One of the most important steps ever made in our knowledge of the nerves is, the distinction which Bichat is supposed to have established, of a ganglionic system, and a cerebral system. And we may add, to the discoveries in nervous anatomy, the remarkable one, made in our own time, that the two offices—of conducting the motive impressions from the central seat of the will to the muscles, and of propagating sensations from the surface of the body and the external organs of sense to the sentient mind—reside in two distinct portions of the nervous substance:—a discovery which has been declared67 to be “doubtless the most important accession to physiological (anatomical) knowledge since the time of Harvey.” This doctrine was first published and taught by Sir Charles Bell: after an interval of some 464 years, it was more distinctly delivered in the publications of Mr. John Shaw, Sir C. Bell’s pupil. Soon afterwards it was further confirmed, and some part of the evidence corrected, by Mr. Mayo, another pupil of Sir C. Bell, and by M. Majendie.68
I shall not attempt to explain the details of these anatomical investigations; and I shall speak very briefly of the speculations which have been suggested by the obvious subservience of the nerves to life, sensation, and volition. Some general inferences from their distribution were sufficiently obvious; as, that the seat of sensation and volition is in the brain. Galen begins his work, On the Anatomy of the Nerves, thus: “That none of the members of the animal either exercises voluntary motion, or receives sensation, and that if the nerve be cut, the part immediately becomes inert and insensible, is acknowledged by all physicians. But that the origin of the nerves is partly from the brain, and partly from the spinal marrow, I proceed to explain.” And in his work On the Doctrines of Plato and Hippocrates, he proves at 465 great length69 that the brain is the origin of sensation and motion, refuting the opinions of earlier days, as that of Chrysippus,70 who placed the hegemonic or master-principle of the soul, in the heart. But though Galen thought that the rational soul resides in the brain, he was disposed to agree with the poets and philosophers, according to whom the heart is the seat of courage and anger, and the liver the seat of love.71 The faculties of the soul were by succeeding physiologists confined to the brain; but the disposition still showed itself, to attribute to them distinct localities. Thus Willis72 places the imagination in the corpus callosum, the memory in the folds of the hemispheres, the perception in the corpus striatum. In more recent times, a system founded upon a similar view has been further developed by Gall and his followers. The germ of Gall’s system may be considered as contained in that of Willis; for Gall represents the hemispheres as the folds of a great membrane which is capable of being unwrapped and spread out, and places the different faculties of man in the different regions of this membrane. The chasm which intervenes between matter and motion on the one side, and thought and feeling on the other, is brought into view by all such systems; but none of the hypotheses which they involve can effectually bridge it over.
The same observation may be made respecting the attempts to explain the manner in which the nerves operate as the instruments of sensation and volition. Perhaps a real step was made by Glisson,73 professor of medicine in the University of Cambridge, who distinguished in the fibres of the muscles of motion a peculiar property, different from any merely mechanical or physical action. His work On the Nature of the Energetic Substance, or on the Life of Nature and of its Three First Faculties, The Perceptive, Appetitive, and Motive, which was published in 1672, is rather metaphysical than physiological. But the principles which he establishes in this treatise he applies more specially to physiology in a treatise On the Stomach and Intestines (Amsterdam, 1677). In this he ascribes to the fibres of the animal body a peculiar power which he calls Irritability. He divides irritation into natural, vital, and animal; and he points out, though briefly, the gradual differences of irritability in different organs. “It is hardly comprehensible,” says Sprengel,74 “how this 466 lucid and excellent notion of the Cambridge teacher was not accepted with greater alacrity, and further unfolded by his contemporaries.” It has, however, since been universally adopted.
But though the discrimination of muscular irritability as a peculiar power might be a useful step in physiological research, the explanations hitherto offered, of the way in which the nerves operate on this irritability, and discharge their other offices, present only a series of hypotheses. Glisson75 assumed the existence of certain vital spirits, which, according to him, are a mild, sweet fluid, resembling the spirituous part of white of egg, and residing in the nerves.—This hypothesis, of a very subtle humor or spirit existing in the nerves, was indeed very early taken up.76 This nervous spirit had been compared to air by Erasistratus, Asclepiades, Galen, and others. The chemical tendencies of the seventeenth century led to its being described as acid, sulphureous or nitrous. At the end of that century, the hypothesis of an ether attracted much notice as a means of accounting for many phenomena; and this ether was identified with the nervous fluid. Newton himself inclines to this view, in the remarkable Queries which are annexed to his Opticks. After ascribing many physical effects to his ether, he adds (Query 23), “Is not vision performed chiefly by the vibrations of this medium, excited in the bottom of the eye by the rays of light, and propagated through the solid, pellucid, and uniform capillamenta of the nerves into the place of sensation?” And (Query 24), “Is not animal motion performed by the vibrations of this medium, excited in the brain by the power of the will, and propagated from thence through the capillamenta of the nerves into the muscles for contracting and dilating them?” And an opinion approaching this has been adopted by some of the greatest of modern physiologists; as Haller, who says,77 that, though it is more easy to find what this nervous spirit is not than what it is, he conceives that, while it must be far too fine to be perceived by the sense, it must yet be more gross than fire, magnetism, or electricity; so that it may be contained in vessels, and confined by boundaries. And Cuvier speaks to the same effect:78 “There is a great probability that it is by an imponderable fluid that the nerve acts on the fibre, and that this nervous fluid is drawn from the blood, and secreted by the medullary matter.”
Without presuming to dissent from such authorities on a point of 467 anatomical probability, we may venture to observe, that these hypotheses do not tend at all to elucidate the physiological principle which is here involved; for this principle cannot be mechanical, chemical, or physical, and therefore cannot be better understood by embodying it in a fluid; the difficulty we have in conceiving what the moving force is, is not got rid of by explaining the machinery by which it is merely transferred. In tracing the phenomena of sensation and volition to their cause, it is clear that we must call in some peculiar and hyperphysical principle. The hypothesis of a fluid is not made more satisfactory by attenuating the fluid; it becomes subtle, spirituous, ethereal, imponderable, to no purpose; it must cease to be a fluid, before its motions can become sensation and volition. This, indeed, is acknowledged by most physiologists; and strongly stated by Cuvier.79 “The impression of external objects upon the me, the production of a sensation, of an image, is a mystery impenetrable for our thoughts.” And in several places, by the use of this peculiar phrase, “the me,” (le moi) for the sentient and volent faculty, he marks, with peculiar appropriateness and force, that phraseology borrowed from the world of matter will, in this subject, no longer answer our purpose. We have here to go from Nouns to Pronouns, from Things to Persons. We pass from the Body to the Soul, from Physics to Metaphysics. We are come to the borders of material philosophy; the next step is into the domain of Thought and Mind. Here, therefore, we begin to feel that we have reached the boundaries of our present subject. The examination of that which lies beyond them must be reserved for a philosophy of another kind, and for the labors of the future; if we are ever enabled to make the attempt to extend into that loftier and wider scene, the principles which we gather on the ground we are now laboriously treading.
Such speculations as I have quoted respecting the nervous fluid, proceeding from some of the greatest philosophers who ever lived, prove only that hitherto the endeavor to comprehend the mystery of perception and will, of life and thought, have been fruitless and vain. Many anatomical truths have been discovered, but, so far as our survey has yet gone, no genuine physiological principle. All the trains of physiological research which we have followed have begun in exact examination of organization and function, and have ended in wide conjectures and arbitrary hypotheses. The stream of knowledge in all such cases is 468 clear and lively at its outset; but, instead of reaching the great ocean of the general truths of science, it is gradually spread abroad among sands and deserts till its course can be traced no longer.
Hitherto, therefore, we must consider that we have had to tell the story of the failures of physiological speculation. But of late there have come into view and use among physiologists certain principles which may be considered as peculiar to organized subjects; and of which the introduction forms a real advance in organical science. Though these have hitherto been very imperfectly developed, we must endeavor to exhibit, in some measure, their history and bearing.
[2nd Ed.] [In order to show that I am not unaware how imperfect the sketch given in this work is, as a History of Physiology, I may refer to the further discussions on these subjects contained in the Philosophy of the Inductive Sciences, Book ix. I have there (Chap. ii.) noticed the successive Biological Hypotheses of the Mystical, the Iatrochemical, and Iatromathematical Schools, the Vital-Fluid School, and the Psychical School. I have (Chaps. iii., iv., v.) examined several of the attempts which have been made to analyze the Idea of Life, to classify Vital Functions, and to form Ideas of Separate Vital Forces. I have considered in particular, the attempts to form a distinct conception of Assimilation and Secretion, of Generation, and of Voluntary Motion; and I have (Chap. vi.) further discussed the Idea of Final Causes as employed in Biology.]
BEFORE we proceed to consider the progress of principles which belong to animal and human life, such as have just been pointed at, we must look round for such doctrines, if any such there be, as apply alike to all organized beings, conscious or unconscious, fixed or locomotive;—to the laws which regulate vegetable as well as animal forms and functions. Though we are very far from being able to present a 469 clear and connected code of such laws, we may refer to one law, at least, which appears to be of genuine authority and validity; and which is worthy our attention as an example of a properly organical or physiological principle, distinct from all mechanical, chemical, or other physical forces; and such as cannot even be conceived to be resolvable into those. I speak of the tendency which produces such results as have been brought together in recent speculations upon Morphology.
It may perhaps be regarded as indicating how peculiar are the principles of organic life, and how far removed from any mere mechanical action, that the leading idea in these speculations was first strongly and effectively apprehended, not by a laborious experimenter and reasoner, but by a man of singularly brilliant and creative fancy; not by a mathematician or chemist, but by a poet. And we may add further, that this poet had already shown himself incapable of rightly apprehending the relation of physical facts to their principles; and had, in trying his powers on such subjects, exhibited a signal instance of the ineffectual and perverse operation of the method of philosophizing to which the constitution of his mind led him. The person of whom we speak, is John Wolfgang Göthe, who is held, by the unanimous voice of Europe, to have been one of the greatest poets of our own, or of any time, and whose Doctrine of Colors we have already had to describe, in the History of Optics, as an entire failure. Yet his views on the laws which connect the forms of plants into one simple system, have been generally accepted and followed up. We might almost be led to think that this writer’s poetical endowments had contributed to this scientific discovery;—the love of beauty of form, by fixing the attention upon the symmetry of plants; and the creative habit of thought, by making constant developement of a familiar process.80
470 But though we cannot but remark the peculiarity of our being indebted to a poet for the discovery of a scientific principle, we must not forget that he himself held, that in making this step, he had been guided, not by his invention, but by observation. He repelled, with extreme repugnance, the notion that he had substituted fancy for fact, or imposed ideal laws on actual things. While he was earnestly pursuing his morphological speculations, he attempted to impress them upon Schiller. “I expounded to him, in as lively a manner as possible, the metamorphosis of plants, drawing on paper, with many characteristic strokes, a symbolic plant before his eyes. He heard me,” Göthe says,81 “with much interest and distinct comprehension; but when I had done, he shook his head, and said, ‘That is not Experience; that is an Idea:’ I stopt with some degree of irritation; for the point which separated us was marked most luminously by this expression.” And in the same work he relates his botanical studies and his habit of observation, from which it is easily seen that no common amount of knowledge and notice of details, were involved in the course of thought which led him to the principle of the Metamorphosis of Plants.
Before I state the history of this principle, I may be allowed to endeavor to communicate to the reader, to whom this subject is new, some conception of the principle itself. This will not be difficult, if he will imagine to himself a flower, for instance, a common wild-rose, or the blossom of an apple-tree, as consisting of a series of parts disposed in whorls, placed one over another on an axis. The lowest whorl is the calyx with its five sepals; above this is the corolla with its five petals; above this are a multitude of stamens, which may be considered as separate whorls of five each, often repeated; above these is a whorl composed of the ovaries, or what become the seed-vessels in the fruit, which are five united together in the apple, but indefinite in number and separate in the rose. Now the morphological view is 471 this;—that the members of each of these whorls are in their nature identical, and the same as if they were whorls of ordinary leaves, brought together by the shortening their common axis, and modified in form by the successive elaboration of their nutriment. Further, according to this view, a whorl of leaves itself is to be considered as identical with several detached leaves dispersed spirally along the axis, and brought together because the axis is shortened. Thus all the parts of a plant are, or at least represent, the successive metamorphoses of the same elementary member. The root-leaves thus pass into the common leaves;—these into bracteæ;—these into the sepals;—these into the petals;—these into the stamens with their anthers;—these into the ovaries with their styles and stigmas;—these ultimately become the fruit; and thus we are finally led to the seed of a new plant.
Moreover the same notion of metamorphosis may be applied to explain the existence of flowers which are not symmetrical like those we have just referred to, but which have an irregular corolla or calyx. The papilionaceous flower of the pea tribe, which is so markedly irregular, may be deduced by easy gradations from the regular flower, (through the mimoseæ,) by expanding one petal, joining one or two others, and modifying the form of the intermediate ones.
Without attempting to go into detail respecting the proofs of that identity of all the different organs, and all the different forms of plants, which is thus asserted, we may observe, that it rests on such grounds as these;—the transformations which the parts of flowers undergo by accidents of nutriment or exposure. Such changes, considered as monstrosities where they are very remarkable, show the tendencies and possibilities belonging to the organization in which they occur. For instance, the single wild-rose, by culture, transforms many of its numerous stamens into petals, and thus acquires the deeply folded flower of the double garden-rose. We cannot doubt of the reality of this change, for we often see stamens in which it is incomplete. In other cases we find petals becoming leaves, and a branch growing out of the centre of the flower. Some pear-trees, when in blossom, are remarkable for their tendencies to such monstrosities.82 Again, we find that flowers which are usually irregular, occasionally become regular, and conversely. The common snap-dragon (Linaria vulgaris) affords a curious instance of this.83 The usual form of this plant is “personate,” the corolla being divided into two lobes, which differ in form, and 472 together present somewhat the appearance of an animal’s face; and the upper portion of the corolla is prolonged backwards into a tube-like “spur.” No flower can be more irregular; but there is a singular variety of this plants termed Peloria, in which the corolla is strictly symmetrical, consisting of a conical tube, narrowed in front, elongated behind into five equal spurs, and containing five stamens of equal length, instead of the two unequal pairs of the didynamous Linaria. These and the like appearances show that there is in nature a capacity for, and tendency to, such changes as the doctrine of metamorphosis asserts.
Göthe’s Metamorphosis of Plants was published 1790: and his system was the result of his own independent course of thoughts. The view which it involved was not, however, absolutely new, though it had never before been unfolded in so distinct and persuasive a manner. Linnæus considered the leaves, calyx, corolla, stamens, each as evolved in succession from the other; and spoke of it as prolepsis or anticipation,84 when the leaves changed accidentally into bracteæ, these into a calyx, this into a corolla, the corolla into stamens, or these into the pistil. And Caspar Wolf apprehended in a more general manner the same principle. “In the whole plant,” says he,85 “we see nothing but leaves and stalk;” and in order to prove what is the situation of the leaves in all their later forms, he adduces the cotyledons as the first leaves.
Göthe was led to his system on this subject by his general views of nature. He saw, he says,86 that a whole life of talent and labor was requisite to enable any one to arrange the infinitely copious organic forms of a single kingdom of nature. “Yet I felt,” he adds, “that for me there must be another way, analogous to the rest of my habits. The appearance of the changes, round and round, of organic creatures had taken strong hold on my mind. Imagination and Nature appeared to me to vie with each other which could go on most boldly yet most consistently.” His observation of nature, directed by such a thought, led him to the doctrine of the metamorphosis.
In a later republication of his work (Zur Morphologie, 1817,) he gives a very agreeable account of the various circumstances which affected the reception and progress of his doctrine. Willdenow87 quoted 473 him thus:—“The life of plants is, as Mr. Göthe very prettily says, an expansion and contraction, and these alternations make the various periods of life.” “This ‘prettily,’” says Göthe, “I can be well content with, but the ‘egregie,’ of Usteri is much more pretty and obliging.” Usteri had used this term respecting Göthe in an edition of Jussieu.
The application of the notion of metamorphosis to the explanation of double and monstrous flowers had been made previously by Jussieu. Göthe’s merit was, to have referred to it the regular formation of the flower. And as Sprengel justly says,88 his view had so profound a meaning, made so strong an appeal by its simplicity, and was so fruitful in the most valuable consequences, that it was not to be wondered at if it occasioned further examination of the subject; although many persons pretend to slight it. The task of confirming and verifying the doctrine by a general application of it to all cases,—a labor so important and necessary after the promulgation of any great principle,—Göthe himself did not execute. At first he collected specimens and made drawings with some such view,89 but he was interrupted and diverted to other matters. “And now,” says he, in his later publication, “when I look back on this undertaking, it is easy to see that the object which I had before my eyes was, for me, in my position, with my habits and mode of thinking, unattainable. For it was no less than this: that I was to take that which I had stated in general, and presented to the conception, to the mental intuition, in words; and that I should, in a particularly visible, orderly, and gradual manner, present it to the eye; so as to show to the outward sense that out of the germ of this idea might grow a tree of physiology fit to overshadow the world.”
Voigt, professor at Jena, was one of the first who adopted Göthe’s view into an elementary work, which he did in 1808. Other botanists labored in the direction which had thus been pointed out. Of those who have thus contributed to the establishment and developement of the metamorphic doctrine. Professor De Candolle, of Geneva, is perhaps the most important. His Theory of Developement rests upon two main principles, abortion and adhesion. By considering some parts as degenerated or absent through the abortion of the buds which might have formed them, and other parts as adhering together, he holds that all plants may be reduced to perfect symmetry: and the actual and constant occurrence of such incidents is shown beyond 474 all doubt. And thus the snap-dragon, of which we have spoken above, is derived from the Peloria, which is the normal condition of the flower, by the abortion of one stamen, and the degeneration of two others. Such examples are too numerous to need to be dwelt on.
The doctrine, being thus fully established, has been applied to solve different problems in botany; for instance, to explain the structure of flowers which appear at first sight to deviate widely from the usual forms of the vegetable world. We have an instance of such an application in Mr. Robert Brown’s explanation of the real structure of various plants which had been entirely misunderstood: as, for example, the genus Euphorbia. In this plant he showed that what had been held to be a jointed filament, was a pedicel with a filament above it, the intermediate corolla having evanesced. In Orchideæ (the orchis tribe), he showed that the peculiar structure of the plant arose from its having six stamens (two sets of three each), of which five are usually abortive. In Coniferæ (the cone-bearing trees), it was made to appear that the seed was naked, while the accompanying appendage, corresponding to a seed-vessel, assumed all forms, from a complete leaf to a mere scale. In like manner it was proved that the pappus, or down of composite plants (as thistles), is a transformed calyx.
Along with this successful application of a profound principle, it was natural that other botanists should make similar attempts. Thus Mr. Lindley was led to take a view90 of the structure of Reseda (mignonette) different from that usually entertained; which, when published, attracted a good deal of attention, and gained some converts among the botanists of Germany and France. But in 1833, Mr. Lindley says, with great candor, “Lately, Professor Henslow has satisfactorily proved, in part by the aid of a monstrosity in the common Mignonette, in part by a severe application of morphological rules, that my hypothesis must necessarily be false.” Such an agreement of different botanists respecting the consequences of morphological rules, proves the reality and universality of the rules.
We find, therefore, that a principle which we may call the Principle of Developed and Metamorphosed Symmetry, is firmly established 475 and recognized, and familiarly and successfully applied by botanists. And it will be apparent, on reflection, that though symmetry is a notion which applies to inorganic as well as to organic things, and is, in fact, a conception of certain relations of space and position, such developement and metamorphosis as are here spoken of, are ideas entirely different from any of those to which the physical sciences have led us in our previous survey; and are, in short, genuine organical or physiological ideas;—real elements of the philosophy of life.
We must, however imperfectly, endeavor to trace the application of this idea in the other great department of the world of life; we must follow the history of Animal Morphology.
~Additional material in the 3rd edition.~
THE most general and constant relations of the form of the organs, both in plants and animals, are the most natural grounds of classification. Hence the first scientific classifications of animals are the first steps in animal morphology. At first, a zoology was constructed by arranging animals, as plants were at first arranged, according to their external parts. But in the course of the researches of the anatomists of the seventeenth century, it was seen that the internal structure of animals offered resemblances and transitions of a far more coherent and philosophical kind, and the Science of Comparative Anatomy rose into favor and importance. Among the main cultivators of this science91 at the period just mentioned, we find Francis Redi, of Arezzo; Guichard-Joseph Duvernay, who was for sixty years Professor of Anatomy at the Jardin du Roi at Paris, and during this lapse of time had for his pupils almost all the greatest anatomists of the greater part of the eighteenth century; Nehemiah Grew, secretary to the Royal Society of London, whose Anatomy of Plants we have already noticed.
But Comparative Anatomy, which had been cultivated with ardor 476 to the end of the seventeenth century, was, in some measure, neglected during the first two-thirds of the eighteenth. The progress of botany was, Cuvier sagaciously suggests,92 one cause of this; for that science had made its advances by confining itself to external characters, and rejecting anatomy; and though Linnæus acknowledged the dependence of zoology upon anatomy93 so far as to make the number of teeth his characters, even this was felt, in his method, as a bold step. But his influence was soon opposed by that of Buffon, Daubenton, and Pallas; who again brought into view the importance of comparative anatomy in Zoology; at the same time that Haller proved how much might be learnt from it in Physiology. John Hunter in England, the two Monros in Scotland, Camper in Holland, and Vicq d’Azyr in France, were the first to follow the path thus pointed out. Camper threw the glance of genius on a host of interesting objects, but almost all that he produced was a number of sketches; Vicq d’Azyr, more assiduous, was stopt in the midst of a most brilliant career by a premature death.
Such is Cuvier’s outline of the earlier history of comparative anatomy. We shall not go into detail upon this subject; but we may observe that such studies had fixed in the minds of naturalists the conviction of the possibility and the propriety of considering large divisions of the animal kingdom as modifications of one common type. Belon, as early as 1555, had placed the skeleton of a man and a bird side by side, and shown the correspondence of parts. So far as the case of vertebrated animals extends, this correspondence is generally allowed; although it required some ingenuity to detect its details in some cases; for instance, to see the analogy of parts between the head of a man and a fish.
In tracing these less obvious correspondencies, some curious steps have been made in recent times. And here we must, I conceive, again ascribe no small merit to the same remarkable man who, as we have already had to point out, gave so great an impulse to vegetable morphology. Göthe, whose talent and disposition for speculating on all parts of nature were truly admirable, was excited to the study of anatomy by his propinquity to the Duke of Weimar’s cabinet of natural history. In 1786, he published a little essay, the object of which was to show that in man, as well as in beasts, the upper jaw contains an intermaxillary bone, although the sutures are obliterated. After 1790,94 animated and impelled by the same passion for natural 477 observation and for general views, which had produced his Metamorphosis of Plants, he pursued his speculations on these subjects eagerly and successfully. And in 1795, he published a Sketch of a Universal Introduction into Comparative Anatomy, beginning with Osteology; in which he attempts to establish an “osteological type,” to which skeletons of all animals may be referred. I do not pretend that Göthe’s anatomical works have had any influence on the progress of the science comparable with that which has been exercised by the labors of professional anatomists; but the ingenuity and value of the views which they contained was acknowledged by the best authorities; and the clearer introduction and application of the principle of developed and metamorphosed symmetry may be dated from about this time. Göthe declares that, at an early period of these speculations, he was convinced95 that the bony head of beasts is to be derived from six vertebræ. In 1807, Oken published a “Program” On the Signification of the Bones of the Skull, in which he maintained that these bones are equivalent to four vertebræ); and Meckel, in his Comparative Anatomy, in 1811, also resolved the skull into vertebræ. But Spix, in his elaborate work Cephalogenesis, in 1815, reduced the vertebræ of the head to three. “Oken,” he says,96 “published opinions merely theoretical, and consequently contrary to those maintained in this work, which are drawn from observation.” This resolution of the head into vertebræ is assented to by many of the best physiologists, as explaining the distribution of the nerves, and other phenomena. Spix further extended the application of the vertebral theory to the heads of all classes of vertebrate animals; and Bojanus published a Memoir expressly on the vertebral structure of the skulls of fishes in Oken’s Isis for 1818. Geoffroy Saint-Hilaire presented a lithographic plate to the French Academy in February 1824, entitled Composition de la Tête osseuse chez l’Homme et les Animaux, and developed his views of the vertebral composition of the skull in two Memoirs published in the Annales des Sciences Naturelles for 1824. We cannot fail to recognize here the attempt to apply to the skeleton of animals the principle which leads botanists to consider all the parts of a flower as transformations of the same organs. How far the application of the principle, as here proposed, is just, I must leave philosophical physiologists to decide.