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THEORY OF FREE CELL-FORMATION. 9
tions, and dependent upon peculiar changes which the element has undergone. Generally, it may be said that, as long as the life of the cell has not been brought to a close,
as long as cells behave as elements still endowed with vital power, the nucleus maintains a very nearly constant form.
The nucleus, in its turn, in completely developed cells, very constantly encloses another structure within itself— the so-called nucleolus. With regard to the question of vital form, it cannot be said of the nucleolus that it appears to be an absolute requisite; and, in a considerable number of young cells, it has as yet escaped detection. On the other hand, we regularly meet with it in fully developed, older forms; and it, therefore, seems to mark a higher degree of development in the cell. According to the view which was put forward in the first instance by Schleiden, and accepted by Schwann, the connection between the three coexistent cellconstituents was long thought to be on this wise: that the nucleolus was the first to show itself in the development of tissues, by separating out of a formative fluid CMastema, cytoblastema), that it quickly attained a certain size, that then fine granules were precipitated out of the blastema and settled around it, and that about these there condensed a membrane. That in this way a nucleus was
Pig. 3. a. Hepatic cell. h. Spindle-shaped cells from connective tissue, e. Capillary vessel, d. Somewhat large stellate cell from a lymphatic gland. «. Ganglion-cell from the cerebellum. The nuclei in every instance similar.
completed, about which new matter gradually gathered, and in due time produced a little membrane (the celebrated watch-glass form, fig. 4, d'). This description of the first development of • ••..*:::.". P cells out of free blastema, according '.•'.•.''leery to which the nucleus was regarded as cy'''<$%£/<3£'\ Prececnng the formation of the cell, d !;^$i&$'\><' and playing the part of a real cellformer (cyioblast), is the one which is usually concisely designated by the name of the celltheory (more accurately, theory of free cell-formation),—a theory of development which has now been almost entirely abandoned, and in support of the correctness of which not one single fact can with certainty be adduced. With respect to the nucleolus, all that we can for the present regard as certain, is, that where we have to deal with large and fully developed cells, we almost constantly see a nucleolus in them; but that, on the contrary, in the case of many young cells it is wanting.
You will hereafter be made acquainted with a series of facts in the history of pathological and physiological development, which render it in a high degree probable that the nucleus plays an extremely important part within the cell—a part, I will here at once remark, less connected with the function and specific office of the cell, than with its maintenance and multiplication as a living part. The specific (in a narrower sense, animal) function is most distinctly manifested in muscles, nerves, and gland-cells; the
Fig. 4. From Schleiden, 'Grundzuge der wiss. Botanik,' I," fig. ]. "Contents of the embryo-sac of Vicia /aba soon after impregnation. In the clear fluid, consisting of gum and sugar, granules of protein-compounds are seen swimming about (a), among which a few larger ones are strikingly conspicuous. Around these latter the former are seen conglomerated into the form of a small disc (b, e). Around other discs a clear, sharply defined border may be distinguished, which gradually recedes farther and farther from the disc (the cytoblast), and, finally, can be distinctly recognised to be a young cell (d, «)." IMPORT OF THE NUCLEUS AND CELL-CONTENTS. 11
peculiar actions of which — contraction, sensation, and secretion—appear to be connected in no direct manner with the nuclei. But that, whilst fulfilling all its functions, the element remains an element, that it is not annihilated nor destroyed by its continual activity—this seems essentially to depend upon the action of the nucleus. All those cellular formations which lose their nucleus, have a more transitory existence; they perish, they disappear, they die away or break up. A human blood-corpuscle, for example, is a cell without a nucleus; it possesses an external membrane and red contents; but herewith the tale of its constituents, so far as we can make them out, is told, and whatever has been recounted concerning a nucleus in blood-cells, has had its foundation in delusive appearances, which certainly very easily can be, and frequently are, occasioned by the production of little irregularities upon the surface (Fig. 52). We should not be able to say, therefore, that blood-corpuscles were cells, if we did not know that there is a certain period during which human bloodcorpuscles also have nuclei; the period, namely, embraced by the first months of intra-uterine life. Then circulate also in the human body nucleated blood-cells, like those which we see in frogs, birds, and fish throughout the whole of their lives. In mammalia, however, this is restricted to a certain period of their development, so that at a later stage the red blood-cells no longer exhibit all the characteristics of a cell, but have lost an important constituent in their composition. But we are also all agreed upon this point, that the blood is one of those changeable constituents of the body, whose cellular elements possess no durability, and with regard to which everybody assumes that they perish, and are replaced by new ones, which in their turn are doomed to annihilation, and everywhere (like the uppermost cells in the cuticle, in which we also can discover no nuclei, as soon as they begin to desquamate) have already reached a stage in their development, when they no longer require that durability in their more intimate composition for which we must regard the nucleus as the guarantee.
On the other hand, notwithstanding the manifold investigations to which the tissues are at present subjected, we are acquainted with no part which grows or multiplies, either in a physiological or pathological manner, in which nucleated elements cannot invariably be demonstrated as the starting-points of the change, and in which the first decisive alterations which display themselves, do not involve the nucleus itself, so that we often can determine from its condition what would possibly have become of the elements.
You see from this description that, at least, two different things are of necessity required for the composition of a cellular element; the membrane, whether round, jagged, or stellate, and the nucleus, which from the outset differs in chemical constitution from the membrane. Herewith, however, we are far from having enumerated all the essential constituents of the cell, for, in addition to the nucleus, it is filled with a relatively greater or less quantity of contents, as is likewise commonly, it seems, the nucleus itself, the contents of which are also wont to differ from those of the cell. Within the cell, for example, we see pigment, without the nucleus' containing any. Within a smooth muscular fibre-cell, the contractile substance is deposited, which appears to be the seat of the contractile force of muscle ; the nucleus, however, remains a nucleus. The cell may develop itself into a nerve-fibre, but the nucleus remains, lying
Fig. 5. a. Pigment-cell from the choroid membrane of the eye. b. Smooth muscular fibre-cell from the intestines, c. Portion of a nerve-fibre with a double contour, axis-cylinder, medullary sheath and parietal, nucleolated nucleus
IMPORT OF THE NUCLEUS AND CELL-CONTENTS. 13
on the outside of the medullary [white1] substance, a constant constituent. Hence it follows, that the special peculiarities which individual cells exhibit in particular places, under particular circumstances, are in general dependent upon the varying properties of the cell-contents, and that it is not the constituents which we have hitherto considered (membrane and nucleus), but the contents (or else the masses of matter deposited without the cell, intercellular), which give rise to the functional'(physiological) differences of tissues. For us it is essential to know that in the most various tissues these constituents, which, in some measure, represent the cell in its abstract form, the nucleus and membrane, recur with great constancy, and that by their combination a simple element is obtained, which, throughout the whole series of living vegetable and animal forms, however different they may be externally, however much their internal composition may be subjected to change, presents us with a structure of quite a peculiar conformation, as a definite basis for all the phenomena of life.
According to my ideas, this is the only possible startingpoint for all biological doctrines. If a definite correspondence in elementary form pervades the whole series of all living things, and if in this series something else which might be placed in the stead of the cell be in vain sought for, then must every more highly developed organism, whether vegetable or animal, necessarily, above all, be regarded as a progressive total, made up of larger or smaller number of similar or dissimilar cells. Just as a tree constitutes a mass arranged in a definite manner, in which, in every single part, in the leaves as in the root, in the trunk as in the blossom, cells are discovered to be the ultimate elements, so is it also with the forms of animal life. Every animal presents itself as a sum of vital unities, every one of
1 All words included in square brackets have been inserted by the Translator, and are intended to be explanatory.