"7. The circulation in the lungs or leaves of plants is very similar to that of fish. In fish the blood, after having pa.s.sed through their gills, does not return to the heart as from the lungs of air-breathing animals, but the pulmonary vein taking the structure of an artery after having received the blood from the gills, which there gains a more florid color, distributes it to the other parts of their bodies. The same structure occurs in the livers of fish, whence we see in those animals two circulations independent of the power of the heart--viz., that beginning at the termination of the veins of the gills and branching through the muscles, and that which pa.s.ses through the liver; both which are carried on by the action of those respective arteries and veins."(6)

Darwin is here a trifle fanciful in forcing the a.n.a.logy between plants and animals. The circulatory system of plants is really not quite so elaborately comparable to that of fishes as he supposed. But the all-important idea of the uniformity underlying the seeming diversity of Nature is here exemplified, as elsewhere in the writings of Erasmus Darwin; and, more specifically, a clear grasp of the essentials of the function of respiration is fully demonstrated.

ZOOLOGY AT THE CLOSE OF THE EIGHTEENTH CENTURY

Several causes conspired to make exploration all the fashion during the closing epoch of the eighteenth century. New aid to the navigator had been furnished by the perfected compa.s.s and quadrant, and by the invention of the chronometer; medical science had banished scurvy, which hitherto had been a perpetual menace to the voyager; and, above all, the restless spirit of the age impelled the venturesome to seek novelty in fields altogether new. Some started for the pole, others tried for a northeast or northwest pa.s.sage to India, yet others sought the great fict.i.tious antarctic continent told of by tradition. All these of course failed of their immediate purpose, but they added much to the world's store of knowledge and its fund of travellers' tales.

Among all these tales none was more remarkable than those which told of strange living creatures found in antipodal lands. And here, as did not happen in every field, the narratives were often substantiated by the exhibition of specimens that admitted no question. Many a company of explorers returned more or less laden with such trophies from the animal and vegetable kingdoms, to the mingled astonishment, delight, and bewilderment of the closet naturalists. The followers of Linnaeus in the "golden age of natural history," a few decades before, had increased the number of known species of fishes to about four hundred, of birds to one thousand, of insects to three thousand, and of plants to ten thousand.

But now these sudden accessions from new territories doubled the figure for plants, tripled it for fish and birds, and brought the number of described insects above twenty thousand. Naturally enough, this wealth of new material was sorely puzzling to the cla.s.sifiers. The more discerning began to see that the artificial system of Linnaeus, wonderful and useful as it had been, must be advanced upon before the new material could be satisfactorily disposed of. The way to a more natural system, based on less arbitrary signs, had been pointed out by Jussieu in botany, but the zoologists were not prepared to make headway towards such a system until they should gain a wider understanding of the organisms with which they had to deal through comprehensive studies of anatomy. Such studies of individual forms in their relations to the entire scale of organic beings were pursued in these last decades of the century, but though two or three most important generalizations were achieved (notably Kaspar Wolff's conception of the cell as the basis of organic life, and Goethe's all-important doctrine of metamorphosis of parts), yet, as a whole, the work of the anatomists of the period was germinative rather than fruit-bearing. Bichat's volumes, telling of the recognition of the fundamental tissues of the body, did not begin to appear till the last year of the century. The announcement by Cuvier of the doctrine of correlation of parts bears the same date, but in general the studies of this great naturalist, which in due time were to stamp him as the successor of Linnaeus, were as yet only fairly begun.

V. ANATOMY AND PHYSIOLOGY IN THE NINETEENTH CENTURY

CUVIER AND THE CORRELATION OF PARTS

We have seen that the focal points of the physiological world towards the close of the eighteenth century were Italy and England, but when Spallanzani and Hunter pa.s.sed away the scene shifted to France. The time was peculiarly propitious, as the recent advances in many lines of science had brought fresh data for the student of animal life which were in need of cla.s.sification, and, as several minds capable of such a task were in the field, it was natural that great generalizations should have come to be quite the fashion. Thus it was that Cuvier came forward with a brand-new cla.s.sification of the animal kingdom, establishing four great types of being, which he called vertebrates, mollusks, articulates, and radiates. Lamarck had shortly before established the broad distinction between animals with and those without a backbone; Cuvier's Cla.s.sification divided the latter--the invertebrates--into three minor groups. And this division, familiar ever since to all students of zoology, has only in very recent years been supplanted, and then not by revolution, but by a further division, which the elaborate recent studies of lower forms of life seemed to make desirable.

In the course of those studies of comparative anatomy which led to his new cla.s.sification, Cuvier's attention was called constantly to the peculiar co-ordination of parts in each individual organism. Thus an animal with sharp talons for catching living prey--as a member of the cat tribe--has also sharp teeth, adapted for tearing up the flesh of its victim, and a particular type of stomach, quite different from that of herbivorous creatures. This adaptation of all the parts of the animal to one another extends to the most diverse parts of the organism, and enables the skilled anatomist, from the observation of a single typical part, to draw inferences as to the structure of the entire animal--a fact which was of vast aid to Cuvier in his studies of paleontology. It did not enable Cuvier, nor does it enable any one else, to reconstruct fully the extinct animal from observation of a single bone, as has sometimes been a.s.serted, but what it really does establish, in the hands of an expert, is sufficiently astonishing.

"While the study of the fossil remains of the greater quadrupeds is more satisfactory," he writes, "by the clear results which it affords, than that of the remains of other animals found in a fossil state, it is also complicated with greater and more numerous difficulties. Fossil sh.e.l.ls are usually found quite entire, and retaining all the characters requisite for comparing them with the specimens contained in collections of natural history, or represented in the works of naturalists. Even the skeletons of fishes are found more or less entire, so that the general forms of their bodies can, for the most part, be ascertained, and usually, at least, their generic and specific characters are determinable, as these are mostly drawn from their solid parts. In quadrupeds, on the contrary, even when their entire skeletons are found, there is great difficulty in discovering their distinguishing characters, as these are chiefly founded upon their hairs and colors and other marks which have disappeared previous to their incrustation. It is also very rare to find any fossil skeletons of quadrupeds in any degree approaching to a complete state, as the strata for the most part only contain separate bones, scattered confusedly and almost always broken and reduced to fragments, which are the only means left to naturalists for ascertaining the species or genera to which they have belonged.

"Fortunately comparative anatomy, when thoroughly understood, enables us to surmount all these difficulties, as a careful application of its principles instructs us in the correspondences and dissimilarities of the forms of organized bodies of different kinds, by which each may be rigorously ascertained from almost every fragment of its various parts and organs.

"Every organized individual forms an entire system of its own, all the parts of which naturally correspond, and concur to produce a certain definite purpose, by reciprocal reaction, or by combining towards the same end. Hence none of these separate parts can change their forms without a corresponding change in the other parts of the same animal, and consequently each of these parts, taken separately, indicates all the other parts to which it has belonged. Thus, as I have elsewhere shown, if the viscera of an animal are so organized as only to be fitted for the digestion of recent flesh, it is also requisite that the jaws should be so constructed as to fit them for devouring prey; the claws must be constructed for seizing and tearing it to pieces; the teeth for cutting and dividing its flesh; the entire system of the limbs, or organs of motion, for pursuing and overtaking it; and the organs of sense for discovering it at a distance. Nature must also have endowed the brain of the animal with instincts sufficient for concealing itself and for laying plans to catch its necessary victims....

"To enable the animal to carry off its prey when seized, a corresponding force is requisite in the muscles which elevate the head, and this necessarily gives rise to a determinate form of the vertebrae to which these muscles are attached and of the occiput into which they are inserted. In order that the teeth of a carnivorous animal may be able to cut the flesh, they require to be sharp, more or less so in proportion to the greater or less quant.i.ty of flesh that they have to cut. It is requisite that their roots should be solid and strong, in proportion to the quant.i.ty and size of the bones which they have to break to pieces.

The whole of these circ.u.mstances must necessarily influence the development and form of all the parts which contribute to move the jaws...."

After these observations, it will be easily seen that similar conclusions may be drawn with respect to the limbs of carnivorous animals, which require particular conformations to fit them for rapidity of motion in general; and that similar considerations must influence the forms and connections of the vertebrae and other bones const.i.tuting the trunk of the body, to fit them for flexibility and readiness of motion in all directions. The bones also of the nose, of the orbit, and of the ears require certain forms and structures to fit them for giving perfection to the senses of smell, sight, and hearing, so necessary to animals of prey. In short, the shape and structure of the teeth regulate the forms of the condyle, of the shoulder-blade, and of the claws, in the same manner as the equation of a curve regulates all its other properties; and as in regard to any particular curve all its properties may be ascertained by a.s.suming each separate property as the foundation of a particular equation, in the same manner a claw, a shoulder-blade, a condyle, a leg or arm bone, or any other bone separately considered, enables us to discover the description of teeth to which they have belonged; and so also reciprocally we may determine the forms of the other bones from the teeth. Thus commencing our investigations by a careful survey of any one bone by itself, a person who is sufficiently master of the laws of organic structure may, as it were, reconstruct the whole animal to which that bone belonged."(1)

We have already pointed out that no one is quite able to perform the necromantic feat suggested in the last sentence; but the exaggeration is pardonable in the enthusiast to whom the principle meant so much and in whose hands it extended so far.

Of course this entire principle, in its broad outlines, is something with which every student of anatomy had been familiar from the time when anatomy was first studied, but the full expression of the "law of co-ordination," as Cuvier called it, had never been explicitly made before; and, notwithstanding its seeming obviousness, the exposition which Cuvier made of it in the introduction to his cla.s.sical work on comparative anatomy, which was published during the first decade of the nineteenth century, ranks as a great discovery. It is one of those generalizations which serve as guideposts to other discoveries.

BICHAT AND THE BODILY TISSUES

Much the same thing may be said of another generalization regarding the animal body, which the brilliant young French physician Marie Francois Bichat made in calling attention to the fact that each vertebrate organism, including man, has really two quite different sets of organs--one set under volitional control, and serving the end of locomotion, the other removed from volitional control, and serving the ends of the "vital processes" of digestion, a.s.similation, and the like.

He called these sets of organs the animal system and the organic system, respectively. The division thus pointed out was not quite new, for Grimaud, professor of physiology in the University of Montpellier, had earlier made what was substantially the same cla.s.sification of the functions into "internal or digestive and external or locomotive"; but it was Bichat's exposition that gave currency to the idea.

Far more important, however, was another cla.s.sification which Bichat put forward in his work on anatomy, published just at the beginning of the last century. This was the division of all animal structures into what Bichat called tissues, and the pointing out that there are really only a few kinds of these in the body, making up all the diverse organs. Thus muscular organs form one system; membranous organs another; glandular organs a third; the vascular mechanism a fourth, and so on. The distinction is so obvious that it seems rather difficult to conceive that it could have been overlooked by the earliest anatomists; but, in point of fact, it is only obvious because now it has been familiarly taught for almost a century. It had never been given explicit expression before the time of Bichat, though it is said that Bichat himself was somewhat indebted for it to his master, Desault, and to the famous alienist Pinel.

However that may be, it is certain that all subsequent anatomists have found Bichat's cla.s.sification of the tissues of the utmost value in their studies of the animal functions. Subsequent advances were to show that the distinction between the various tissues is not really so fundamental as Bichat supposed, but that takes nothing from the practical value of the famous cla.s.sification.

It was but a step from this scientific cla.s.sification of tissues to a similar cla.s.sification of the diseases affecting them, and this was one of the greatest steps towards placing medicine on the plane of an exact science. This subject of these branches completely fascinated Bichat, and he exclaimed, enthusiastically: "Take away some fevers and nervous trouble, and all else belongs to the kingdom of pathological anatomy."

But out of this enthusiasm came great results. Bichat practised as he preached, and, believing that it was only possible to understand disease by observing the symptoms carefully at the bedside, and, if the disease terminated fatally, by post-mortem examination, he was so arduous in his pursuit of knowledge that within a period of less than six months he had made over six hundred autopsies--a record that has seldom, if ever, been equalled. Nor were his efforts fruitless, as a single example will suffice to show. By his examinations he was able to prove that diseases of the chest, which had formerly been cla.s.sed under the indefinite name "peripneumonia," might involve three different structures, the pleural sac covering the lungs, the lung itself, and the bronchial tubes, the diseases affecting these organs being known respectively as pleuritis, pneumonia, and bronchitis, each one differing from the others as to prognosis and treatment. The advantage of such an exact cla.s.sification needs no demonstration.

LISTER AND THE PERFECTED MICROSCOPE

At the same time when these broad macroscopical distinctions were being drawn there were other workers who were striving to go even deeper into the intricacies of the animal mechanism with the aid of the microscope.

This undertaking, however, was beset with very great optical difficulties, and for a long time little advance was made upon the work of preceding generations. Two great optical barriers, known technically as spherical and chromatic aberration--the one due to a failure of the rays of light to fall all in one plane when focalized through a lens, the other due to the dispersive action of the lens in breaking the white light into prismatic colors--confronted the makers of microscopic lenses, and seemed all but insuperable. The making of achromatic lenses for telescopes had been accomplished, it is true, by Dolland in the previous century, by the union of lenses of crown gla.s.s with those of flint gla.s.s, these two materials having different indices of refraction and dispersion. But, aside from the mechanical difficulties which arise when the lens is of the minute dimensions required for use with the microscope, other perplexities are introduced by the fact that the use of a wide pencil of light is a desideratum, in order to gain sufficient illumination when large magnification is to be secured.

In the attempt to overcome those difficulties, the foremost physical philosophers of the time came to the aid of the best opticians. Very early in the century, Dr. (afterwards Sir David) Brewster, the renowned Scotch physicist, suggested that certain advantages might accrue from the use of such gems as have high refractive and low dispersive indices, in place of lenses made of gla.s.s. Accordingly lenses were made of diamond, of sapphire, and so on, and with some measure of success. But in 1812 a much more important innovation was introduced by Dr. William Hyde Wollaston, one of the greatest and most versatile, and, since the death of Cavendish, by far the most eccentric of English natural philosophers. This was the suggestion to use two plano-convex lenses, placed at a prescribed distance apart, in lieu of the single double-convex lens generally used. This combination largely overcame the spherical aberration, and it gained immediate fame as the "Wollaston doublet."

To obviate loss of light in such a doublet from increase of reflecting surfaces, Dr. Brewster suggested filling the inters.p.a.ce between the two lenses with a cement having the same index of refraction as the lenses themselves--an improvement of manifest advantage. An improvement yet more important was made by Dr. Wollaston himself in the introduction of the diaphragm to limit the field of vision between the lenses, instead of in front of the anterior lens. A pair of lenses thus equipped Dr.

Wollaston called the periscopic microscope. Dr. Brewster suggested that in such a lens the same object might be attained with greater ease by grinding an equatorial groove about a thick or globular lens and filling the groove with an opaque cement. This arrangement found much favor, and came subsequently to be known as a Coddington lens, though Mr.

Coddington laid no claim to being its inventor.

Sir John Herschel, another of the very great physicists of the time, also gave attention to the problem of improving the microscope, and in 1821 he introduced what was called an aplanatic combination of lenses, in which, as the name implies, the spherical aberration was largely done away with. It was thought that the use of this Herschel aplanatic combination as an eyepiece, combined with the Wollaston doublet for the objective, came as near perfection as the compound microscope was likely soon to come. But in reality the instrument thus constructed, though doubtless superior to any predecessor, was so defective that for practical purposes the simple microscope, such as the doublet or the Coddington, was preferable to the more complicated one.

Many opticians, indeed, quite despaired of ever being able to make a satisfactory refracting compound microscope, and some of them had taken up anew Sir Isaac Newton's suggestion in reference to a reflecting microscope. In particular, Professor Giovanni Battista Amici, a very famous mathematician and practical optician of Modena, succeeded in constructing a reflecting microscope which was said to be superior to any compound microscope of the time, though the events of the ensuing years were destined to rob it of all but historical value. For there were others, fortunately, who did not despair of the possibilities of the refracting microscope, and their efforts were destined before long to be crowned with a degree of success not even dreamed of by any preceding generation.

The man to whom chief credit is due for directing those final steps that made the compound microscope a practical implement instead of a scientific toy was the English amateur optician Joseph Jackson Lister.

Combining mathematical knowledge with mechanical ingenuity, and having the practical aid of the celebrated optician Tulley, he devised formulae for the combination of lenses of crown gla.s.s with others of flint gla.s.s, so adjusted that the refractive errors of one were corrected or compensated by the other, with the result of producing lenses of hitherto unequalled powers of definition; lenses capable of showing an image highly magnified, yet relatively free from those distortions and fringes of color that had heretofore been so disastrous to true interpretation of magnified structures.

Lister had begun his studies of the lens in 1824, but it was not until 1830 that he contributed to the Royal Society the famous paper detailing his theories and experiments. Soon after this various continental opticians who had long been working along similar lines took the matter up, and their expositions, in particular that of Amici, introduced the improved compound microscope to the attention of microscopists everywhere. And it required but the most casual trial to convince the experienced observers that a new implement of scientific research had been placed in their hands which carried them a long step nearer the observation of the intimate physical processes which lie at the foundation of vital phenomena. For the physiologist this perfection of the compound microscope had the same significance that the, discovery of America had for the fifteenth-century geographers--it promised a veritable world of utterly novel revelations. Nor was the fulfilment of that promise long delayed.

Indeed, so numerous and so important were the discoveries now made in the realm of minute anatomy that the rise of histology to the rank of an independent science may be said to date from this period. Hitherto, ever since the discovery of magnifying-gla.s.ses, there had been here and there a man, such as Leuwenhoek or Malpighi, gifted with exceptional vision, and perhaps unusually happy in his conjectures, who made important contributions to the knowledge of the minute structure of organic tissues; but now of a sudden it became possible for the veriest tyro to confirm or refute the laborious observations of these pioneers, while the skilled observer could step easily beyond the barriers of vision that hitherto were quite impa.s.sable. And so, naturally enough, the physiologists of the fourth decade of the nineteenth century rushed as eagerly into the new realm of the microscope as, for example, their successors of to-day are exploring the realm of the X-ray.

Lister himself, who had become an eager interrogator of the instrument he had perfected, made many important discoveries, the most notable being his final settlement of the long-mooted question as to the true form of the red corpuscles of the human blood. In reality, as everybody knows nowadays, these are biconcave disks, but owing to their peculiar figure it is easily possible to misinterpret the appearances they present when seen through a poor lens, and though Dr. Thomas Young and various other observers had come very near the truth regarding them, unanimity of opinion was possible only after the verdict of the perfected microscope was given.

These blood corpuscles are so infinitesimal in size that something like five millions of them are found in each cubic millimetre of the blood, yet they are isolated particles, each having, so to speak, its own personality. This, of course, had been known to microscopists since the days of the earliest lenses. It had been noticed, too, by here and there an observer, that certain of the solid tissues seemed to present something of a granular texture, as if they, too, in their ultimate const.i.tution, were made up of particles. And now, as better and better lenses were constructed, this idea gained ground constantly, though for a time no one saw its full significance. In the case of vegetable tissues, indeed, the fact that little particles encased a membranous covering, and called cells, are the ultimate visible units of structure had long been known. But it was supposed that animal tissues differed radically from this construction. The elementary particles of vegetables "were regarded to a certain extent as individuals which composed the entire plant, while, on the other hand, no such view was taken of the elementary parts of animals."

ROBERT BROWN AND THE CELL NUCLEUS

In the year 1833 a further insight into the nature of the ultimate particles of plants was gained through the observation of the English microscopist Robert Brown, who, in the course of his microscopic studies of the epidermis of orchids, discovered in the cells "an opaque spot,"

which he named the nucleus. Doubtless the same "spot" had been seen often enough before by other observers, but Brown was the first to recognize it as a component part of the vegetable cell and to give it a name.

"I shall conclude my observations on Orchideae," said Brown, "with a notice of some points of their general structure, which chiefly relate to the cellular tissue. In each cell of the epidermis of a great part of this family, especially of those with membranous leaves, a single circular areola, generally somewhat more opaque than, the membrane of the cell, is observable. This areola, which is more or less distinctly granular, is slightly convex, and although it seems to be on the surface is in reality covered by the outer lamina of the cell. There is no regularity as to its place in the cell; it is not unfrequently, however, central or nearly so.

"As only one areola belongs to each cell, and as in many cases where it exists in the common cells of the epidermis, it is also visible in the cutaneous glands or stomata, and in these is always double--one being on each side of the limb--it is highly probable that the cutaneous gland is in all cases composed of two cells of peculiar form, the line of union being the longitudinal axis of the disk or pore.

"This areola, or nucleus of the cell as perhaps it might be termed, is not confined to the epidermis, being also found, not only in the p.u.b.escence of the surface, particularly when jointed, as in cypripedium, but in many cases in the parenchyma or internal cells of the tissue, especially when these are free from the deposition of granular matter.

"In the compressed cells of the epidermis the nucleus is in a corresponding degree flattened; but in the internal tissue it is often nearly spherical, more or less firmly adhering to one of the walls, and projecting into the cavity of the cell. In this state it may not unfrequently be found in the substance of the column and in that of the perianthium.

"The nucleus is manifest also in the tissue of the stigma, where in accordance with the compression of the utriculi, it has an intermediate form, being neither so much flattened as in the epidermis nor so convex as it is in the internal tissue of the column.

"I may here remark that I am acquainted with one case of apparent exception to the nucleus being solitary in each utriculus or cell--namely, in Bletia Tankervilliae. In the utriculi of the stigma of this plant, I have generally, though not always, found a second areola apparently on the surface, and composed of much larger granules than the ordinary nucleus, which is formed of very minute granular matter, and seems to be deep seated.

"Mr. Bauer has represented the tissue of the stigma, in the species of Bletia, both before and, as he believes, after impregnation; and in the latter state the utriculi are marked with from one to three areolae of similar appearance.