Ammann, Professor in Leipsic, adopted Morison’s views in his ‘Character Plantarum Naturalis’ (1685), and Paul Hermann, Professor in Leyden from 1679 to 1695, after collecting plants in Ceylon for eight years, proposed a system founded on that of Morison, but which can scarcely be called an improvement upon it.
In contrast to Morison, John Ray[23] (1628 to 1705) not only knew how to adopt all that was good and true in the works of his predecessors, and to criticise and complete them from his own observations, but could also joyfully acknowledge the services of others, and combine their results and his own into a harmonious whole. He wrote many botanical works; but none display his character as a man and a naturalist better than his comprehensive ‘Historia Plantarum,’ published in three large folio volumes without plates in the period from 1686 to 1704. This work contains a series of descriptions of all plants then known; but the first volume commences with a general account of the science in fifty-eight pages, which, printed in ordinary size, would itself make a small volume, and which treats of the whole of theoretic botany in the style of a modern text-book. If morphology, anatomy, and physiology, in which latter subject he relies on the authority of Malpighi and Grew, are not kept strictly apart in his exposition, yet it is easy to separate the morphological part, and his theory of systematic botany is in fact given separately. Jung’s definitions of the subject-matter of each of the chapters on morphology are first given, and Ray then adds his own remarks, in which he criticises, expands, and supplements those of his predecessor. Omitting all that is not his own, and the anatomical and physiological portions, we will describe some of the more important results of his studies on system. First and foremost Ray adopted the idea which Grew had conceived, but in a very clumsy form, that difference of sex prevails in the vegetable kingdom, and hence the flower had a different meaning and importance for him from what it had had for his predecessors, though his views on the subject were still indistinct. Ray perceived more clearly than Cesalpino that many seeds contain not only an embryo but also a substance, which he calls ‘pulpa’ or ‘medulla,’ and which is now known as the endosperm, and that the embryo has not always two cotyledons, but sometimes only one or none; and though he was not quite clear as regards the distinction, which we now express by the words dicotyledonous and monocotyledonous embryo, yet he may claim the great merit of having founded the natural system in part upon this difference in the formation of the embryo. He displays more conspicuously than any systematist before Jussieu the power of perceiving the larger groups of relationship in the vegetable kingdom, and of defining them by certain marks; these marks moreover he determines not on a priori grounds, but from acknowledged affinities; but it is only in the great divisions of his system that he is thus true to the right course; in the details he commits many and grievous offences against his own method, as we shall see below when we come to an enumeration of his classes. Modern writers have often attributed to Ray the merit of having first taught the transmutation of species, and of being thus one of the founders of the theory of descent. Let us see how much truth there is in this assertion. Though plants, says Ray, which spring from the same seed and produce their species again through seed, belong to the same species, yet cases may occur in which the specific character is not perpetual and infallible. Seeds may sometimes degenerate and produce plants specifically distinct from the mother-plant, though this may not often happen, and so there would be a transmutation of species, as experience teaches. It is true that he considered the statements of various writers, that Triticum may change into Lolium, Sisymbrium into Mentha, Zea into Triticum, etc., to be very doubtful, yet there were, he thought, other cases which were well ascertained; it was in evidence in a court of law that a gardener in London had sold cauliflower seed which had produced only common cabbage. It is to be observed, he says, that such transmutations only occur between nearly allied species and such as belong to the same genus, and some perhaps would not allow that such plants are specifically distinct. These words, especially when judged by Ray’s general views, appear only to express the opinion that certain inconsiderable variations are possible within a narrow circle of affinity, especially in cultivated plants. Ray does not speak of the appearance of new forms, but says that a known form changes into another already existing and known form, which is the reverse of that which the theory of descent requires.
In his development of the principles of his system, among other errors we encounter one that leads to very important consequences in his application of the dictum, ‘natura non facit saltus,’ which he interprets as though all affinities must present themselves in a series that would be represented by a straight line,—an error which has misled systematists even in recent times, and was first recognised as an error by Pyrame de Candolle. Ray overlooked the fact that the dictum holds good even when the affinities arrange themselves in the form of branching series, that is, after the manner of a genealogical tree. Much more sound is his remark, that the framing of the true system had previously been impossible, because the differences and agreements of forms were not sufficiently known; and another saying of his, that nature refuses to be forced into the fetters of a precise system, shows the dawn of the knowledge which afterwards led in Linnaeus to a strict separation of the natural and artificial systems.
It excites no small astonishment after all Ray’s judicious and clear-sighted utterances on the nature and method of the natural system to find him adopting the division into woody plants and herbs; nor is the matter improved by his making the distinctive mark of trees and shrubs to be the forming of buds, that is, distinct winter buds, which is a mistake into the bargain. Yet we feel ourselves in some degree compensated for this serious error by his dividing trees and herbs into those with a two-leaved and those with a one-leaved or leafless embryo, in modern language into Dicotyledons and Monocotyledons. Ray’s system is undoubtedly the one which in the time preceding Linnaeus does most justice to natural affinities. The following synopsis of his Classes will serve to show the progress made since Cesalpino. The names in brackets are the Linnaean names for some of the genera in particular classes.
A. Plantae gemmis carentes (herbae).
(a) Imperfectae.
- I. Plantae submarinae (chiefly Polypes, Fucus).
- II. Fungi.
- III. Musci (Confervae, Mosses, Lycopods).
- IV. Capillares (Ferns, Lemna, Equisetum).
(b) Perfectae.
Dicotyledones (binis cotyledonibus).
- V. Apetalae.
- VI. Planipetalae lactescentes.
- VII. Discoideae semine papposo.
- VIII. Corymbiferae.
- IX. Capitalae (vi-ix are Compositae).
- X. Semine nudo solitario (Valerianeae, Mirabilis, Thesium, etc.).
- XI. Umbelliferae.
- XII. Stellatae.
- XIII. Asperifoliae.
- XIV. Verticillatae (Labiatae).
- XV. Semine nudo polyspermo (Ranunculus, Rosa, Alisma!).
- XVI. Pomiferae (Cucurbitaceae).
- XVII. Bacciferae (Rubus, Smilax, Bryonia, Solanum, Menyanthes).
- XVIII. Multisiliquae (Sedum, Helleboreae, Butomus, Asclepias).
- XIX. Vasculiferae monopetalae (various).
- XX. Vasculiferae dipetalae (various).
- XXI. Tetrapetalae siliquosae (Cruciferae, Ruta, Monotropa).
- XXII. Leguminosae.
- XXIII. Pentapetalae vasculiferae enangiospermae (various).
Monocotyledones (singulis aut nullis cotyledonibus).
- XXIV. Graminifoliae floriferae vasculo tricapsulari (Liliaceae, Orchideae, Zingiberaceae).
- XXV. Stamineae (Grasses).
- XXVI. Anomalae incertae sedis.
B. Plantae gemmiferae (arbores).
(a) Monocotyledones.
- XXVII. Arbores arundinaceae (Palms, Dracaena).
(b) Dicotyledones.
- XXVIII. Arbores fructu a flore remoto seu apetalae (Coniferae and various others).
- XXIX. Arbores fructu umbilicato (various).
- XXX. Arbores fructu non umbilicato (various).
- XXXI. Arbores fructu sicco (various).
- XXXII. Arbores siliquosae (woody Papilionaceae).
- XXXIII. Arbores anomalae (Ficus).
Of these classes only the Fungi, Capillares, Stellatae, Labiatae, Pomiferae, Tetrapetalae, Siliquosae, Leguminosae, Floriferae, and Stamineae can pass as wholly or approximately natural groups, and there are mistakes even in these; moreover the majority of them had long been recognised. The examples annexed in brackets show how open the others are to objection. If it must be allowed on the one side that Ray, like Jung, doubts whether the Cryptogams are propagated without seeds, it is on the other side obvious that he makes as little objection as his predecessors, contemporaries, and immediate successors to the idea that Polypes and Sponges are vegetables. But worse than this is the extremely faulty subordination and coordination in his system; while the class of Mosses contains the Confervae, Lichens, Liverworts, Mosses, and Clubmosses, and therefore objects as distinct from one another as Infusoria, Worms, Crabs, and Mollusks, we find on the contrary the one family of Compositae split up into four classes founded on quite petty and unimportant differences. Finally, if Ray recognised the general importance to the system of the leaf-formation in the embryo, he was still far from strictly separating all Monocotyledons and Dicotyledons.
Ray’s chief merit is that he to some extent recognised natural affinities in their broader features; the systematic separation of the smaller groups was but little advanced by him. He too, like Morison, found two adherents in Germany in the persons of Christopher Knaut (1638-1694), who published a flora of Halle in 1687 arranged after Ray’s method, and Christian Schellhammer (1649-1716), professor at Helmstädt and afterwards at Jena.
Augustus Quirinus Bachmann (Rivinus)[24] (1652-1725) was for Germany what Morison and Ray were for England, and Tournefort for France. From the year 1691 he was Professor of botany, physiology, materia medica, and chemistry in Leipsic; he applied himself with such ardour to astronomy that he injured his eyesight by observing spots in the sun. With such a variety of occupations it is not surprising that his special knowledge of plants was inconsiderable when compared with that of the three just named; but he was better able than they to appreciate the principles of morphology laid down by Jung, and to use them for deciding questions of systematic botany. He did most service by his severe strictures on the more prominent errors which botanists up to his time had persisted in, his own positive contributions, at least as far as the recognition of affinities is concerned, being inconsiderable. His ‘Introductio universalis in rem herbariam,’ which appeared in 1690, and contains 39 pages of the largest size, is the most interesting for us; in it he declines the great quantity of unnecessary work with which botanists occupied themselves, and declares the scientific study of plants to be the only end and aim of botany. He first treats of naming, and lays down with respect to generic and specific names the principles which Linnaeus afterwards consistently applied, whereas Bachmann himself did not follow his own precepts, but injured his reputation as a botanist by a tasteless nomenclature. Nevertheless he declared distinctly that the best plan is to designate each plant by two words, one of which should be the name of the genus, the other that of the species, and he ingeniously pointed out the great convenience of this binary nomenclature in dealing with medicinal plants, and in the writing of prescriptions. He refused to regard cultivated varieties as species, though Tournefort and others continued to do so.
In his system he rejected the division into trees, shrubs, and herbs, showing by good examples that there is no real distinction of the kind in nature. From many of his remarks in his critical dissertations we might infer that he possessed a very fine feeling for natural relationship, but at the same time expressions occur which seem to show that he did not at all appreciate its importance in the system; we notice this in Tournefort also. Because flowers come before the fruit he jumps with curious logic to the conclusion that the main divisions in the system should be derived from the flower, and in following this rule he makes use of exactly that mark in the corolla which has the least value for classification, namely, regularity or irregularity of form. It is strange, moreover, that Bachmann, who spent a considerable fortune on the production of copper-plate figures of plants without any special object, though he founded his system on the form of the flower, should yet have devoted only a superficial study to its construction; his account of it is very inferior to that of any one before or since his time. His classification thus founded cannot be said to be an advance in systematic botany; nevertheless, he had no lack of adherents, and among them in Germany, Heucher, Knaut, Ruppius, Hebenstreit, and Ludwig; in England, Hill and others, who made alterations here and there in his system, but any real development of it was from its nature an impossibility; he endeavoured to defend it against the assaults of Ray and Dillen; Rudbeck also declared against him.
Joseph Pitton de Tournefort[25] (1656-1708) founded his system also on the form of the corolla, but his views are to some extent opposed to those of Bachmann. While the latter was pre-eminently critical and deficient in knowledge of species, Tournefort was more inclined to dogmatise, and atoned in the eyes of his contemporaries for want of morphological insight by his extensive acquaintance with individual plants. He is commonly regarded as the founder of genera in the vegetable kingdom; but it has been already shown that the conceptions of genera and species had been framed as early as the 16th century from the describing of plants, and that Kaspar Bauhin also, in naming his plants, consistently distinguished genera and species; moreover Bachmann in 1690 had supported the claims of the binary nomenclature as the most suitable for the designation of plants, though he did not himself adopt it; Tournefort did adopt it, but in an entirely different way from that of Bauhin. Bauhin gave only the name of the genus, and supplied the species with characters; Tournefort, on the other hand, provided his genera with names and characters, and added the species and varieties without special description. Tournefort therefore was not the first who established genera; he merely transferred the centre of gravity, so to speak, in descriptive botany to the definition of the genera; but in doing so he committed the great fault of treating specific differences within the genus as a matter of secondary importance. How little depth there was in his botanical ideas may be seen not only from his very poor theory of the flower, the imperfections in which, as in the case of Bachmann, are the more remarkable, since he founded his system on the outward form of the flower, but still more from the expression which he uses at the end of his history of botany, a work otherwise of considerable merit; he says there that the science of botany has been so far advanced since the age of Hippocrates, that hardly anything is still wanting except an exact establishing of genera. His general propositions on the subject of systematic botany, together with much that is good, but which is generally not new and is better expressed in the works of Morison, Ray, and Bachmann, contain strange misconceptions; for instance, he classes plants which have no flower and fruit with those in which these parts are to be seen only with the microscope, that is, the smallness of the organs is equivalent to their absence. It may seem strange that his theory of the flower should be so imperfect, when the excellent investigations of Malpighi and Grew into the structure of flowers, fruit, and seed were already before the world (1700), and Rudolph Jacob Camerarius had made known his discovery of sexuality in the vegetable kingdom. This doctrine, however, Tournefort expressly refused to admit. But the reproach of neglecting the labours of Malpighi and Grew is equally applicable to Bachmann and the systematists up to A. L. de Jussieu; we have here only the first example of the fact since so often confirmed, that professed systematists shrank with a certain timidity from the results of more delicate morphological research, and rested their classifications as far as possible on obvious external features in plants,—a proceeding which more than anything else delayed the construction of the natural system.
Tournefort’s system is thoroughly artificial, if possible, more artificial than that of Bachmann, and certainly inferior to Ray’s. If we meet with single groups that are really natural, it is simply because in some families the genera so agree together in all their marks, that they necessarily remain united, whatever mark we select for the systematic purpose. We do not find in Tournefort the distinction between Phanerogams and Cryptogams already established by Ray, nor the division of woody plants and herbs into Monocotyledons and Dicotyledons; if his chief work, to which we confine ourselves here, the ‘Institutiones rei herbariae,’ did not bear the date of 1700, we might conclude that it was written before the ‘Historia Plantarum’ of Ray, and the chief work of Bachmann. Yet it has one merit of a purely formal kind; it is pervaded by a rigorous spirit of system; every class is divided into sections, these into genera, and these again into species; figures of the leaves and of the parts of the flower, very beautifully engraved on copper-plate and filling a whole volume, are perspicuously arranged; the whole work therefore is easy to consult and understand. But to form an idea of the confusion as regards natural affinities that reigns in his system, we need only examine the first three sections of his first class, when we shall find Atropa and Mandragora together in the first section, Polygonatum and Ruscus in the second, Cerinthe, Gentiana, Soldanella, Euphorbia, and Oxalis in the third. The handiness of the book, the little interest taken by most of the botanists of the time in the question of natural relationship, and the continually increasing eagerness for a knowledge of individual plants, are evidently the reasons why Tournefort gained over to his side most of the botanists not only of France, but also of England, Italy, and Germany; and why later attempts in systematic botany during the first thirty or forty years of the 18th century were almost exclusively founded on his system, as they were afterwards on the sexual system of Linnaeus. Boerhaave, among others, proposed a system in 1710, which may be regarded as a combination of those of Ray, Hermann, and Tournefort, but it met with no support on any other grounds.
We here take our leave of the systematists of the 17th century, and, passing over the mere plant-collectors of the first thirty years of the 18th, turn at once to Linnaeus.
Carl Linnaeus[26], called Carl von Linné after 1757, was born in 1707 at Rashult in Sweden, where his father was preacher. He began the study of theology, but was soon drawn away from it by his preference for botany, and in this pursuit he was encouraged by Dr. Rothmann, who sent him to the works of Tournefort. In Lund, where he now studied medicine, he became acquainted with Vaillant’s treatise, ‘De sexu plantarum,’ and had his attention drawn by it to the sexual organs. In 1730, when he was only twenty-three years old, the aged Professor Rudbeck gave up to him his botanical lectures and the management of the botanic gardens, and here Linnaeus began the composition of the ‘Bibliotheca Botanica,’ the ‘Classes Plantarum,’ and the ‘Genera Plantarum.’ In the year 1732 he made a botanical journey to Lapland, and in 1734 to Dalecarlia; in 1735 he went to Holland, where he obtained a degree; in that country he remained three years, and printed the works above-named, together with the ‘Systema Naturae,’ the ‘Fundamenta Botanica,’ and other treatises. From Holland he visited England and France. In the year 1738 he returned to Stockholm and was compelled to gain a livelihood as a physician, till in 1741 he became Professor of Botany in Upsala, where he died in the year 1778.
Linnaeus is commonly regarded as the reformer of the natural sciences which are distinguished by the term descriptive, and it is usual to say that a new epoch in the history of our science begins with him, as a new astronomy began with Copernicus, and new physics with Galileo. This conception of Linnaeus’ historical position, as far at least as his chief subject, botany, is concerned, can only be entertained by one who is not acquainted with the works of Cesalpino, Jung, Ray, and Bachmann, or who disregards the numerous quotations from them in Linnaeus’ theoretical writings. On the contrary, Linnaeus is pre-eminently the last link in the chain of development represented by the above-named writers; the field of view and the ideas of Linnaeus are substantially the same as theirs; he shares with them in the fundamental errors of the time, and indeed essentially contributed to transmit them to the 19th century. But to maintain that Linnaeus marks not the beginning of a new epoch, but the conclusion of an old one, does not at all imply that his labours had no influence upon the time that followed him. Linnaeus stands in the same relation to the systematists of the period we are considering that Kaspar Bauhin does to the botanists of the 16th century; as Bauhin gathered up all that was serviceable in his predecessors, Cesalpino only excepted, while the botanists of our second period drew again from him, though they set out from other points of view than his; so Linnaeus adopted all that the systematists of the 17th century had built upon the foundation of Cesalpino’s ideas, gave it unity and fashioned it into a system without introducing into it anything that was fundamentally and essentially new; all that had been developed in systematic botany from Cesalpino to Tournefort culminated in him, and the results, which he put together in a very original form and with the power of a master, were no more unfruitful for the further development of botany than the contents of Kaspar Bauhin’s works for the successors of Cesalpino.
Whoever carefully compares the works of Cesalpino, Jung, Morison, Ray, Bachmann, and Tournefort with Linnaeus, ‘Fundamenta Botanica’ (1736), his ‘Classes Plantarum’ (1738), and his ‘Philosophia Botanica’ (1751), must be thoroughly convinced that the ideas on which his theories are based are to be found scattered up and down in the works of his predecessors; further, whoever has traced the history of the sexual theory from the time of Camerarius (1694), must allow that Linnaeus added nothing new to it, though he contributed essentially to its recognition, and that even after Koelreuter’s labours he continued to entertain some highly obscure and even mystical notions on the subject.
But that which gave Linnaeus so overwhelming an importance for his own time was the skilful way in which he gathered up all that had been done before him; this fusing together of the scattered acquisitions of the past is the great and characteristic merit of Linnaeus.
Cesalpino was the first who introduced Aristotelian modes of thought into botany; his system was intended to be a natural one, but it was in reality extremely unnatural; Linnaeus, in whose works the profound impression which he had received from Cesalpino is everywhere to be traced, retained all that was important in his predecessor’s views, but perceived at the same time what no one before him had perceived, that the method pursued by Cesalpino, Morison, Ray, Tournefort, and Bachmann could never do justice to those natural affinities which it was their object to discover, and that in this way only an artificial though very serviceable arrangement could be attained, while the exhibition of natural affinities must be sought by other means.
As regards the terminology of the parts of plants, which was all that the morphology of the day attempted, Linnaeus simply adopted all that was contained in the Isagoge of Jung, but gave it a more perspicuous form, and advanced the theory of the flower by accepting without hesitation the sexual importance of the stamens, which was still but little attended to; he thus arrived at a better general conception of the flower, and this bore fruit again in a terminology which is as clear as it is convenient; the terms monoecious, dioecious, triandrous, monogynous, etc., still used in the science, and the later-invented expressions dichogamous, protandrous, protogynous, etc., owe their origin to this correct conception of the sexual relations in plants. But there was one great misconception in the matter, which has not a little contributed to increase Linnaeus’ reputation. He called his artificial system, founded on the number, union, and grouping of the stamens and carpels, the sexual system of plants, because he rested its supposed superiority on the fact, that it was founded upon organs the function of which lays claim to the very highest importance. But it is obvious that the sexual system of Linnaeus would have the same value for the purposes of classification, if the stamens had nothing whatever to do with propagation, or if their sexual significance were quite unknown. For it is exactly those characters of the stamens which Linnaeus employs for purposes of classification, their number and mode of union, which are matter of entire indifference as regards the sexual function.
But though the notion that this artificial system has any important connection with the doctrine of the sexuality of plants is evidently due to a confusion of ideas, yet the progress of the science has shown, that Linnaeus’ sexual system did often and necessarily lead to the establishing of natural groups for the very reason, that the characters of the stamens which he employed are entirely independent of their function; for we must regard it as an important result of the labours of systematists, that those characters of organisms are shown to be of the greatest value for classification, which are entirely or in a very great measure independent of the functions of the organs. The error, which led Cesalpino to make the functional importance of the parts of fructification the principle of his division, reappears therefore in Linnaeus in another form; to find a principle of division, he turns to those organs, whose function appears to him the most important, but he takes his characters not from differences of function, but from the number and mode of union, which are of no importance for the sexual function. We meet with this error in Leibnitz and Burckhard, who are mentioned here merely to defend Linnaeus from the charge repeatedly brought against him by his contemporaries that he was indebted to these two writers for the idea of his sexual system. They erroneously found in the great physiological importance of the sexual organs a reason for deriving from their differences the principles of division that were to found a system; this error in theory Linnaeus shared with them, but they did not correct it in practice, as Linnaeus did, by confining himself to purely morphological features in working out his system. What the renowned philosopher[27] incidentally uttered in the year 1701 on the matter in question is moreover so unimportant and so indistinct, that Linnaeus could not gain much from it; what Burckhard[28] says on the subject in his often-quoted letter to Leibnitz (1702) is indeed much better, and comes near to Linnaeus’ idea; but it is a very long way from the hints there given to the completion of the well-articulated and highly practical system which Linnaeus constructed.
The botanists of the 16th century, and in the main even Morison and Ray, had in one-sided fashion devoted their chief attention to distinguishing species, Bachmann and Tournefort to the establishment of generic characters, while they neglected species; Linnaeus, on the contrary, applied equal care and much greater skill to describing both genera and species. He reduced to practical shape the suggestion which Bachmann had left to his successors, and so must be regarded, if not as the inventor, at least as the real founder of the binary nomenclature of organisms.
It is only fulfilling the duty of a historian to state the sources from which Linnaeus drew, but it would be a misapprehension to see in this any depreciation of a great man; it were to be desired that all naturalists would, like Linnaeus, adopt all that is good in the contributions of their predecessors, and improve or adapt it as he did. Linnaeus himself has repeatedly quoted the sources of his knowledge as far as they were known to him, and has in many cases estimated the services of his predecessors with a candour which never betrays a trace of jealousy, but often displays a warm respect, as may be seen especially in the short introductions to the several systems given in the ‘Classes Plantarum.’ Linnaeus could not only recognise what was good in his predecessors and occasionally make use of it, but he imparted life and fruitfulness to the thoughts of others by applying them as he applied his own thoughts, and bringing out whatever theoretical value they possessed. It was evidently this freshness of life that often misled his successors into believing that Linnaeus thought out and discovered everything for himself. We learn to appreciate the contributions of Cesalpino and his successors in the 17th century, and even of Kaspar Bauhin for the first time in the works of Linnaeus; we are astonished to see the long-known thoughts of these writers, which in their own place look unimportant and incomplete, fashioned by Linnaeus into a living whole; thus he was at once and in the best sense both receptive and productive, and he might perhaps have done more for the theory of the science if he had not been entangled in one grave error, which was more sharply pronounced in him than in his predecessors and contemporaries, that, namely, of supposing that the highest and only worthy task of a botanist is to know all species of the vegetable kingdom exactly by name. Linnaeus distinctly declared that this was his view, and his school in Germany and England adhered to it so firmly that it established itself with the general public, who to the present day consider it as a self-evident proposition that a botanist exists essentially for the purpose of at once designating any and every plant by a name. Like his predecessors, Linnaeus regarded morphology and general theoretical botany only as means to be used for discovering the principles of terminology and definition, with a view to the improvement of the art of describing plants.
We have hitherto spoken chiefly of the manner in which Linnaeus dealt with his subject in matters of detail; in his inner nature he was a schoolman, and that in a higher degree than even Cesalpino himself, who should rather be called an Aristotelian in the strict sense of the word. But to say that Linnaeus’ mode of thought is thoroughly scholastic is virtually saying that he was not an investigator of nature in the modern meaning of the word; we might point to the fact that Linnaeus never made a single important discovery throwing light on the nature of the vegetable world; but that would still not prove that he was a schoolman.
True investigation of nature consists not only in deducing rules from exact and comparative observation of the phenomena of nature, but in discovering the genetic forces from which the causal connexion, cause and effect may be derived. In the pursuit of these objects, it is compelled to be constantly correcting existing conceptions and theories, producing new conceptions and new theories, and thus adjusting our own ideas more and more to the nature of things. The understanding does not prescribe to the objects, but the objects to the understanding. The Aristotelian philosophy and its medieval form, scholasticism, proceeds in exactly the contrary way; it is not properly concerned with acquiring new conceptions and new theories by means of investigation, for conceptions and theories have been once for all established; experience must conform itself to the ready-made system of thought; whatever does not so conform must be dialectically twisted and explained till it apparently fits in with the whole. From this point of view the intellectual task consists essentially in this twisting and turning of facts, for the general idea of the whole is already made and needs not to be altered. Experience in the higher sense of investigation of nature is rendered impossible by the fact, that we are supposed to know all the ultimate principles of things; but these ultimate principles of scholasticism are at bottom only words with extremely indefinite meaning, abstractions obtained by a series of jumps from every-day experience, which has not been tried and refined in the crucible of science, and is therefore worthless; and the higher the abstraction is raised, the farther it withdraws from the guiding hand of experience, the more venerable and more important do these ‘abstracta’ appear, and we can finally come to a mutual understanding about them, though again only through figures and metaphors[29]. Science, according to the scholastic method, is a playing with abstract conceptions; the best player is he who can so combine them together, that the real contradictions are skilfully concealed. On the contrary, the object of true investigation, whether in philosophy or in natural science, is to make unsparing discovery of existing contradictions and to question the facts until our conceptions are cleared up, and if necessary the whole theory and general view is replaced by a better. In the Aristotelian philosophy and in scholasticism facts are merely examples for the illustration of fixed abstract conceptions, but in the real investigation of nature they are the fruitful soil from which new conceptions, new combinations of thought, new theories, and general views spring and grow. The most pernicious feature in scholasticism and the Aristotelian philosophy is the confounding of mere conceptions and words with the objective reality of the things denoted by them; men took a special pleasure in deducing the nature of things from the original meaning of the words, and even the question of the existence or non-existence of a thing was answered from the idea of it. This way of thinking is found everywhere in Linnaeus, not only where he is busy as systematist and describer, but where he wishes to give information on the nature of plants and the phenomena of their life, as in his ‘Fundamenta,’ his ‘Philosophia Botanica,’ and especially in his ‘Amoenitates Academicae.’ From among many instances we may select his mode of proving sexuality in plants. Linnaeus knew and lauded the services rendered to botany by Rudolph Jacob Camerarius, who as a genuine investigator of nature had demonstrated the sexuality of plants in the only possible way, namely, that of experiment. But Linnaeus cares little for this experimental proof; he just notices it in passing, and expends all his art on a genuine scholastic demonstration intended to prove the existence of sexuality as arising necessarily from the nature of the plant. He connects his demonstration with the dictum ‘omne vivum ex ovo,’ which Harvey had founded on an imperfect induction, and which he evidently takes for an a priori principle, and concludes from it that plants also must proceed from an ‘ovum,’ overlooking the fact that in ‘omne vivum ex ovo’ plants already form a half of the ‘omne vivum’; then he continues, ‘reason and experience teach us that plants proceed from an ‘ovum,’ and the cotyledons confirm it’; reason, experience, and cotyledons! Surely a remarkable assemblage of proofs. In the next sentence he confines himself at first to the cotyledons, which according to him spring in animals from the yolk of the egg, in which the life-point is found; consequently, he says, the seed-leaves of plants, which envelope the ‘corculum,’ are the same thing; but that the progeny is formed not simply from the ‘ovum,’ nor from the fertilising matter in the male organs, but from the two combined, is shown by animals, hybrids, reason, and anatomy. By reason in this and the previous sentence he understands the necessity, concluded from the nature, that is, the conception of the thing, that it must be so; animals supply him with the analogy, and anatomy can prove nothing, as long as it is not known what is the design of the anatomical arrangements. But the weakest side of this proof lies in the hybrids, for Linnaeus, when he wrote the ‘Fundamenta,’ knew of none except the mule; hybrids in plants were first described by Koelreuter in 1761, and these Linnaeus nowhere mentions; and what amount of proof can be drawn from the vegetable hybrids, which Linnaeus afterwards supposed himself to have observed, but which were no hybrids, we shall see in the history of the sexual theory; here we need only remark that he arrives at the existence of these hybrids from the idea of sexuality exactly as he arrived at that of sexuality from the idea of hybridisation. Then he goes on with his demonstration; ‘that an egg germinates without fecundation is denied by experience, and this must hold good therefore of the eggs[30] of plants—every plant is provided with flower and fruit, even where these are not visible to the eye’; with Linnaeus, of course, this is logically concluded from the conception of the plant or of the ‘ovum’; he alleges indeed certain observations as well, but they are incorrect. He continues, ‘The fructification consists of the sexual organs of the flowers; that the anthers are the male organs, the pollen the fertilising matter, is proved by their nature, further by the fact that the flower precedes the fruit, as also by their position, the time, the loculaments (anthers), by castration, and by the structure of the pollen.’ Here too the main point with Linnaeus is the nature of the male organs, and that we may know what this nature is he refers to a former paragraph, where we learn that the essence of the flower is in the anthers and stigma. Almost all his demonstrations consist of such reasonings in a circle and in arguing from the thing to be proved. And while the passages quoted show how much he did for the doctrine of sexuality, we find this sophistical style of reasoning still more copiously displayed in the essay entitled ‘Sponsalia Plantarum’ in the ‘Amoenitates’ (i. p. 77), and in a worse form still in the essay, ‘Plantae Hybridae’ (Amoen. iii. p. 29). That Linnaeus had not the remotest conception of the way in which the truth of a hypothetical fact is proved on the principles of strict inductive investigation is shown by these and many other examples, and by his enquiry into the seeds of mosses (Amoen. ii. p. 266), upon which he prided himself not a little, but which is really inconceivably bad even for that time (1750). It was not Linnaeus’ habit to occupy himself with what we should call an enquiry; whatever escaped the first critical glance he left quietly alone; it did not occur to him to examine into the causes of the phenomena that interested him; he classified them and had done with them; as for instance in his ‘Somnus Plantarum,’ as he called the periodical movements of plants. We cannot read much of the ‘Philosophia Botanica’ or the ‘Amoenitates’ without feeling that we are transported into the literature of the middle ages by the kind of scholastic sophistry which is all that his argumentation amounts to; and yet these works of Linnaeus date from the middle of the last century, from a time when Malpighi, Grew, Camerarius, and Hales had already carried out their model investigations, and his contemporaries Duhamel, Koelreuter, and others were experimenting in true scientific manner. This peculiarity in Linnaeus explains why men like Buffon, Albert Haller, and Koelreuter treated him with a certain contempt; and also why his strict adherents in Germany, who lived on his writings and were unable to separate what was really good in him from his mode of reasoning, came to make their own botany like anything rather than a science of nature. Linnaeus was in fact a dangerous guide for weak minds, for his curious logic, among the worst to be met with in the scholastic writers, was combined with the most brilliant powers of description; the enormous extent of his knowledge of particulars, and above all the pre-eminent firmness and certainty which distinguished his mode of dealing with systematic botany, could not fail to make the profoundest impression on those who judged of the powers of an investigator of nature by these qualities alone. One of his greatest gifts was without doubt the power which he possessed of framing precise and striking descriptions of species and genera in the animal and vegetable kingdoms by means of a few marks contained in the smallest possible number of words; in this point he was a model of unrivalled excellence to all succeeding botanists.
On the whole the superiority of Linnaeus lay in his natural gift for discriminating and classifying the objects which engaged his attention; he might almost be said to have been a classifying, co-ordinating, and subordinating machine. He dealt with everything about which he wrote in the way in which he dealt with objects of natural history. The systematic botanists whom he mentions in the ‘Classes Plantarum’ are classified then and there as fructists, corollists, and calycists. All who occupy themselves in any way with botany are divided into two great classes, the true botanists and mere botanophils, and it is very characteristic of his way of thinking that he places anatomists, gardeners, and physicians in the latter class. True botanists again are either mere collectors or systematists. To the collectors belong all who add to the number of known plants, also authors of monographs and floras, and the botanical explorers of foreign countries, whom we should now more courteously call systematists. By systematists Linnaeus understands those who occupy themselves with the classification and naming of plants, and he divides them into philosophers, systematists proper, and nomenclators; the philosophers are those who study the theory of the science on principles founded on reason and observation, and are subdivided into orators, institutors, erystics, and physiologists; the latter are those who discovered the mystery of sexuality in plants, and hence Malpighi, Hales, and such men are not physiologists in Linnaeus’ sense. The second class of systematists, the systematists proper, he distinguishes into orthodox and heterodox, the former taking the grounds of division exclusively from the organs of fructification, while the latter use other marks as well. In this manner Linnaeus treats every subject of which he has to speak, and wherever he can in short, numbered sentences, which look like descriptions of genera and species. His mind and character were fully formed in 1736 when he wrote his ‘Fundamenta,’ and he preserved his peculiarities of style from that time forward; we find the same modes of expression in the ‘Nemesis Divina,’ a treatise on religion and morals addressed as a legacy to his son. Where these peculiarities of manner and expression are suitable they make a favourable impression on the reader, as for instance in the short accounts he gives of the various systems in the ‘Classes Plantarum,’ a work in which Linnaeus was quite in his element; there he traces with a fine instinct the guiding principles of each system, pronounces upon its merits and defects, and sets it before the reader in numbered sentences of epigrammatic brevity. This manner is strictly adhered to in the ‘Philosophia’ also, and it has certainly helped not a little to withdraw the attention of his reader from his many fallacies in argument, especially his oft-recurring reasonings in a circle.
This remarkable combination of an unscientific philosophy with mastery over the classification of things and conceptions, this mixture of consistency in carrying out his scholastic principles with gross inaccuracies of thought, impart to his style an originality, which is rendered still more striking by the native freshness and directness, and not unfrequently by the poetic feeling, which animate his periods.
In any attempt to estimate the advance which the science owes to the labours of Linnaeus, the chief prominence must be assigned to two points; first to his success in carrying out the binary nomenclature in connection with the careful and methodical study which he bestowed on the distinguishing of genera and species; this system of nomenclature he endeavoured to extend to the whole of the then known vegetable world, and thus descriptive botany in its narrower sense assumed through his instrumentality an entirely new form, which, serving as a model for the naming and defining of the larger groups, could be applied without modification to the founding and completing the natural system. When at a later time Jussieu and De Candolle marked out their families and groups of families, their mode of proceeding was in the main that of Linnaeus when distinguishing his genera by abstraction of specific differences. This merit has been always assigned to Linnaeus without reserve. The second merit has been less recognised, and yet it is at least of equal importance; it is that of having first perceived that the attempt made by Cesalpino and his successors to found a system, that shall do justice to natural affinities, on predetermined marks can never succeed. Linnaeus framed his artificial sexual system, but he exhibited a fragment of a natural system by its side, while he repeatedly declared that the chief task of botanists is to discover the natural system. Thus he cleared the ground for systematic botany. He made use of his own system, because it was extremely convenient for describing individual plants, but he ascribed all true scientific value exclusively to the natural system; and with what success he laboured to advance it may be gathered from the fact, that Bernard de Jussieu founded his improved series of families on the fragment of Linnaeus, and that his nephew, A. L. de Jussieu, by simply adopting Linnaeus’ conception of the principle which lies at the foundation of the natural system, succeeded in carrying it on to a further stage of development.
The main features of Linnaeus’ theoretical botany can best be learned from the ‘Philosophia Botanica,’ which may be regarded as a text-book of that which Linnaeus called botany, and which far surpasses all earlier compositions of the kind in perspicuity and precision, and in copiousness of material; and indeed it would be difficult to find in the ninety years after 1781 a text-book of botany which treats what was known on the subject at each period with equal clearness and completeness. In giving the reader some idea of the way in which Linnaeus deals with his subject, it will be well to pass over the first two chapters, which discuss the literature and the various systems which had been proposed, and turn to the third, which under the heading ‘Plantae’ treats of the general nature of plants, and specially of the organs of vegetation. The vegetable world, says Linnaeus, comprises seven families, Fungi, Algae, Mosses, Ferns, Grasses, Palms, and Plants. All are composed of three kinds of vessels, sap-vessels which convey the fluids, tubes which store up the sap in their cavities, and tracheae which take in air; these statements Linnaeus adopts from Malpighi and Grew. He gives no characteristic marks for the Fungi; of the Algae he says that in them root, leaf, and stem are all fused together; to the Mosses he ascribes an anther without a filament, and separate from the female flower which has no pistil; the seeds of the Mosses have no integument or cotyledons; this characteristic of the Mosses is explained in his paper entitled ‘Semina Muscorum’ in the ‘Amoenitates Academicae,’ ii. The Ferns are marked by the fructification on the under side of the fronds, which are therefore not conceived of as leaves. The very simple leaves, the jointed stalk, the ‘calyx glumosus,’ and the single seed mark the Grasses. The simple stem, the rosette of leaves at the summit, and the spathe of the inflorescence are characteristic of the Palms. All vegetable forms which do not belong to any of the previous families he names Plants. He rejects the customary division into herbs, shrubs, and trees as unscientific. This arrangement of the vegetable kingdom must not be confounded with Linnaeus’ fragment of a natural system, in which he adopts sixty-seven families (orders), the Fungi, Algae, Mosses, and Ferns forming each a family. He evidently introduces the divisions in the ‘Philosophia,’ in order that it may be seen how far the statements that follow are applicable to all the Vegetabilia or only to certain sections of them. The parts in the individual plant which the beginner must distinguish are three; the root, the herb[31], and the parts of fructification, in which enumeration Linnaeus departs from his predecessors, by whom the fructification and the herb together are opposed to the root. In the central part of the plant is the pith, enclosed by the wood which is formed from the bast; the bast is distinct from the rind, which again is covered by the epidermis; these anatomical facts are from Malpighi; the statement that the pith grows by extending itself and its envelopes is borrowed from Mariotte. Cesalpino’s view on the formation of the bud is expressed by Linnaeus in the statement, that the end of a thread of the pith passing through the rind is resolved into a bud, etc. The bud is a compressed stem, capable of unlimited extension till fructification puts a term to vegetation. The fructification is formed by the leaves uniting into a calyx, from which the apex of a branch issues as a flower about one year in advance, while the fruit arising from the substance of the pith cannot begin a new life till the woody substance of the stamens has been absorbed by the fluids of the pistil. In this way Linnaeus corrected Cesalpino’s theory of the flower, that he might take into account the sexual importance of the stamens discovered by Camerarius. He concludes by saying that there is no new creation but only a continuous generation, for which he gives the remarkable and thoroughly Cesalpinian reason, ‘cum corculum seminis constat parte radicis medullari.’
The root, which takes up the food, and produces the stem and the fructification, consists of pith, wood, bast, and rind, and is divided into the two parts, ‘caudex’ and ‘radicula.’ The ‘caudex’ answers pretty nearly to our primary root and rhizomes, the ‘radicula’ to what we now call secondary roots.
The herb springs from the root, and is terminated by the fructification; it consists of the stem, leaves, leaf-supports (‘fulcrum’), and the organs of hibernation (‘hibernaculum’). Then follow the further distinctions of stem and leaves; the terminology, still partly in use and resting essentially on the definitions of Jung, is here set forth in great detail. Linnaeus however does not mention the remarkable distinction between stem and leaf which Jung founded on relations of symmetry, and in general he shows less depth of conception than Jung, confining himself more to the direct impression on the senses, and so distinguishing sometimes where there is no real difference. Examples of this are furnished by the paragraph devoted to ‘fulcra.’ By this term he designates the subsidiary organs of plants, among which he reckons stipules, bracts, spines, thorns, tendrils, glands, and hairs. It appears from this, that Linnaeus did not extend the idea of the leaf (‘folium’) to stipules and bracts, and the examples he gives of tendrils show at the same time that he was ignorant of the different morphological character of the organ in Vitis and Pisum. The putting the seven organs above-named together under the idea of ‘fulcrum’ shows plainly enough that Linnaeus, in framing his terminology, aimed only at distinguishing what was different to the sense by fixed words, in order to obtain means for short diagnoses of species and genera. He had no thought of arriving at more general propositions from a comparison of forms in plants, in order to attain to a deeper insight into their nature. The same thing appears from his notion of ‘hibernaculum,’ by which he understands a part of the plant which envelopes the stem in its embryonal state and protects it from harm from without; he here distinguishes bulbs from the winter buds of woody plants. In this course of mixing up morphological and biological relations of organs he was followed by botanists till late into our own century.
Linnaeus goes far beyond his predecessors in distinguishing and naming the organs of fructification, the subject of the fourth chapter of the ‘Philosophia Botanica.’ The fructification, he says, is a temporary part in plants devoted to propagation, terminating the old and beginning the new. He distinguishes the following seven parts: (1) the calyx, which represents the rind, including in this term the involucre of the Umbelliferae, the spathe, the calyptra of Mosses, and even the volva of certain Fungi,—another instance of the way in which Linnaeus was guided by external appearance in his terminology of the parts of plants; (2) the corolla, which represents the inner rind (bast) of the plant; (3) the stamen, which produces the pollen; (4) the pistil, which is attached to the fruit and receives the pollen; here for the first time the ovary, style, and stigma are clearly distinguished. But next comes as a special organ (5) the pericarp, the ovary which contains the seed. As bulbs and buds were treated not simply as young shoots, but as separate organs, so here too the ripe fruit is regarded not merely as the developed ovary, but as a special organ. Nevertheless, Linnaeus distinguishes the different forms of fruit much better than his predecessors had done. (6) The seed is a part of the plant that falls off from it, the rudiment of a new plant, and it is excited to active life by the pollen. The treatment of the seed and its parts is the feeblest of all Linnaeus’ efforts; he follows Cesalpino, but his account of the parts of the seed is much more imperfect than that of Cesalpino and his successors. The embryo is called the ‘corculum,’ and two parts are distinguished in it, the ‘plumula’ and the ‘rostellum’ (radicle). The cotyledon is co-ordinated with the ‘corculum,’ and is regarded therefore not as part of the embryo but as a distinct organ of the seed; it is defined as ‘corpus laterale seminis bibulum caducum.’ Nothing could be worse, and it seems almost incredible that so bad a definition and distinction could be given in 1751, and again in 1770, by the first botanist of his time, when Malpighi and Grew, nearly a hundred years earlier, had illustrated the parts of the seed and even the history of its development and its germination by numerous figures. He does not mention the endosperm, evidently confounding it with the cotyledon, though Ray had already distinguished it clearly from the other parts of the seed. Linnaeus’ terminology of the seed supplies more than sufficient corroboration of our previous remark, that he shows incapacity for the careful investigation of any object at all difficult to observe, and it will now seem a small matter that he, like most of the earlier botanists, treats one-seeded indehiscent fruits as seeds, and hence makes the pappus a part of the seed. (7) By the word ‘receptaculum’ he understands everything by which the parts of the fructification are connected together, both the ‘receptaculum proprium,’ which unites the parts of the single flower, and the ‘receptaculum commune,’ under which term he comprises the most diverse forms of inflorescence (umbel, cyme, spadix).
He concludes with the remark that the essence of the flower consists in the anther and the stigma, that of the fruit in the seed, that of the fructification in the flower and the fruit, and that of all vegetable forms in the fructification, and he adds a long list of distinctions between the organs of fructification with their names; among these organs appear the nectaries, which he was the first to distinguish.
In the fifth chapter he discusses the question of difference of sex in plants. His views on this subject have been already mentioned in order to show that they were entirely founded on worthless scholastic deductions; here we may quote a few of the propositions which were famous in after times. We assume, he says, that two individuals of different sexes were created in the beginning of things in every kind of living creatures. Plants, though they are without sensation, yet live as do animals, for they have a beginning and an advance in age (aetas), and are liable to disease and death; they have also a power of movement, a natural appetency (propulsio), an anatomy, and an organic structure (organismus). Simple explanations are given of these words, but they prove nothing about the matter. He then expounds the whole theory of sexuality, which is made to rest entirely on scholastic arguments, and in doing this he spins out to excessive length the parallel which he draws between the conditions of sexuality in animals and plants. It is manifestly this chapter of the ‘Philosophia Botanica,’ together with the treatise ‘Sponsalia Plantarum,’ which led the adherents of Linnaeus, who were ignorant of the older literature of the subject and were much impressed by his scholastic dexterity, to celebrate him as the founder of the sexual theory of plants; whereas a more careful study of history shows incontrovertibly that Linnaeus helped in this way to disseminate the doctrine, but did absolutely nothing to establish it.
The writings of Linnaeus which we have hitherto examined are occupied with the nature of plants, and of this he knew nothing more than he gathered from the investigations and reflections of his predecessors; and it is here especially that his peculiar scholasticism is exhibited in contrast with the facts obtained by induction which he communicated to his readers. But the strong side of his intellect appears with splendid effect in the succeeding chapters of the ‘Philosophia,’ which treat of the principles of systematic botany; here, where he has no longer to establish facts, but to arrange ideas, to dispose and summarise, we find Linnaeus thoroughly in his element.
The groundwork of botanical science, he begins, is twofold, classification and naming. The constituting of classes, orders, and genera he calls theoretical classification; the constituting of species and varieties is practical classification. The work of classification carried out by Cesalpino, Morison, Tournefort, and others leads to the establishing of a system; the mere practice of describing species may be carried on by those who know nothing of systematic botany. These expressions of Linnaeus are interesting, because like other remarks of his they show that he placed the establishment and arrangement of the larger groups above the mere distinguishing of individual forms; his disciples to a great extent forgot their master’s teaching, and fancied that the collecting and distinguishing of species was systematic botany. He opposes the system itself, which deals with the relative conceptions of classes, orders, genera, species, and varieties, to a mere synoptical view, serving with its dichotomy only to practical ends. Then comes the often-quoted sentence, ‘We reckon so many species as there were distinct forms created “in principio.”’ In a former place he had said ‘ab initio’ instead of ‘in principio’; instead therefore of a beginning in time he here posits an ideal, theoretical beginning, which is more in accordance with his philosophical views. That new species can arise is, he continues, disproved by continuous generation and propagation, and by daily observation, and by the cotyledons. It is hard to understand how the Linnaean school till far into our own century could have remained firm in a doctrine resting on such arguments as these. Linnaeus’ definition of varieties shows that he understood by the word species fundamentally distinct forms; there are, he says, as many varieties as there are different plants growing from the seed of the same species; and he adds that a variety owes its origin to an accidental cause, such as climate, soil, warmth, the wind; but this is evidently mere arbitrary assumption. Judging by all he says, his view is that species differ in their inner nature, varieties only in outward form. Here, where we find the dogma of the constancy of species for the first time expressed in precise terms,—a dogma generally accepted till the appearance of the theory of descent, we should be justified in demanding proof; but since dogmas as a rule do not admit of proof, Linnaeus simply states his view[32], unless we are to take the sentence, ‘negat generatio continuata, propagatio, observationes quotidianae, cotyledones,’ as proving the assertion that new species never appear. We shall see further on to what surprising conclusions Linnaeus was himself led by his dogma, when he had to take into account the relations of affinity in genera and larger groups. The species and the genus, he continues, are always the work of nature, the variety is often that of cultivation; the class and the order depend both on nature and on art, which must mean that the larger groups of the vegetable kingdom have not the same objective reality as the species and the genus, but rest partly on opinion. That Linnaeus estimated the labours of the systematists after Cesalpino and the contributions of the German fathers of botany up to Bauhin, as they have been judged of in the present work, is shown by paragraph 163, where he explains the word habit, and adds that Kaspar Bauhin and the older writers had excellently divined (divinarunt) the affinities of plants from their habit, and even real systematists had often erred, where the habit pointed out to them the right way. But he says that the natural arrangement, which is the ultimate aim of botany, is founded, as the moderns have discovered, on the fructification, though even this will not determine all the classes. It is interesting therefore to observe how Linnaeus further on (paragraph 168) directs, that in forming genera, though they must rest on the fructification, yet it is needful to attend to the habit also, lest an incorrect genus should be established on some insignificant mark (levi de causâ): but this attention to the habit must be managed with reserve, so as not to disturb the scientific diagnosis.
Linnaeus next lays down with great detail each several rule, which must be observed in establishing species, genera, orders, and classes, and it is here that he displays his unrivalled skill as a systematist. These rules were strictly observed by himself in his numerous descriptive works, and thus a spirit of order and clearness was introduced into the art of describing plants, which gave it at once a different appearance from that which it had received at the hands of his predecessors. Whoever therefore compares the ‘Genera Plantarum,’ the ‘Systema Naturae,’ and other descriptive works of Linnaeus with those of Morison, Ray, Bachmann, or Tournefort, finds so great a revolution effected by them, that he is impressed with the persuasion that botany first became a science in the hands of Linnaeus; all former efforts seem to be so unskilful and without order in comparison with his method. Without doubt the greatest and most lasting service which Linnaeus rendered both to botany and to zoology lies in the certainty and precision which he introduced into the art of describing. But if a reformation was thus effected in botany, as Linnaeus himself took pleasure in saying, it must not be overlooked that the knowledge of the nature of plants was rather hindered than advanced by him. Ray, Bachmann, and in part also Morison and Tournefort, had already liberated themselves to a great extent from the influence of scholasticism, and they still give us the impression of having been genuine investigators of nature; but Linnaeus fell back again into the scholastic modes of thought, and these were so intimately combined with his brilliant performances in systematic botany, that his successors were unable to separate the one from the other.
The feeling for order and perspicuity, which made Linnaeus a reformer of the art of describing, combined with his scholasticism, was evidently the cause of his not bestowing more energetic labour on the natural system. It has been repeatedly mentioned that it was he who first established sixty-five truly natural groups in his fragment of the early date of 1738; and a certain feeling for natural affinity is shown in the establishment of his seven families, Fungi, Algae, Mosses, Ferns, Grasses, Palms, and Plants properly so-called. Moreover in paragraph 163 of the ‘Philosophia Botanica,’ he carries out the division of the whole vegetable kingdom into Acotyledons, Monocotyledons, and Polycotyledons with their subdivisions very admirably; and thus we see him continually impelled towards a natural arrangement, but never bestowing upon it the necessary labour and thought.
And so two different conceptions of a system of plants continued to subsist side by side with each other in the mind of Linnaeus; one more superficial, and adapted for practical use, expressed in his artificial sexual system, and one more profound and scientifically valuable, embodied in his fragment and in the natural groups above-mentioned.
The same may be said also of Linnaeus’ morphological views; here, too, a more superficial pursued its way along with a more profound conception. He formed his terminology of the parts of plants for practical use in describing them, and convenient as it is, it seems nevertheless shallow or superficial, because its foundations are not more deeply laid in the comparative study of forms. But we discover from very various passages in his writings that he felt the need of a more profound conception of plant-form, and what he was able to say on the subject he put together under the head of ‘metamorphosis plantarum.’ His doctrine of metamorphosis is entirely based on the views of Cesalpino, with which we have already become acquainted, though he did not adopt them in their original form, but endeavoured to develop them in true Cesalpinian fashion; for on the one hand he derived leaves and parts of flowers from the tissues of the stem, and on the other conceived of the parts of the flower as only altered leaves. This doctrine of metamorphosis appears in somewhat confused form in the last page of his ‘Philosophia Botanica.’ There he says that the whole of the herb is a continuation of the medullary substance of the root; the principle of the flowers and leaves is the same, because both spring from the tissue-layers surrounding the pith, as Cesalpino had taught. The statement which follows, that the principle of the bud and the leaves is identical, would be a departure from Cesalpino, and in any case inconsistent, without the explanation that the bud consists of rudimentary leaves; but this again puts the axial portion of the bud out of sight. The perianth, he says, comes from concrescent rudiments of leaves. How closely Linnaeus adhered to Cesalpino in his later years appears in his explanation of the catkin, which comes next and which is taken entirely from Cesalpino’s theory. That a more superficial and a more profound conception pursue their way together unadjusted in Linnaeus’ speculations on form is specially shown by the fact, that in the text of the ‘Philosophia Botanica,’ paragraph 84, he places the ‘stipulae’ under the idea of ‘fulcra’ and not under that of ‘folia,’ while on the contrary at the end of the same work, where he brings together the different paragraphs respecting metamorphosis, he speaks of the ‘stipulae’ as appendages of the leaves.
The idea of Cesalpino, that the parts of the flower which surround the fruit arise like the ordinary leaves from the tissues that enclose the pith, is further developed by Linnaeus in his ‘Metamorphosis Plantarum,’ in the fourth volume of the ‘Amoenitates Academicae’ (1759), in a very strange manner. He compares the formation of the flower with the metamorphosis of animals, and especially of insects, and after describing the changes that take place in animals, he says at page 370 that plants are subject to similar change. The metamorphosis of insects consists in the putting off different skins, so that they finally come forth naked in their true and perfect form. This metamorphosis we also find in most plants, for they consist, at least in the truly living part of the root, of rind, bast, wood, and pith. The rind is to the plant what the skin is to the larva of an insect, and after putting this skin off there remains a naked insect. When the flower is produced in the plant the rind opens and forms the calyx (exactly Cesalpino’s view), and from out of this the inner parts of the plant issue to form the flower, so that the bast, the wood, and the pith issue forth naked in the form of corolla, stamens, and stigma. So long as the plant lies concealed within the rind and clothed only with leaves, it appears to us as unrecognisable and obscure as a butterfly, which in its larva-condition is covered with skin and spines.
In this doctrine of metamorphosis, which Linnaeus founded on Cesalpino, the chief point to observe is, that the ordinary leaves are identical with the exterior parts of the flower, because both originate in the outer tissues of the stem. The pertinent fact, which may easily be observed without a microscope, that the concentric arrangement of outer and inner rind, wood, and pith occurs only in some flowering plants, that the case is quite different with Monocotyledons, and that Cesalpino’s theory of the flower cannot properly be applied to them,—these are things which we must not expect to find Linnaeus with his peculiar modes of thought taking into consideration.
The want of firm standing-ground in experience is shown also by the fact, that with his own and Cesalpino’s theory of the flower he combined another view of its nature, which under the name of ‘prolepsis plantarum’ was set forth in two dissertations in 1760 and 1763, but the two theories are scarcely compatible with one another. While the last paragraph in the ‘Philosophia Botanica’ says, ‘Flos ex gemmâ annuo spatio foliis praecocior est,’ the dissertations contain the doctrine[33], that the flower is nothing but the synchronous appearance of leaves, which properly belong to the bud-formations of six consecutive years, in such a way that the leaves of the bud destined to be unfolded in the second year of the plants become bracts, the leaves of the third year the calyx, those of the fourth the corolla, those of the fifth the stamens, those of the sixth the pistil. Here we see once more how Linnaeus moves in the sphere of arbitrary assumptions with no thought of exact observation, for this whole theory of prolepsis rests on nothing that can be called a well-ascertained fact.
Yet a third time we find in Linnaeus the juxtaposition of a superficial view resting on every-day perception, and a more profound and to some extent a philosophical view; this is the case where he is concerned on the one hand with the dogma of the constancy of species, and on the other hand has to explain the fact of natural relationship and its gradations. Apart from some insignificant verbal explanations, Linnaeus adduced nothing in support of the dogma but the every-day perception of the unchangeableness of species, and to this he held fast to the end of his life; but it was important to find an explanation of the fact, to which he himself repeatedly drew attention, that genera, orders, and classes do not merely rest on opinion but indicate really existing affinities. His mode of solving the difficulty was a very remarkable one; not only does the scholastic manner of thought appear here again quite unalloyed by modern science, but he grounds his explanation once more on the old a priori notion that the pith is the vital principle in the plant, and also on his own assumption, that in the sexual act the woody substance of the anthers combines with the pith-substance of the pistil. Hugo Mohl has given a clear account of the matter in No. 46 of the ‘Botanische Zeitung’ for 1870, although neither he nor Wigand nor most of Linnaeus’ biographers seem to know, that his theories are all to be traced to Cesalpino. Linnaeus’ theory of natural affinities, as he gave it in 1762 in the ‘Fundamentum Fructificationis,’ and in 1764 in the sixth edition of the ‘Genera Plantarum,’ is as follows: At the creation of plants (in ipsa creatione) one species was made as the representative of each natural order, and these plants so corresponding to the natural orders were distinct from one another in habit and fructification, that is, absolutely distinct. In the communication of 1764 the following words occur:—
1. Creator T.O. in primordio vestiit vegetabile medullare principiis constitutivis diversi corticalis, unde tot difformia individua, quot ordines naturales, prognata.
2. Classicas has plantas Omnipotens miscuit inter se, unde tot genera ordinum, quot inde plantae.
3. Genericas has miscuit natura, unde tot species congeneres, quot hodie existunt.
4. Species has miscuit casus, unde totidem quot passim occurrunt varietates.
Hugo Mohl was right in rejecting Heufler’s assumption that a view resembling the modern theory of descent was contained in these paragraphs. It must be plain to any one who knows the ideas of Aristotle, Theophrastus, and Cesalpino, within the sphere of which Linnaeus is here moving, what he understands by his ‘vegetabile medullare’ and ‘corticale’; that he does not for a moment mean a plant of simplest organisation, but that both expressions indicate only the original elements of vegetation which the Creator, according to Linnaeus, united to one another at the first. He assumed that plants of the highest and of the lowest grades of organisation were originally created at the same time and alongside of one another; no new class-plants were afterwards created, but from the mingling together of the existing ones by the act of the Creator generically distinct forms were produced, and the natural mingling of these gave birth to species, while varieties were mere chance deviations from species. But it is to be noticed that in these minglings or hybridisations the woody substance of the one form which supplies the pollen is united with the pith-substance of the other form, whose pistil is thus fertilised; and so in these supposed crossings it is always the two original elements of the plant, the medullary and the cortical, which are mingled together.
No further proof is wanting that this theory of Linnaeus is no precursor of our theory of descent, but is most distinctly opposed to it; it is utterly and entirely the fruit of scholasticism, while the essential feature in Darwin’s theory of descent is that scholasticism finds no place in it.
[CHAPTER III.]
Development of the Natural System under the Influence of the Dogma of the Constancy of Species.
1759-1850.
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[34] 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[35] (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.
| CLASS. | ||||||||
| Acotyledones | I. | |||||||
| Monocotyledones | ||||||||
| Stamina | ||||||||
| hypogyna | II. | |||||||
| perigyna | III. | |||||||
| epigyna | IV. | |||||||
| Dicotyledones. | ||||||||
| Apetalae | Stamina | |||||||
| hypogyna | V. | |||||||
| perigyna | VI. | |||||||
| epigyna | VII. | |||||||
| Monopetalae | Corolla | |||||||
| hypogyna | VIII. | |||||||
| perigyna | IX. | |||||||
| epigyna | ||||||||
| antheris | ||||||||
| connatis | X. | |||||||
| distinctis | XI. | |||||||
| Polypetalae | Stamina | |||||||
| epigyna | XII. | |||||||
| hypogyna | XIII. | |||||||
| perigyna | XIV. | |||||||
| Diclines irregulares | XV. | |||||||
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.
- Class I.
- 1. Fungi.
- 2. Algae.
- 3. Hepaticae.
- 4. Musci.
- 5. Filices.
- 6. Naiades.
- Class II.
- 7. Aroideae.
- 8. Typhae.
- 9. Cyperoideae.
- 10. Gramineae.
- Class III.
- 11. Palmae.
- 12. Asparagi.
- 13. Junci.
- 14. Lilia.
- 15. Bromeliae.
- 16. Asphodeli.
- 17. Narcissi.
- 18. Irides.
- Class IV.
- 19. Musae.
- 20. Cannae.
- 21. Orchides.
- 22. Hydrocharides.
- Class V.
- 23. Aristolochiae.
- Class VI.
- 24. Elaeagni.
- 25. Thymeleae.
- 26. Proteae.
- 27. Lauri.
- 28. Polygoneae.
- 29. Atriplices.
- Class VII.
- 30. Amaranthi.
- 31. Plantagines.
- 32. Nyctagines.
- 33. Plumbagines.
- Class VIII.
- 34. Lysimachiae.
- 35. Pediculares.
- 36. Acanthi.
- 37. Jasmineae.
- 38. Vitices.
- 39. Labiatae.
- 40. Scrophulariae.
- 41. Solaneae.
- 42. Borragineae.
- 43. Convolvuli.
- 44. Polemonia.
- 45. Bignoniae.
- 46. Gentianeae.
- 47. Apocyneae.
- 48. Sapotae.
- Class IX.
- 49. Guajacanae.
- 50. Rhododendra.
- 51. Ericae.
- 52. Campanulaceae.
- Class X.
- 53. Cichoraceae.
- 54. Cinarocephalae.
- 55. Corymbiferae.
- Class XI.
- 56. Dipsaceae.
- 57. Rubiaceae.
- 58. Caprifolia.
- Class XII.
- 59. Araliae.
- 60. Umbelliferae.
- Class XIII.
- 61. Ranunculaceae.
- 62. Papaveraceae.
- 63. Cruciferae.
- 64. Capparides.
- 65. Sapindi.
- 66. Acera.
- 67. Malpighiae.
- 68. Hyperica.
- 69. Guttiferae.
- 70. Aurantia.
- 71. Meliae.
- 72. Vites.
- 73. Gerania.
- 74. Malvaceae.
- 75. Magnoliae.
- 76. Anonae.
- 77. Menisperma.
- 78. Berberides.
- 79. Tiliaceae.
- 80. Cisti.
- 81. Rutaceae.
- 82. Caryophylleae.
- Class XIV.
- 83. Sempervivae.
- 84. Saxifragae.
- 85. Cacti.
- 86. Portulaceae.
- 87. Ficoideae.
- 88. Onagrae.
- 89. Myrti.
- 90. Melastomae.
- 91. Salicariae.
- 92. Rosaceae.
- 93. Leguminosae.
- 94. Terebinthaceae.
- 95. Rhamni.
- Class XV.
- 96. Euphorbiae.
- 97. Cucurbitaceae.
- 98. Urticae.
- 99. Amentaceae.
- 100. Coniferae.
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[36] ‘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 Schmeidel’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[37] (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 (adhérence). 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 flower-stalks 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 subdivisions 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).
- 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[38] (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 coordination 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[39], 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[40] 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; (1) 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[41]. 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 pre-eminent; 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.
[CHAPTER IV.]
Morphology under the Influence of the Doctrine of Metamorphosis and of the Spiral Theory.
1790-1850.
The efforts of Jussieu, De Candolle, and Robert Brown were directed to the discovery of the relationship between different species of plants by comparing them together; the doctrine of metamorphosis founded by Goethe set itself from the first to bring to light the hidden relationship between the different organs of one and the same plant. As De Candolle’s doctrine of symmetry derived the different species of plants from an ideal plan of symmetry or type, so the doctrine of metamorphosis assumed an ideal fundamental organ, from which the different leaf-forms in a plant could be derived. The stem came into consideration only as carrying the leaves, the root was almost entirely disregarded. As the resemblance of nearly allied species of plants suggests itself naturally and unsought to the mind of the unbiassed observer, so also does the connection between different organs of a leafy nature in one and the same plant. Cesalpino called the corolla simply a ‘folium’ (leaf); he and Malpighi regarded the cotyledons also as leaves; Jung called attention to the variety of the leaf-forms, which are found in many plants at different heights on the same stem; Caspar Friedrich Wolff, the first who bestowed systematic consideration on the subject, declared in 1766, that he saw nothing ultimately in the plant but leaves and stem, including the root in the stem[42].
Long before Goethe’s time speculation had busied itself with attempts to explain these observations; we saw how Cesalpino and Linnaeus, starting from the old view that the pith is the seat of the soul in plants, regarded the seeds as metamorphosed pith, the floral envelopes with the stamens and the true leaves as metamorphosed layers of the rind and wood of the stem. The word metamorphosis from their point of view had a very plain meaning; it was really the cylindrical pith whose upper end changed into seeds, it was the actual substance of the cortex which produced both the ordinary leaves and the parts of the flower. Wolff on the other hand from a point of view of his own gave an apparently intelligible physical explanation of the proposition, that all appendages of the stem are leaves, but the explanation had the fault of not being true; he attributed the metamorphosis of leaves to altered nourishment, the flowers especially to his ‘vegetatio languescens.’
Goethe’s conception of the matter was from the first much less clear, and chiefly because he was never able to bring the abnormal into its true connection with the normal or ascending metamorphosis. In the first sentence of his ‘Doctrine of metamorphosis’ (1790) he says, ‘that it is open to observation that certain exterior parts of plants sometimes change and pass into the form of adjacent parts, either wholly or in a greater or less degree.’ In the cases of which Goethe is here thinking a distinct meaning can be affixed to the word metamorphosis; if, for example, the seeds of a plant with normal flowers produce a plant which has petals in place of stamens, or in which the ovaries are resolved into green expanded leaves, it is actually the case that a plant of a known form has given rise to another plant of a different form, in other words, a change or metamorphosis has really taken place. But we cannot reason in this way in the case of that which Goethe calls normal or ascending metamorphosis. When in a given species, which has remained constant with all its marks for countless generations, the cotyledons, the leaves, the bracts, and the parts of the flower are called leaves, this must be merely the result of abstraction, which has led to the generalising of the idea of a leaf; if we make abstraction of the physiological characters of the carpels, stamens, floral envelopes, and cotyledons, and regard only the way in which they originate on the stem, we are justified in including them in one general idea with ordinary leaves, and to this idea we quite arbitrarily give the name leaf. But this does not justify us in speaking of a change of these organs, so long as we consider the whole plant in question as a hereditary and constant form. For the plant therefore taken as constant the idea of metamorphosis has only a figurative meaning; the abstraction performed by the mind is transferred to the object itself, if we ascribe to it a metamorphosis which has really taken place only in our conception. The case would be different, if here as well as in the abnormal instances above-mentioned we could assume that the stamens and other organs of the plants lying before us were ordinary leaves in their progenitors. So long as this assumption of an actual change is not even hypothetically made, the expression change or metamorphosis is purely figurative, the metamorphosis is a mere ‘idea.’ This distinction Goethe has not made; he did not clearly see that his normal ascending metamorphosis can only have the meaning of a scientific fact, if a real change is assumed to take place in the course of propagation in this case, as in that of abnormal metamorphosis or misformation. A comparison of his various expressions shows that he took the word metamorphosis sometimes in its literal, sometimes in its ideal and figurative sense; for instance, he says expressly, ‘We may say that a stamen is a folded petal, just as we may say that a petal is a stamen in a state of expansion.’ This sentence shows that Goethe did not regard a particular leaf-form as first in time, and that others proceeded from it by change; he uses the word metamorphosis in a purely ideal sense. At other times his remarks may be interpreted as though he really considered the normal ascending metamorphosis to be a real change in the organs, arising from a transmutation of the species. With this confusion of notion and thing, idea and reality, subjective conception and objective existence, Goethe took up exactly the position of the so-called nature-philosophy.
Goethe’s doctrine could only make its way to logical consistency and clearness of thought by deciding for the one or the other way; he must either assume that the different leaf-forms, which were regarded as alike only in the idea, were really produced by change of a previous form,—a conception that at once presupposes a change of species in time; or he must entirely adopt the position of the idealistic philosophy, in which idea and reality coincide. In this case the assumption of a change in time was not necessary; the metamorphosis remained an ideal one, a mere mode of view; the word leaf then signifies only an ideal fundamental form from which the different forms of leaves actually observed may be derived, as De Candolle’s constant species from an ideal type.
If now we read Goethe’s further remarks on the doctrine of metamorphosis attentively[43], we perceive that he really arrived at neither of these conclusions, but perpetually vacillated between the two; a number of his sayings might be collected, which might be taken for precursors of a theory of descent, as they have been taken by some modern writers; but it is quite as easy to make a selection which would carry us back to the position of the ideal philosophy and the constancy of species. In the later years of his life the idea of a physical metamorphosis accomplished in time, and involving a change of species, does appear more distinctly in Goethe’s writings. This explains the lively, nay passionate, interest which he took in the dispute between Cuvier and Geoffrey de St. Hilaire in 1830[44]. We gather from it that Goethe, in spite of all his wanderings in the mists of the nature-philosophy of the time, felt a growing need for some clearer insight into the nature of metamorphosis, both in plants and animals, without ever being able to make his way into the clear light.
But these better motions remained without importance for the history of botany; the adherents of his doctrine of metamorphosis all apprehended it in the sense of the nature-philosophy, and Goethe himself did not remonstrate against the frightful way in which it was distorted by them. Its further development therefore was in accordance with the principles of that philosophy, which was accustomed to apply the results of purely idealistic views in an uncritical way to imperfectly observed facts. Above all the difficulty remained unsolved, how the dogma of the constancy of species was to be brought into logical connection with the idea of the metamorphosis of organs. The supranatural, which Elias Fries found in the natural system, subsisted still in the doctrine of metamorphosis in comparing the organs of a plant.
Still more obscure and entirely the product of the nature-philosophy is Goethe’s view of the spiral tendency in vegetation. At p. 194 of his essay entitled ‘Spiraltendenz der Vegetation’ (1831) he says: ‘Having fully grasped the idea of metamorphosis we next turn our attention to the vertical tendency, in order to gain a nearer acquaintance with the development of the plant. This tendency must be looked upon as an immaterial staff, which supports the existence.... This principle of life (!) manifests itself in the longitudinal fibres which we use as flexible threads for many purposes; it is this which forms the wood in trees, which keeps annual and biennial plants erect, and even produces the extension from node to node in climbing and creeping plants. Next we have to observe the spiral direction which winds round the other.’ This spiral direction which passes at once with Goethe into a ‘spiral tendency,’ is seen in various phenomena of vegetation, as in spiral vessels, in twining stems, and sometimes in the position of leaves. The closing remarks of this short essay, in which he explains the vertical tendency as the male, the spiral as the female principle in the plant, show how far Goethe lost himself in the profundities of the nature-philosophy. Thus he introduced his readers into the deepest depths of mysticism.
It would be as useless as it would be wearisome to follow out in detail to its extremest point of absurdity the progressive transformation which the doctrine of metamorphosis underwent in the hands of the botanists of the nature-philosophy school, and to see how its catchwords, polarity, contraction and expansion, the stem-like and the fistular, anaphytosis and life-nodes, and others, were compounded with the results of the most every-day observation into meaningless conglomerates; rough obscure impressions of the sense, as well as incidental fancies, were regarded as ideas and principles. A full account of these inconceivable aberrations is to be found in Wigand’s ‘Geschichte und Kritik der Metamorphose.’ Our own countrymen certainly, Voigt, Kieser, Nees von Esenbeck, C. H. Schulz, and Ernst Meyer (the historian of botany) bear off the palm of absurdity, but there were others also, among them the Swedish botanist Agardh, and some Frenchmen, Turpin, for instance, and Du Petit-Thouars[45], who were not altogether free from this weakness. Even the best German botanists of the time, such as Ludolph Treviranus, Link, G. W. Bischoff, and others, managed to escape the influence of this philosophy of nature, only where they confined themselves to the most barren empiricism. Strange phenomenon! that as soon as gifted and understanding men began to talk of the metamorphosis of plants, they fell into senseless phrase-mongering; Ernst Meyer, for instance, was it is true no great botanist, but he shows in his ‘Geschichte der Botanik’ that he possessed a clever and cultivated intellect. The painful impression, which the treatment of the doctrine of metamorphosis by these writers makes upon us, is due partly to the fact that the deeper meaning of the idealistic philosophy never attained to logical expression in their hands, and still more to their indulgence in an unmeaning play of phrases, combining the highest abstractions with the most negligent and rudest empiricism, and sometimes with utterly incorrect observations. Oken can claim the merit of more correct observation and greater philosophical consistency, and if we reject his views, yet his mode of presenting them has at least the pleasing appearance of more consequential reasoning. We perceive for the first time the full greatness of the debt which modern botany owes to men like Pyrame de Candolle, Robert Brown, von Mohl, Schleiden, Nägeli, and Unger, the latter of whom only slowly worked his way out of the trammels of the nature-philosophy, when we compare the literature of the doctrine of metamorphosis before the year 1840 with the present condition of our science, for which they paved the way.
In spite of the real and apparent differences between Goethe’s doctrine of metamorphosis and De Candolle’s doctrine of a plan of symmetry, these writers agreed in this, that they set out alike from the doctrine of the constancy of species, and led up equally to the result, that alongside of manifold physiological differences in the organs of plants certain points of formal agreement can be discovered, which are expressed chiefly in the order of their succession and in their relative positions. In this distinction lay the good kernel of the doctrine of metamorphosis in Goethe, and Wolff, and even in Linnaeus and Cesalpino: it was only necessary to set this free from the dross with which the nature-philosophy had surrounded it, and to make the relations of position in organs the subject of earnest investigation, in order to secure important results in this branch of morphology. The first step in this direction was taken by Carl Friedrich Schimper, who was followed by Alexander Braun; both adopted the main idea of the doctrine of metamorphosis in the form in which it can be reconciled with the doctrine of constancy, that is, in a purely idealistic sense. Both liberated themselves from the gross errors of the nature-philosophers, and thus gave a more logical expression to the purely idealistic morphological consideration of form in plants.
Karl Friedrich Schimper[46] founded before the year 1830 the theory of the arrangement of leaves which is named after him, and which he expounded to the naturalists assembled at Stuttgart in 1834 as a complete and perfected system. Alexander Braun, in a review of Schimper’s exposition in ‘Flora’ of 1835, gave a clear and simple account of the theory, having already himself published an excellent and comprehensive treatise on the same subject. The doctrine of phyllotaxis appeared in these publications with a formal completeness which could not fail to attract the attention of the botanical world and indeed of a larger audience; and justly so, for, as unfortunately so very seldom happens in botanical subjects, a scientific idea was in this case not merely incidentally suggested, but was worked out in all its consequences as a complete structure, and this structure gained in external splendour from the circumstance that its propositions, dealing with geometrical constructions, could be expressed in numbers and formulae,—a thing hitherto unknown in botanical science.
That the leaves are arranged on the stems that produce them according to fixed geometrical rules had been noticed by Cesalpino and by Bonnet in the middle of the eighteenth century; but nothing more resulted than weak attempts at mere description of different cases. Schimper’s theory is marked by that which is at once its greatest merit and its fundamental error, the referring of all relations of position to a single principle. This principle lies in the idea that growth in a stem has an upward direction in a spiral line, and that the formation of leaves is a local exaggeration of this spiral growth. The direction of the spiral line may change in the same species, or in the same axis, and may even change from leaf to leaf. The important variations in the arrangement of leaves are not shown in their longitudinal distances, but in the measure of their lateral deviations on the stem. The characteristic point in this theory is the mode of considering these lateral deviations or divergences of the leaves as they follow one another on an axis, the referring them to a more general law of position. Means were at the same time skilfully supplied for discovering the true conditions of arrangement, the genetic spiral, in cases where the genetic succession of the leaves, and consequently their divergence, could not be immediately recognised. After innumerable observations, it appeared that there is a wonderful variety in the disposition of leaves, but that at the same time a comparatively small number of these variations commonly occur, and that these ordinary divergences 1/2, 2/3, 3/8, 8/13, 13/21, etc. have this remarkable relation to one another, that both the numerator and denominator of each successive fraction are obtained by adding together the numerators and denominators of the two preceding fractions, or the individual fractions named are the successive convergents of a continuous fraction:—
- 1
- 1 + 1
- ——-
- 1 + 1
- ——-
- 1....
By change of single cyphers in this, the simplest of all continuous fractions, the expressions were also obtained for all measures of position that deviate from the usual main series. The common occurrence of so-called leaf-whorls seemed at once to be opposed to the principle of special growth and to the doctrine of position founded upon it, especially in the cases in which it was supposed that all the leaves of a whorl arise simultaneously. But the founders of the doctrine, relying on their geometrical constructions, declared that every theory is incorrect, which sets out from the whorl as a simultaneous formation. But the way in which the different leaf-whorls of a stem are arranged among themselves, and are connected with continuous spiral positions, required new geometrical constructions; it was necessary to assume a supplementary relation (prosenthesis), which the measure of the phyllotaxis adopts in the transition from the last leaf of one cycle to the first of the next. Artificial as this construction appears, it has the advantage of saving the spiral principle, and the prosenthetic relation itself admits of being again expressed in highly simple fractions,—a great advantage for the formal consideration of the relative positions of the parts of the flower, and their relation to the preceding positions of the leaves. The great skill shown by the founders of the doctrine in the morphological consideration of the whole plant-form appears equally in the establishment of the rules, according to which the relations of position of the leaves of a side-shoot connect with those of the mother-axis, and which made it possible to represent the nature of inflorescences especially with extreme clearness by means of geometrical figures. An expressive and elegant terminology not only made the whole theory attractive, but fitted it in a high degree to supply a suitable, plain, and precise phraseology for describing the most varied forms of plants. That the theory possesses such advantages as these may be gathered from the fact, that since 1835 the morphological examination and comparison not only of flowers and inflorescences, but also of vegetative shoots and their ramification, has reached great formal completeness. A thorough acquaintance with the principle of this doctrine has made it possible to explain to reader or hearer the most intricate forms of plants so clearly, that they may be said to reveal the law of their formation themselves, and to grow before the eye of the observer, while at the same time the most recondite relations of the organs of the same or of different plants were brought out distinctly and in elegant phraseology. When this mode of description was combined with De Candolle’s views on abortion, degeneration, and adherence, and at the same time took into consideration the chief physiological forms of leaf-structures, according as these were developed as scales, foliage-leaves, bracts, floral envelopes, staminal and carpellary leaves, it was possible to give such an artistic account of every form of plant, as made it visible to sense in its entirety, and at the same time brought out the morphological law of its construction. Whoever reads the writings of Alexander Braun and Wydler, and especially of Thilo Irmisch (after 1873), who knew how to combine his descriptions in a variety of ways with remarks on the biological relations of plants, cannot fail to admire the extraordinary skill displayed by these men in describing plants. Compared with the dry diagnoses of the systematists, their descriptions attain to the dignity of an art, and present the commonest forms to the reader in a new and attractive light. But the theory had a further advantage; it seemed not only to present the form of the plant in its matured state, but to treat it genetically; and in fact it did possess an element of historical development, inasmuch as it made the genetic succession of the leaves and of their axillary shoots, which is at the same time the succession from the base to the summit, the foundation of all consideration of the plant-form. But it is also true that in this lay one of the weak sides of the theory; as long as it was a question only of continuous spirals, the succession of matured leaves does also represent the succession of their formation in time; but this was not actually proved in the case of leaf-whorls, and here, to save the theory, genetic relations had to be presupposed for which no further proof was forthcoming, while fresh researches have repeatedly shown that a strict application of Schimper’s theory is found frequently to contradict the facts of development as directly observed[47]. Moreover, regard was had only to those measurements of divergence on the continuous genetic spiral which were taken on the matured stem, while there was always the possibility that the divergences might have been different at the first, and been afterwards modified, as Nägeli subsequently suggested[48]. And again, the theory had a dangerous adversary to encounter in the frequent occurrence of leaves that are strictly alternate or crossed in pairs, and to conceive of this as a spiral arrangement must at once appear to be an arbitrary proceeding both from the mathematical point of view and from that of historical development; the assumption of a return of the genetic spiral from leaf to leaf, as for instance in the Grasses, like the prosenthesis in the change of divergence, afforded, it is true, a construction which was geometrically correct, but which could hardly be made to agree with the history of development and the mechanical forces concerned. Again, it was a great and essential defect in the theory, that in assuming the spiral arrangement it entirely neglected the relations of symmetry of the plant-form, which are in many cases clearly expressed, and their connection with the outer world, on which Hugo von Mohl had already published some excellent remarks in 1836,—a defect, which unhappily is not yet sufficiently appreciated. A due consideration of these objections, and of the cases in which the history of development is opposed to the constructions of the theory, must have led to the conviction that the idea of a spiral tendency in the growth of plants is at least not borne out in all cases, and more profound reflexion would show, that a scientific principle, really explaining the phenomena, is no more to be found in the assumption of such a general tendency, than in a like assumption with regard to the heavenly bodies, that they have a tendency to elliptic movement because they commonly move in ellipses. Hence Hofmeister, the latest investigator of the doctrine of phyllotaxis on the basis of the history of development, comes to the conclusion that the notion of a screw-shaped or spiral course of evolution of lateral members of plants is not merely an unsuitable hypothesis, but an error. Its unreserved abandonment is, he considers, the first condition for attaining an insight into the proximate causes of the varieties of relative position in the vegetable kingdom. But this judgment, correct as it is, was pronounced thirty years after the appearance of Schimper’s theory; history, which speaks from another point of view, and not only enquires into the correctness of a theory but has to appraise its historical importance, speaks in a less unfavourable manner. The chief point here is not whether the theory was right, but how far it contributed to the advance of the science. It was distinctly fruitful in results, for it brought the important question of the relative positions of organs for the first time into the front rank in the study of morphology; we may even say that a large part of the results of the study of the history of development were first brought into the true light by the consistent application of the theory, or in the effort to disprove it. With all its fundamental errors, Schimper’s theory remains one of the most interesting phenomena in the history of morphology, because it was carried out with thorough logical consistency. We should as little wish to omit it from our literature, as modern astronomy would wish to see the old theory of epicycles disappear from its history. Both theories served to connect together the facts that were known in their time.
The fundamental error of the theory lies much deeper than appears at first sight. Here too we have the idealistic conception of nature, which refuses to know anything of the causal nexus, because it takes organic forms for the ever-recurring copies of eternal ideas, and in accordance with this platonic sphere of thought confounds the abstractions of the mind with the objective existence of things. This confusion shows itself in Schimper’s doctrine, inasmuch as he takes the geometrical constructions, which he transfers to his plants and which, though they may be highly suitable from his point of view, are nevertheless purely arbitrary, for actual characters of the plants themselves, in other words, takes the subjective connection of the leaves by a spiral line for a tendency inherent in the nature of the plant. Schimper in making his constructions overlooked the fact that, because a circle can be described by turning a radius round one of its extremities, it does not follow that circular surfaces in nature must really have been formed in this way; in other words, he did not see that the geometrical consideration of arrangements in space, useful as it may otherwise be, gives no account of the causes to which they are due. But this was not properly an oversight in Schimper’s case, for he would have scarcely admitted efficient causes in the true scientific sense into his explanations of the form of plants. How far Schimper was from regarding plants as something coming into being in time and according to natural laws, how profoundly he despised the principles of modern natural science is shown in his judgment of Darwin’s theory of descent and of the modern atomic theory, the coarseness of which is the more surprising, because Schimper was a man of refined and even poetic feeling. ‘Darwin’s doctrine of breeding,’ he says, ‘is, as I discovered at once and could not help perceiving more and more after repeated and careful perusal, the most shortsighted possible, most stupidly mean and brutal, much more paltry even than that of the tesselated atoms with which a modern buffoon and hired forger has tried to entertain us.’ Here is the old platonic view of nature flying at modern science; the sternest ‘opposites’ that culture has ever produced.
The theory of Schimper, which should rather be called the theory of Schimper and Braun, considering the active part which Braun took from the first in framing and applying it, was capable of further development only in the mathematical and formal direction, as was shown especially in Naumann’s essay, ‘Ueber den Quincunx als Grundgesetz der Blattstellung vieler Pflanzen’ (1845). The defects above described, but not the merits of the theory were shared by the doctrine of phyllotaxis laid down about ten years later by the brothers Louis and Auguste Bravais. Their theory makes use of mathematical formulae to even a greater extent than that of Schimper without paying any attention to genetic conditions, and yet it is less consistent with itself, for it assumes two thoroughly different kinds of phyllotaxis, the positions in which are arranged in a straight and in a curved line; for the latter without any apparent reason a purely ideal original divergence is assumed which stands in irrational relation to the circumference of the stem, and from it all other divergences should be derivable; and this ultimately degenerates into mere playing with figures which in this form afford no deeper insight into the causes of the relations of position. As regards serviceableness in the methodic description of plants the theory of the brothers Bravais is much inferior to that of Schimper[49].
The genetic morphology founded about the year 1840 had to make the best terms it could with the doctrine of phyllotaxis, which was constructed on a totally different principle; the two went their way on the whole side by side without disturbance from one another till the year 1868, when Hofmeister in his general morphology attacked the principle of Schimper’s theory, and endeavoured to substitute a genetic and mechanical explanation of the relative positions for the purely formal account of them; this attempt however, which from the nature of the case has not yet led to a finished theory but nevertheless contains the germ of a further development of this important doctrine, does not come within the scope of this history.
The doctrine of phyllotaxis of Schimper and Braun, as it appeared after 1830, had clearly presented only one side of the theory of metamorphosis; what other elements there were in it capable of being turned to speculative account were further cultivated by Alexander Braun between the years 1840 and 1860. In this period fresh points of view were asserting themselves in botanical research; the founding of the doctrine of cells, the study of the more delicate anatomy of plants and of the history of development, and increased methodical knowledge of the Cryptogams were enlarging the repertory of botanical facts, while the physico-mechanical method of investigation was being more and more adopted. Braun, who took an active part by his own researches in this revolution in morphological botany, remained true nevertheless to idealistic views; and in his frequent and comprehensive discussions of the general results of the new investigations in accordance with these views he has shown how far the idealistic platonising contemplation of nature is in a condition to do justice to the results of exact inductive enquiry. The opposition between his point of view and that of the most eminent representatives of the inductive method became more and more pronounced as years went on, and must be treated here as a historical fact. But if the new tendency in botany pursued especially by von Mohl, Schleiden, Nägeli, Unger, and Hofmeister may be called inductive in the absence of a better term, and be contrasted with the idealistic tendency represented by Braun and his school, it must not be supposed that the latter did not equally contribute in matters of detail to the enriching of the science by the method of induction; on the contrary, Braun himself was the author of a series of important works conceived in this spirit. When the new method is here called inductive, it should be understood that the word is used in a higher than the usual sense, and some explanation of this point will not be superfluous in this place. Idealistic views of nature of all times, whether they present themselves as Platonism, Aristotelian logic, Scholasticism or modern Idealism, have all of them this in common, that they regard the highest knowledge attainable by man as something already won and established; the highest axioms, the most comprehensive truths are supposed to be already known, and the task of inductive enquiry is essentially that of verifying them; the results of observation serve to elucidate already received views, to illustrate already known truths; inductive enquiry has only to establish individual facts. But in the sense in which inductive enquiry was understood by Bacon, Locke, Hume, Kant, and Lange, its task is one that goes essentially farther than this; it must not be content with establishing individual facts, but it must employ them in the critical examination of the most general notions that have come down to us, and do its best to deduce new and comprehensive theories from them, even where these may be entirely opposed to traditional views. But it is part of the very nature of this method of investigation, that its general results are subject to constant modification and improvement; each more general truth has only a temporary value, and endures as long as no new facts militate against it. The distinction therefore between idealism and the inductive method in the domain of natural science comes to this, that the former fits new facts into a scheme of old conceptions, the latter deduces new conceptions from new facts; the one is in its nature dogmatic and intolerant, the other eminently critical; the one is conservative, the other always pressing forwards; the one inclines to philosophic contemplation, the other to vigorous and productive investigation. To this must be added one point of great importance; the idealistic view of nature, rejecting causality, explains nature from notions of design, and is teleological; ethical and even theological elements are thus introduced into natural science.
It is in this form that the distinction between the idealistic view represented by Braun and the modern inductive morphology presents itself to us. If it were the task of this history only to record the discovery of new facts, it would be superfluous to allude to these differences here; but then it would also be impossible to estimate rightly that portion of Braun’s long scientific labours which is at once the most original and the most interesting from the historical point of view, and which is to be found not so much in his many descriptive and monographic works, as in his philosophic efforts in the domain of morphology; these moreover deserve our consideration, because they carry out Goethe’s half-explained conceptions to their remotest consequences, and express in purer form the idealism which lies at the foundation of the older nature-philosophy. No botanist since Cesalpino has so thoroughly endeavoured to leaven the entire results of inductive investigation with the principles of an idealistic philosophy, and to explain them in its light.
Braun’s philosophical views not only accompany his knowledge of facts, but everywhere permeate and colour it; in his writings, contributions, and monographs on the most various subjects, facts are regarded from the point of view of his philosophy. He has given a general view of his philosophical principles and illustrated them by a vast variety of facts in his famous book, ‘Betrachtungen über die Erscheinung der Verjüngung in der Natur, insbesondere in der Lebens-und Bildungsgeschichte der Pflanze’ (1849-50). He himself directs attention to the opposition between his own stand-point and the modern induction in the tenth page of the preface, where he replies to the obvious objection, that his ideas may be regarded as antiquated, in the words, ‘A more living contemplation of nature, such as is here attempted, which seeks in natural bodies not merely the operation of dead forces, but the expression of a living fact, does not lead, as is supposed, to airy structures of fancy, for it does not pretend to gain a knowledge of life in nature in any other way than as it is revealed in phenomena,’ etc. This thought is still more distinctly uttered in page 13 of the text; ‘As external nature without mankind presents to us only the spectacle of a labyrinth without a guide, so too scientific contemplation, which denies the inner spiritual principle in nature and the intimate connection of nature with the informing spirit[50], leads to a chaos of substances and forces, which are unknown because divorced from spirit, or, to speak more precisely, to a chaos of nothing but unknown causes, which work together in an inexplicable manner.’ In a note to this passage he points expressly to ‘the comfortless character of such an unreal mode of viewing nature, which must necessarily endeavour to root out everything in the conceptions and language of science which appears from its own point of view to be anthropopathic,’ and he requires a tender, ethical element as essential to botanical investigation. The chief object of the volume is to prove that everything in organic life may be resolved into rejuvenescence, of which idea no definition is actually given, though the whole contents of the book are a search after a definition. We may regard the idea of rejuvenescence, as presented by Braun, as an extension of the idea of metamorphosis, in which extended form it is adapted to take in even the results of the cell-theory, of the history of development, and of the modern knowledge of the Cryptogams from the idealistic point of view. One peculiarity of his mode of expounding his views is observed here, as on other occasions, namely, that he gives no precise and arbitrary definition to a word, for instance, like rejuvenescence in the present place, and in a later work to the word individual, but looks behind the word for a profound or even mysterious meaning, which is to be perceived and brought to light by contemplation of the phenomena. In page 5 he says, ‘Thus we see youth and age appear alternately in one and the same history of development; we see youth burst through age, and by growth or transformation step into the middle of the development. This is the phenomenon of rejuvenescence, which is repeated in endless multiplicity in every province of life, but nowhere appears more clearly expressed or more accessible to investigation than in the vegetable kingdom. Without rejuvenescence there is no history of development.’—‘If then we ask for the causes of the phenomena of rejuvenescence (page 7), we shall indeed allow that nature, into which special life enters in its various manifestations, excites, awakes, and works by the influences which the years and even the days bring with them; but the true and inner cause can only be found in the desire after perfection which belongs to every being in its kind, and urges it to bring the outer world, which is strange to it, more and more into complete subjection to itself, and to fashion itself in it as independently as its specific nature admits.’ Further on he says (page 17), ‘The impulse or tendency to development in each creature is likewise no direction of activity impressed from without, but one given from within and working as an inner determination and force from the depth of the inner nature.’ A passage also from page 111 of his treatise on polyembryony, published in 1860, may be quoted here; ‘Though the organism, in the process of realising itself, is subject to physical conditions, yet the proper causes of its morphological and biological characteristics do not lie in these conditions; its laws belong to a higher stage of development of its being, to a sphere in which the faculty of self-determination is distinctly manifested. If this is so, the laws of an organic being appear as tasks imposed, the fulfilling of which is not absolutely necessary but only in relation to the attainment of a definite end, as precepts, to which strict obedience may possibly not be paid.’ To return once more to the idea of rejuvenescence, we find at page 18 the words, ‘As regards the idea of rejuvenescence, from the foregoing considerations we draw the conclusion, that the surrender of growths already accomplished and the going back to new beginnings, the commencement of rejuvenescence, indicate only the outer side of the proceeding, while the essential part of it is an inner gathering up of forces, a new creating, as it were, out of the individual principle of life, a fresh reflecting upon the specific task or the gaining renewed hold upon the type which is to be presented in the outer organism. By this means rejuvenescence maintains its fixed relation to development, which can and ought to present in gradually attained perfection that only which lies in the nature of the creature, and is most intimately its own.’ And at the conclusion of the work (page 347) he says, ‘The way in which the inner spiritual nature of life is specially manifested in the phenomenon of rejuvenescence may be defined as reminiscence in the true sense of the word, as the power of grasping anew in the phenomenon the inner destination of life as contrasted with its daily alienation and decay, and applying it with renewed strength towards that which is without,’ etc.
This conception of rejuvenescence is, then, applied to all the phenomena of life in plants; not only the metamorphosis of leaves, the formation of shoots and their ramification, and the different modes of cell-formation, but even palaeontological facts are manifestations of rejuvenescence, which in the sequel puts off the form of an abstract idea, and becomes personified into an active personality, as is seen in page 8 in the expression, ‘activity of rejuvenescence.’
The relation of Braun’s views to the question of the constancy of species may to some extent appear doubtful; some utterances of his may be interpreted to admit a transmutation of species accomplished in the course of ages, while others are opposed to this, and it is the latter which appear to be consistent with the idealistic position. We read, for instance, at page 9, ‘The appearance, as though the like was always repeating itself in nature, is suggested when we glance back from our station in time upon the succession of former epochs. Here we find the real first beginnings of species and genera, and even of orders and classes in the vegetable and animal kingdoms; we see at the same time that more or less thorough transformations are connected with the appearance of the higher grades in the organic kingdom, so that genera and species of the old world disappear, and new ones step into their place. All this change expresses not the mere accident of convulsions, which, while they destroy, at the same time prepare new ground for the prosperity of organic nature, but rather definite laws whose action pervades all the individual detail of the development of organic life.’ On the other hand we find at the conclusion of the treatise on polyembryony, written a short time before the appearance of Darwin’s memorable work, a sentence which makes the assumption of a transmutation of species appear very doubtful; it says (page 257), ‘If we are justified in assuming a general organic connection in the history of development in plant-forms, can we imagine that the type of the Mosses and of the Ferns has come from the Algae, or vice versa, that the Alga-form owes its origin to the Mosses and Ferns?’