by Henry Smith Williams, M.D., LL.D.
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 pubescence 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.
"The nucleus may even be supposed to exist in the pollen of this family. In the early stages of its formation, at least a minute areola is of ten visible in the simple grain, and in each of the constituent parts of cells of the compound grain. But these areolae may perhaps rather be considered as merely the points of production of the tubes.
"This nucleus of the cell is not confined to orchideae, but is equally manifest in many other monocotyledonous families; and I have even found it, hitherto however in very few cases, in the epidermis of dicotyledonous plants; though in this primary division it may perhaps be said to exist in the early stages of development of the pollen. Among monocotyledons, the orders in which it is most remarkable are Liliaceae, Hemerocallideae, Asphodeleae, Irideae, and Commelineae.
"In some plants belonging to this last-mentioned family, especially in Tradascantia virginica, and several nearly related species, it is uncommonly distinct, not in the epidermis and in the jointed hairs of the filaments, but in the tissue of the stigma, in the cells of the ovulum even before impregnation, and in all the stages of formation of the grains of pollen, the evolution of which is so remarkable in tradascantia.
"The few indications of the presence of this nucleus, or areola, that I have hitherto met with in the publications of botanists are chiefly in some figures of epidermis, in the recent works of Meyen and Purkinje, and in one case, in M. Adolphe Broigniart's memoir on the structure of leaves. But so little importance seems to be attached to it that the appearance is not always referred to in the explanations of the figures in which it is represented. Mr. Bauer, however, who has also figured it in the utriculi of the stigma of Bletia Tankervilliae has more particularly noticed it, and seems to consider it as only visible after impregnation."[2]
SCHLEIDEN AND SCHWANN AND THE CELL THEORY
That this newly recognized structure must be important in the economy of the cell was recognized by Brown himself, and by the celebrated German Meyen, who dealt with it in his work on vegetable physiology, published not long afterwards; but it remained for another German, the professor of botany in the University of Jena, Dr. M. J. Schleiden, to bring the nucleus to popular attention, and to assert its all-importance in the economy of the cell.
Schleiden freely acknowledged his indebtedness to Brown for first knowledge of the nucleus, but he soon carried his studies of that structure far beyond those of its discoverer. He came to believe that the nucleus is really the most important portion of the cell, in that it is the original structure from which the remainder of the cell is developed. Hence he named it the cytoblast. He outlined his views in an epochal paper published in Muller's Archives in 1838, under title of "Beitrage zur Phytogenesis." This paper is in itself of value, yet the most important outgrowth of Schleiden's observations of the nucleus did not spring from his own labors, but from those of a friend to whom he mentioned his discoveries the year previous to their publication. This friend was Dr. Theodor Schwann, professor of physiology in the University of Louvain.
At the moment when these observations were communicated to him Schwann was puzzling over certain details of animal histology which he could not clearly explain. His great teacher, Johannes Muller, had called attention to the strange resemblance to vegetable cells shown by certain cells of the chorda dorsalis (the embryonic cord from which the spinal column is developed), and Schwann himself had discovered a corresponding similarity in the branchial cartilage of a tadpole. Then, too, the researches of Friedrich Henle had shown that the particles that make up the epidermis of animals are very cell-like in appearance. Indeed, the cell-like character of certain animal tissues had come to be matter of common note among students of minute anatomy. Schwann felt that this similarity could not be mere coincidence, but he had gained no clew to further insight until Schleiden called his attention to the nucleus. Then at once he reasoned that if there really is the correspondence between vegetable and animal tissues that he suspected, and if the nucleus is so important in the vegetable cell as Schleiden believed, the nucleus should also be found in the ultimate particles of animal tissues.
Schwann's researches soon showed the entire correctness of this assumption. A closer study of animal tissues under the microscope showed, particularly in the case of embryonic tissues, that "opaque spots" such as Schleiden described are really to be found there in abundance--forming, indeed, a most characteristic phase of the structure. The location of these nuclei at comparatively regular intervals suggested that they are found in definite compartments of the tissue, as Schleiden had shown to be the case with vegetables; indeed, the walls that separated such cell-like compartments one from another were in some cases visible. Particularly was this found to be the case with embryonic tissues, and the study of these soon convinced Schwann that his original surmise had been correct, and that all animal tissues are in their incipiency composed of particles not unlike the ultimate particles of vegetables in short, of what the botanists termed cells. Adopting this name, Schwann propounded what soon became famous as his cell theory, under title of Mikroskopische Untersuchungen uber die Ubereinstimmung in der Structur und dent Wachsthum der Thiere und Pflanzen. So expeditious had been his work that this book was published early in 1839, only a few months after the appearance of Schleiden's paper.
As the title suggests, the main idea that actuated Schwann was to unify vegetable and animal tissues. Accepting cell-structure as the basis of all vegetable tissues, he sought to show that the same is true of animal tissues, all the seeming diversities of fibre being but the alteration and development of what were originally simple cells. And by cell Schwann meant, as did Schleiden also, what the word ordinarily implies--a cavity walled in on all sides. He conceived that the ultimate constituents of all tissues were really such minute cavities, the most important part of which was the cell wall, with its associated nucleus. He knew, indeed, that the cell might be filled with fluid contents, but he regarded these as relatively subordinate in importance to the wall itself. This, however, did not apply to the nucleus, which was supposed to lie against the cell wall and in the beginning to generate it. Subsequently the wall might grow so rapidly as to dissociate itself from its contents, thus becoming a hollow bubble or true cell; but the nucleus, as long as it lasted, was supposed to continue in contact with the cell wall. Schleiden had even supposed the nucleus to be a constituent part of the wall, sometimes lying enclosed between two layers of its substance, and Schwann quoted this view with seeming approval. Schwann believed, however, that in the mature cell the nucleus ceased to be functional and disappeared.
The main thesis as to the similarity of development of vegetable and animal tissues and the cellular nature of the ultimate constitution of both was supported by a mass of carefully gathered evidence which a multitude of microscopists at once confirmed, so Schwann's work became a classic almost from the moment of its publication. Of course various other workers at once disputed Schwann's claim to priority of discovery, in particular the English microscopist Valentin, who asserted, not without some show of justice, that he was working closely along the same lines. Put so, for that matter, were numerous others, as Henle, Turpin, Du-mortier, Purkinje, and Muller, all of whom Schwann himself had quoted. Moreover, there were various physiologists who earlier than any of these had foreshadowed the cell theory--notably Kaspar Friedrich Wolff, towards the close of the previous century, and Treviranus about 1807, But, as we have seen in so many other departments of science, it is one thing to foreshadow a discovery, it is quite another to give it full expression and make it germinal of other discoveries. And when Schwann put forward the explicit claim that "there is one universal principle of development for the elementary parts, of organisms, however different, and this principle is the formation of cells," he enunciated a doctrine which was for all practical purposes absolutely new and opened up a novel field for the microscopist to enter. A most important era in physiology dates from the publication of his book in 1839.
THE CELL THEORY ELABORATED
That Schwann should have gone to embryonic tissues for the establishment of his ideas was no doubt due very largely to the influence of the great Russian Karl Ernst von Baer, who about ten years earlier had published the first part of his celebrated work on embryology, and whose ideas were rapidly gaining ground, thanks largely to the advocacy of a few men, notably Johannes Muller, in Germany, and William B. Carpenter, in England, and to the fact that the improved microscope had made minute anatomy popular. Schwann's researches made it plain that the best field for the study of the animal cell is here, and a host of explorers entered the field. The result of their observations was, in the main, to confirm the claims of Schwann as to the universal prevalence of the cell. The long-current idea that animal tissues grow only as a sort of deposit from the blood-vessels was now discarded, and the fact of so-called plantlike growth of animal cells, for which Schwann contended, was universally accepted. Yet the full measure of the affinity between the two classes of cells was not for some time generally apprehended.
Indeed, since the substance that composes the cell walls of plants is manifestly very different from the limiting membrane of the animal cell, it was natural, so long as the, wall was considered the most essential part of the structure, that the divergence between the two classes of cells should seem very pronounced. And for a time this was the conception of the matter that was uniformly accepted. But as time went on many observers had their attention called to the peculiar characteristics of the contents of the cell, and were led to ask themselves whether these might not be more important than had been supposed. In particular, Dr. Hugo von Mohl, professor of botany in the University of Tubingen, in the course of his exhaustive studies of the vegetable cell, was impressed with the peculiar and characteristic appearance of the cell contents. He observed universally within the cell "an opaque, viscid fluid, having granules intermingled in it," which made up the main substance of the cell, and which particularly impressed him because under certain conditions it could be seen to be actively in motion, its parts separated into filamentous streams.
Von Mohl called attention to the fact that this motion of the cell contents had been observed as long ago as 1774 by Bonaventura Corti, and rediscovered in 1807 by Treviranus, and that these observers had described the phenomenon under the "most unsuitable name of 'rotation of the cell sap.' Von Mohl recognized that the streaming substance was something quite different from sap. He asserted that the nucleus of the cell lies within this substance and not attached to the cell wall as Schleiden had contended. He saw, too, that the chlorophyl granules, and all other of the cell contents, are incorporated with the "opaque, viscid fluid," and in 1846 he had become so impressed with the importance of this universal cell substance that be gave it the name of protoplasm. Yet in so doing he had no intention of subordinating the cell wall. The fact that Payen, in 1844, had demonstrated that the cell walls of all vegetables, high or low, are composed largely of one substance, cellulose, tended to strengthen the position of the cell wall as the really essential structure, of which the protoplasmic contents were only subsidiary products.
Meantime, however, the students of animal histology were more and more impressed with the seeming preponderance of cell contents over cell walls in the tissues they studied. They, too, found the cell to be filled with a viscid, slimy fluid capable of motion. To this Dujardin gave the name of sarcode. Presently it came to be known, through the labors of Kolliker, Nageli, Bischoff, and various others, that there are numerous lower forms of animal life which seem to be composed of this sarcode, without any cell wall whatever. The same thing seemed to be true of certain cells of higher organisms, as the blood corpuscles. Particularly in the case of cells that change their shape markedly, moving about in consequence of the streaming of their sarcode, did it seem certain that no cell wall is present, or that, if present, its role must be insignificant.
And so histologists came to question whether, after all, the cell contents rather than the enclosing wall must not be the really essential structure, and the weight of increasing observations finally left no escape from the conclusion that such is really the case. But attention being thus focalized on the cell contents, it was at once apparent that there is a far closer similarity between the ultimate particles of vegetables and those of animals than had been supposed. Cellulose and animal membrane being now regarded as more by-products, the way was clear for the recognition of the fact that vegetable protoplasm and animal sarcode are marvellously similar in appearance and general properties. The closer the observation the more striking seemed this similarity; and finally, about 1860, it was demonstrated by Heinrich de Bary and by Max Schultze that the two are to all intents and purposes identical. Even earlier Remak had reached a similar conclusion, and applied Von Mohl's word protoplasm to animal cell contents, and now this application soon became universal. Thenceforth this protoplasm was to assume the utmost importance in the physiological world, being recognized as the universal "physical basis of life," vegetable and animal alike. This amounted to the logical extension and culmination of Schwann's doctrine as to the similarity of development of the two animate kingdoms. Yet at the, same time it was in effect the banishment of the cell that Schwann had defined. The word cell was retained, it is true, but it no longer signified a minute cavity. It now implied, as Schultze defined it, "a small mass of protoplasm endowed with the attributes of life." This definition was destined presently to meet with yet another modification, as we shall see; but the conception of the protoplasmic mass as the essential ultimate structure, which might or might not surround itself with a protective covering, was a permanent addition to physiological knowledge. The earlier idea had, in effect, declared the shell the most important part of the egg; this developed view assigned to the yolk its true position.
In one other important regard the theory of Schleiden and Schwann now became modified. This referred to the origin of the cell. Schwann had regarded cell growth as a kind of crystallization, beginning with the deposit of a nucleus about a granule in the intercellular substance--the cytoblastema, as Schleiden called it. But Von Mohl, as early as 1835, had called attention to the formation of new vegetable cells through the division of a pre-existing cell. Ehrenberg, another high authority of the time, contended that no such division occurs, and the matter was still in dispute when Schleiden came forward with his discovery of so-called free cell-formation within the parent cell, and this for a long time diverted attention from the process of division which Von Mohl had described. All manner of schemes of cell-formation were put forward during the ensuing years by a multitude of observers, and gained currency notwithstanding Von Mohl's reiterated contention that there are really but two ways in which the formation of new cells takes place--namely, "first, through division of older cells; secondly, through the formation of secondary cells lying free in the cavity of a cell."
But gradually the researches of such accurate observers as Unger, Nageli, Kolliker, Reichart, and Remak tended to confirm the opinion of Von Mohl that cells spring only from cells, and finally Rudolf Virchow brought the matter to demonstration about 1860. His Omnis cellula e cellula became from that time one of the accepted data of physiology. This was supplemented a little later by Fleming's Omnis nucleus e nucleo, when still more refined methods of observation had shown that the part of the cell which always first undergoes change preparatory to new cell-formation is the all-essential nucleus. Thus the nucleus was restored to the important position which Schwann and Schleiden had given it, but with greatly altered significance. Instead of being a structure generated de novo from non-cellular substance, and disappearing as soon as its function of cell-formation was accomplished, the nucleus was now known as the central and permanent feature of every cell, indestructible while the cell lives, itself the division-product of a pre-existing nucleus, and the parent, by division of its substance, of other generations of nuclei. The word cell received a final definition as "a small mass of protoplasm supplied with a nucleus."
In this widened and culminating general view of the cell theory it became clear that every animate organism, animal or vegetable, is but a cluster of nucleated cells, all of which, in each individual case, are the direct descendants of a single primordial cell of the ovum. In the developed individuals of higher organisms the successive generations of cells become marvellously diversified in form and in specific functions; there is a wonderful division of labor, special functions being chiefly relegated to definite groups of cells; but from first to last there is no function developed that is not present, in a primitive way, in every cell, however isolated; nor does the developed cell, however specialized, ever forget altogether any one of its primordial functions or capacities. All physiology, then, properly interpreted, becomes merely a study of cellular activities; and the development of the cell theory takes its place as the great central generalization in physiology of the nineteenth century. Something of the later developments of this theory we shall see in another connection.
Introduction- A History of Science
I have included extracts from the 4 volume set of A HISTORY OF SCIENCE by Henry Smith Williams, M.D., LL.D. assisted by Edward H. Williams, M.D. A set of texts published early in the 20th century. I recommend reading the 4 volumes as they are an interesting read and written with a different "feel" in language to those texts written today where brevity is carried to such lengths as to make most writings either bland or uninteresting.
This section of notes provides an historic context for the notes covering recent developments in Embryology Research.
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