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.