Chapter XIV: Introduction (4)

The general plastic appearances in the cells are, as we have seen, the following: at first a minute corpuscle is formed (the nucleolus); a layer of substance (the nucleus) is then precipitated around it, which becomes more thickened and expanded by the continual deposition of fresh molecules between those already present. Deposition goes on more vigorously at the outer part of this layer than at the inner. Frequently the entire layer, or in other instances the outer part of it only, becomes condensed to a membrane, which may continue to take up new molecules in such a manner that it increases more rapidly in superficial extent than in thickness, and thus an intervening cavity is necessarily formed between it and the nucleolus. A second layer (cell) is next precipitated around this first, in which precisely the same phenomena are repeated, with merely the difference that in this case the processes, especially the growth of the layer and the formation of the space intervening between it and the first layer (the cell-cavity), go on more rapidly and more completely. Such were the phenomena in the formation of most cells; in some, however, there appeared to be only a single layer formed, while in others (those especially in which the nucleolus was hollow) there were three. The other varieties in the development of the elementary parts were (as we saw) reduced to these--that if two neighbouring cells commence their formation so near to one another that the boundaries of the layers forming around each of them meet at any spot, a common layer may be formed enclosing the two incipient cells. So at least the origin of nuclei, with two or more nucleoli, seemed explicable, by a coalescence of the first layers (corresponding to the nucleus), and the union of many primary cells into one secondary cell by a similar coalescence of the second layers (which correspond to the cell). But the further development of these common layers proceeds as though they were only an ordinary single layer. Lastly, there were some varieties in the progressive development of the cells, which were referable to an unequal deposition of the new molecules between those already present in the separate layers. In this way modifications of form and division of the cells were explained. And among the number of the plastic phenomena in the cells we may mention, lastly, the formation of secondary deposits; for instances occur in which one or more new layers, each on the inner surface of the previous one, are deposited on the inner surface of a simple or of a secondary cell.

These are the most important phenomena observed in the formation and development of cells. The unknown cause, presumed to be capable of explaining these processes in the cells, may be called the plastic power of the cells. We will, in the next place, proceed to determine how far a more accurate definition of this power may be deduced from these phenomena.

In the first place, there is a power of attraction exerted in the very commencement of the cell, in the nucleolus, which occasions the addition of new molecules to those already present. We may imagine the nucleolus itself to be first formed by a sort of crystallization from out of a concentrated fluid. For if a fluid be so concentrated that the molecules of the substance in solution exert a more powerful mutual attraction than is exerted between them and the molecules of the fluid in which they are dissolved, a part of the solid substance must be precipitated. One can readily understand that the fluid must be more concentrated when new cells are being formed in it than when those already present have merely to grow. For if the cell is already partly formed, it exerts an attractive force upon the substance still in solution. There is then a cause for the deposition of this substance, which does not co-operate when no part of the cell is yet formed. Therefore, the greater the attractive force of the cell is, the less concentration of the fluid is required; while, at the commencement of the formation of a cell, the fluid must be more than concentrated. But the conclusion which may be thus directly drawn, as to the attractive power of the cell, may also be verified by observation. Wherever the nutrient fluid is not equally distributed in a tissue, the new cells are formed in that part into which the fluid penetrates first, and where, consequently, it is most concentrated. Upon this fact, as we have seen, depended the difference between the growth of organized and unorganized tissues. And this confirmation of the foregoing conclusion by experience speaks also for the correctness of the reasoning itself.

The attractive power of the cells operates so as to effect the addition of new molecules in two ways,--first, in layers, and secondly, in such a manner in each layer that the new molecules are deposited between those already present. This is only an expression of the fact; the more simple law, by which several layers are formed and the molecules are not all deposited between those already present, cannot yet be explained. The formation of layers may be repeated once, twice, or thrice. The growth of the separate layers is regulated by a law, that the deposition of new molecules should be greatest at the part where the nutrient fluid is most concentrated. Hence the outer part particularly becomes condensed into a membrance both in the layer corresponding to the nucleus and in that answering to the cell, because the nutrient fluid penetrates from without, and consequently is more concentrated at the outer than at the inner part of each layer. For the same reason the nucleus grows rapidly, so long as the layer of the cell is not formed around it, but it either stops growing altogether, or at least grows much more slowly as soon as the cell-layer has surrounded it; because then the latter receives the nutrient matter first, and, therefore, in a more concentrated form. And hence the cell becomes, in a general sense, much more completely developed, while the nucleus-layer usually remains at a stage of development, in which the cell-layer had been in its earlier period. The addition of new molecules is so arranged that the layers increase more considerably in superficial extent than in thickness; and thus an intervening space is formed between each layer and the one preceding it, by which cells and nuclei are formed into actual hollow vesicles. From this it may be inferred that the deposition of new molecules is more active between those which lie side by side along the surface of the membrane, than between those which lie one upon the other in its thickness. Were it otherwise, each layer would increase in thickness, but there would be no intervening cavity between it and the previous one, there would be no vesicles, but a solid body composed of layers.

Attractive power is exerted in all the solid parts of the cell. This follows, not only from the fact that new molecules may be deposited everywhere between those already present, but also from the formation of secondary deposits. When the cavity of a cell is once formed, material may be also attracted from its contents and deposited in layers; and as this deposition takes place upon the inner surface of the membrane of the cell, it is probably that which exerts the attractive influence. This formation of layers on the inner surface of the cell-membrane is, perhaps, merely a repetition of the same process by which, at an earlier period, nucleus and cell were precipitated as layers around the nucleolus. It must, however, be remarked that the identity of these two processes cannot be so clearly proved as that of the processes by which nucleus and cell are formed; more especially as there is a variety in the phenomena, for the secondary deposits in plants occur in spiral forms, while this has at least not yet been demonstrated in the formation of the cell-membrane and the nucleus, although by some botanical writers the cell-membrane itself is supposed to consist of spirals.

The power of attraction may be uniform throughout the whole cell, but it may also be confined to single spots; the deposition of new molecules is then more vigorous at these spots, and the consequence of this uneven growth of the cell-membrane is a change in the form of the cell.

The attractive power of the cells manifest a certain form of election in its operation. It does not take up all the substances contained in the surrounding cytoblastema, but only particular ones, either those which are analogous with the substance already present in the cell (assimilation), or such as differ from it in chemical properties. The several layers grow by assimilation, but when a new layer is being formed, different material from that of the previously-formed layer is attracted: for the nucleolus, the nucleus and cell-membrane are composed of materials which differ in their chemical properties.

Such are the peculiarities of the plastic power of the cells, so far as they can as yet be drawn from observation. But the manifestations of this power presuppose another faculty of the cells. The cytoblastema, in which the cells are formed, contains the elements of the materials of which the cell is composed, but in other combinations; it is not a mere solution of cell-material, but it contains only certain organic substances in solution. The cells, therefore, not only attract materials from out of the cytoblastema, but they must have the faculty of producing chemical changes in its constituent particles. Besides which, all the parts of the cell itself may be chemically altered during the process of its vegetation. The unknown cause of all these phenomena, which we comprise under the term metabolic phenomena of the cells, we will denominate the metabolic power.

The next point which can be proved is, that this power is an attribute of the cells themselves, and that the cytoblastema is passive under it. We may mention vinous fermentation as an instance of this. A decoction of malt will remain for a long time unchanged; but as soon as some yeast is added to it, which consists partly of entire fungi and partly of a number of single cells, the chemical change immediately ensues. Here the decoction of malt is the cytoblastema; the cells clearly exhibit activity, the cytoblastema, in this instance even a boiled fluid, being quite passive during the change. The same occurs when any simple cells, as the spores of the lower plants, are sown in boiled substances.

In the cells themselves again, it appears to be the solid parts, the cell-membrane and the nucleus, which produce the change. The contents of the cell undergo similar and even more various changes than the external the cytoblastema, and it is at least probable that these changes originate with the solid parts composing the cells, especially the cell-membrane, because the secondary deposits are formed on the inner surface of the cell-membrane, and other precipitates are generally formed in the first instance around the nucleus. It may therefore, on the whole, be said that the solid component particles of the cells possess the power of chemically altering the substances in contact with them.

The substances which result from the transformation of the contents of the cell are different from those which are produced by change in the external cytoblastema. What is the cause of this difference, if the metamorphosing power of the cell-membrane be limited to its immediate neighbourhood merely? Might we not much rather expect that converted substance would be found without distinction on the inner as on the outer surface of the cell-membrane? It might be said that the cell-membrane converts the substance in contact with it without distinction, and that the variety in the products of this conversion depends only upon a difference between the convertible substance contained in the cell and the external cytoblastema. But the question then arises, as to how it happens that the contents of the cell differ from the external cytoblastema. If it be true that the cell-membrane, which at first closely surrounds the nucleus, expands in the course of its growth, so as to leave an interspace between it and the cell, and that the contents of the cell consist of fluid which has entered this space merely by imbibition, they cannot differ essentially from the external cytoblastema. I think therefore that, in order to explain the distinction between the cell-contents and the external cytoblastema, we must ascribe to the cell-membrane not only the power in general of chemically altering the substances which it is either in contact with, or has imbibed, but also of so separating them that certain substances appear on its inner, and others on its outer surface. The secretion of substances already present in the blood, as, for instance, of urea, by the cells with which the urinary tubes are lined, cannot be explained without such a faculty of the cells. There is, however, nothing so very hazardous in it, since it is a fact that different substances are separated in the decompositions produced by the galvanic pile. It might perhaps be conjectured from this peculiarity of the metabolic phenomena in the cells, that a particular position of the axes of the atoms composing the cell-membrane is essential for the production of these appearances.

Chemical changes occur, however, not only in the cytoblastema and the cell-contents, but also in the solid parts of which the cells are composed, particularly the cell-membrane. Without wishing to assert that there is any intimate connexion between the metabolic power of the cells and galvanism, I may yet, for the sake of making the representation of the process more clear, remark that the chemical changes produced by a galvanic pile are accompanied by corresponding changes in the pile itself.

The more obscure the cause of the metabolic phenomena in the cells is, the more accurately we must mark the circumstances and phenomena under which they occur. One condition to them is a certain temperature, which has a maximum and a minimum. The phenomena are not produced in a temperature below 0° or above 80° R.; boiling heat destroys this faculty of the cells permanently; but the most favorable temperature is one between 10° and 32° R. Heat is evolved by the process itself.

Oxygen, or carbonic acid, in a gaseous form or lightly confined, is essentially necessary to the metabolic phenomena of the cells. The oxygen disappears and carbonic acid is formed, or _vice versa_, carbonic acid disappears, and oxygen is formed. The universality of respiration is based entirely upon this fundamental condition to the metabolic phenomena of the cells. It is so important that, as we shall see further on, even the principal varieties of form in organized bodies are occasioned by this peculiarity of the metabolic process in the cells.

Each cell is not capable of producing chemical changes in every organic substance contained in solution, but only in particular ones. The fungi of fermentation, for instance, effect no changes in any other solutions than sugar; and the spores of certain plants do not become developed in all substances. In the same manner it is probable that each cell in the animal body converts only particular constituents of the blood.

The metabolic power of the cells is arrested not only by powerful chemical actions, such as destroy organic substances in general, but also by matters which chemically are less uncongenial; for instance, concentrated solutions of neutral salts. Other substances, as arsenic, do so in less quantity. The metabolic phenomena may be altered in quality by other substances, both organic and inorganic, and a change of this kind may result even from mechanical impressions on the cells.

Such are the most essential characteristics of the fundamental powers of the cell, so far as they can as yet be deduced from the phenomena. And now, in order to comprehend distinctly in what the peculiarity of the formative process of a cell, and therefore in what the peculiarity of the essential phenomenon in the formation of organized bodies consist, we will compare this process with a phenomenon of inorganic nature as nearly as possible similar to it. Disregarding all that is specially peculiar to the formation of cells, in order to find a more general definition in which it may be included with a process occurring in inorganic nature, we may view it as a process in which a solid body of definite and regular shape is formed in a fluid at the expense of a substance held in solution by that fluid. The process of crystallization in inorganic nature comes also within this definition, and is, therefore, the nearest analogue to the formation of cells.

Let us now compare the two processes, that the difference of the organic process may be clearly manifest. First, with reference to the plastic phenomena, the forms of cells and crystals are very different. The primary forms of crystals are simple, always angular, and bounded by plane surfaces; they are regular, or at least symmetrical, and even the very varied secondary forms of crystals are almost, without exception, bounded by plane surfaces. But manifold as is the form of cells, they have very little resemblance to crystals; round surfaces predominate, and where angles occur, they are never quite sharp, and the polyhedral crystal-like form of many cells results only from mechanical causes. The structure too of cells and of crystals is different. Crystals are solid bodies, composed merely of layers placed one upon another; cells are hollow vesicles, either single, or several inclosed one within another. And if we regard the membranes of these vesicles as layers, there will still remain marks of difference between them and crystals; these layers are not in contact, but contain fluid between them, which is not the case with crystals; the layers in the cells are few, from one to three only; and they differ from each other in chemical properties, while those of crystals consist of the same chemical substance. Lastly, there is also a great difference between crystals and cells in their mode of growth. Crystals grow by apposition, the new molecules are set only upon the surface of those already deposited, but cells increase also by intussusception, that is to say, the new molecules are deposited also between those already present.

But greatly as these plastic phenomena differ in cells and in crystals, the metabolic are yet more different, or rather they are quite peculiar to cells. For a crystal to grow, it must be already present as such in the solution, and some extraneous cause must interpose to diminish its solubility. Cells, on the contrary, are capable of producing a chemical change in the surrounding fluid, of generating matters which had not previously existed in it as such, but of which only the elements were present in another combination. They therefore require no extraneous influence to effect a change of solubility; for if they can produce chemical changes in the surrounding fluid, they may also produce such substances as could not be held in solution under the existing circumstances, and therefore need no external cause of growth. If a crystal be laid in a pretty strong solution, of a substance similar even to itself, nothing ensues without our interference, or the crystal dissolves completely: the fluid must be evaporated for the crystal to increase. If a cell be laid in a solution of a substance, even different from itself, it grows and converts this substance without our aid. And this it is from which the process going on in the cells (so long as we do not separate it into its several acts) obtains that magical character, to which attaches the idea of Life.

From this we perceive how very different are the phenomena in the formation of cells and of crystals. Meanwhile, however, the points of resemblance between them should not be overlooked. They agree in this important point, that solid bodies of a certain regular shape are formed in obedience to definite laws at the expense of a substance contained in solution in a fluid; and the crystal, like the cell, is so far an active and positive agent as to cause the substances which are precipitated to be deposited on itself, and nowhere else. We must, therefore, attribute to it as well as to the cell a power to attract the substance held in solution in the surrounding fluid. It does not indeed follow that these two attractive powers, the power of crystallization--to give it a brief title--and the plastic power of the cells, are essentially the same. This could only be admitted, if it were proved that both powers acted according to the same laws. But this is seen at the first glance to be by no means the case: the phenomena in the formation of cells and crystals, are, as we have observed, very different, even if we regard merely the plastic phenomena of the cells, and leave their metabolic power (which may possibly arise from some other peculiarity of organic substance) for a time entirely out of the question.

Is it, however, possible that these distinctions are only secondary, that the power of crystallization and the plastic power of the cells are identical, and that an original difference can be demonstrated between the substance of cells and that of crystals, by which we may perceive that the substance of cells must crystallize as cells according to the laws by which crystals are formed, rather than in the shape of the ordinary crystals? It may be worth while to institute such an inquiry.

In seeking such a distinction between the substance of cells and that of crystals, we may say at once that it cannot consist in anything which the substance of cells has in common with those organic substances which crystallize in the ordinary form. Accordingly, the more complicated arrangement of the atoms of the second order in organic bodies cannot give rise to this difference; for we see in sugar, for instance, that the mode of crystallization is not altered by this chemical composition.

Another point of difference by which inorganic bodies are distinguished from at least some of the organic bodies, is the faculty of imbibition. Most organic bodies are capable of being infiltrated by water, and in such a manner that it penetrates not so much into the interspaces between the elementary tissues of the body, as into the simple structureless tissues, such as areolar tissue, &c.; so that they form an homogeneous mixture, and we can neither distinguish particles of organic matter, nor interspaces filled with water. The water occupies the infiltrated organic substances, just as it is present in a solution, and there is as much difference between the capacity for imbibition and capillary permeation, as there is between a solution and the phenomena of capillary permeation. When water soaks through a layer of glue, we do not imagine it to pass through pores, in the common sense of the term; and this is just the condition of all substances capable of imbibition. They possess, therefore, a double nature, they have a definite form like solid bodies; but like fluids, on the other hand, they are also permeable by anything held in solution. As a specifically lighter fluid poured on one specifically heavier so carefully as not to mix with it, yet gradually penetrates it, so also, every solution, when brought into contact with a membrane already infiltrated with water, bears the same relations to the membrane, as though it were a solution. And crystallization being the transition from the fluid to the solid state, we may conceive it possible, or even probable, that if bodies, capable of existing in an intermediate state between solid and fluid could be made to crystallize, a considerable difference would be exhibited from the ordinary mode of crystallization. In fact, there is nothing, which we call a crystal, composed of substance capable of imbibition; and even among organized substances, crystallization takes place only in those which are capable of imbibition, as fat, sugar, tartaric acid, &c. The bodies capable of imbibition, therefore, either do not crystallize at all, or they do so under a form so different from the crystal that they are not recognized as such.

Let us inquire what would most probably ensue if material capable of imbibition crystallized according to the ordinary laws, what varieties from the common crystals would be most likely to show themselves, assuming only that the solution has permeated through the parts of the crystal already formed, and that new molecules can therefore be deposited between them. The ordinary crystals increase only by apposition; but there may be an important difference in the mode of this apposition. If the molecules were all deposited symmetrically one upon another, we might indeed have a body of a certain external form like a crystal; but it would not have the structure of one, it would not consist of layers. The existence of this laminated structure in crystals presupposes a double kind of apposition of their molecules; for in each layer the newly-deposited molecules coalesce, and become continuous with those of the same layer already present; but those molecules which form the adjacent surfaces of two layers do not coalesce. This is a remarkable peculiarity in the formation of crystals, and we are quite ignorant of its cause. We cannot yet perceive why the new molecules, which are being deposited on the surface of a crystal (already formed up to a certain point), do not coalesce and become continuous with those already deposited, like the molecules in each separate layer, instead of forming, as they do, a new layer; and why this new layer does not constantly increase in thickness, instead of producing a second layer around the crystal, and so on. In the meantime we can do no more than express the fact in the form of a law, that the coalescing molecules are deposited rather along the surface beside each other, than in the thickness upon one another, and thus, as the breadth of the layer depends upon the size of the crystal, so also the layer can attain only a certain thickness, and beyond this, the molecules which are being deposited cannot coalesce with it, but must form a new layer.

If we now assume that bodies capable of imbibition could also crystallize, the two modes of junction of the molecules should be shown also by them. Their structure should also be laminated, at least there is no perceptible reason for a difference in this particular, as the very fact of layers being formed in common crystals shows that the molecules need not be all joined together in the most exact manner possible. The closest possible conjunction of the molecules takes place only in the separate layers. In the common crystals this occurs by apposition of the new molecules on the surface of those present and coalescence with them. In bodies capable of imbibition, a much closer union is possible, because in them the new molecules may be deposited by intussusception between those already present. It is scarcely, therefore, too bold an hypothesis to assume, that when bodies capable of imbibition crystallize, their separate layers would increase by intussusception; and that this does not happen in ordinary crystals, simply because it is impossible.

Let us then imagine a portion of the crystal to be formed: new molecules continue to be deposited, but do not coalesce with the portion of the crystal already formed; they unite with one another only, and form a new layer, which, according to analogy with the common crystals, may invest either the whole or a part of the crystal. We will assume that it invests the entire crystal. Now, although this layer be formed by the deposition of new molecules between those already present instead of by apposition, yet this does not involve any change in the law, in obedience to which the deposition of the coalescing molecules goes on more vigorously in two directions, that is, along the surface, than it does in the third direction corresponding to the thickness of the layer; that is to say, the molecules which are deposited by intussusception between those already present, must be deposited much more vigorously between those lying together along the surface of the layer than between those which lie over one another in its thickness. This deposition of molecules side by side is limited in common crystals by the size of the crystal, or by that of the surface on which the layer is formed; the coalescence of molecules therefore ceases as regards that layer, and a new one begins. But if the layers grow by intussusception in crystals capable of imbibition, there is nothing to prevent the deposition of more molecules between those which lie side by side upon the surface, even after the lamina has invested the whole crystal; it may continue to grow without the law by which the new molecules coalesce requiring to be altered. But the consequence is, that the layer becomes, in the first instance more condensed, that is, more solid substance is taken into the same space; and afterwards it will expand and separate from the completed part of the crystal so as to leave a hollow space between itself and the crystal; this space fills with fluid by imbibition, and the first-formed portion of the crystal adheres to a spot on its inner surface. Thus, in bodies capable of imbibition, instead of a new layer attached to the part of the crystal already formed, we obtain a hollow vesicle. At first this must have the shape of the body of the crystal around which it is formed, and must, therefore, be angular, if the crystal is angular. If, however, we imagine this layer to be composed of soft substance capable of imbibition, we may readily comprehend how such a vesicle must very soon become round or oval. But the first-formed part of the crystal also consists of substance capable of imbibition, so that it is very doubtful whether it must have an angular form at all. In common crystals atoms of some one particular substance are deposited together, and we can understand how a certain angular form of the crystal may result if these atoms have a certain form, or if in certain axes they attract each other differently. But in bodies capable of imbibition, an atom of one substance is not set upon another atom of the same substance, but atoms of water come between; atoms of water, which are not united with an atom of solid substance, so as to form a compound atom, as in the water of crystallization, but which exist in some other unknown manner between the atoms of solid substance. It is not possible, therefore, to determine whether that part of the crystal which is first formed must have an angular figure or not.

An ordinary crystal consists of a number of laminæ; when so small as to be but just discernible, it has the form which the whole crystal afterwards exhibits, at least as far as regards the angles; we must therefore suppose that the first layer is formed around a very small corpuscle, which is of the same shape as the subsequent crystal. We will call this the primitive corpuscle. It is doubtful what may be the shape of this corpuscle in the crystals which are capable of imbibition. The first layer, then, is formed around the corpuscle in the way mentioned; it grows by intussusception, and thus forms a hollow, round or oval vesicle, to the inner surface of which the primitive corpuscle adheres. As all the new molecules that are being deposited may be placed in this layer without any alteration being required in the law which regulates the coalescence of the molecules during crystallization, we must conclude that it remains the only layer, and becomes greatly expanded, so as to represent all the layers of an ordinary crystal. It is, however, a question whether there may not exist some reasons why several layers can be formed. We can certainly conceive such to be the case. The quantity of the solid substance that must crystallize in a given time, depends upon the concentration of the fluid; the number of molecules that may, in accordance with the law already mentioned, be deposited in the layer in a given time depends upon the quantity of the solution which can penetrate the membrane by imbibition during that time. If in consequence of the concentration of the fluid there must be more precipitated in the time than can penetrate the membrane, it can only be deposited as a new layer on the outer surface of the vesicle. When this second layer is formed, the new molecules are deposited in it, and it rapidly becomes expanded into a vesicle, on the inner surface of which the first vesicle lies with its primitive corpuscle. The first vesicle now either does not grow at all, or at any rate much more slowly, and then only when the endosmosis into the cavity of the second vesicle proceeds so rapidly that all that might be precipitated while passing through it, is not deposited. The second vesicle, when it is developed at all, must needs be developed relatively with more rapidity than the first; for as the solution is in the most concentrated state at the beginning, the necessity for the formation of a second layer then occurs sooner; but when it is formed, the concentration of the fluid is diminished, and this necessity occurs either later or not at all. It is possible, however, that even a third, or fourth, and more, may be formed; but the outermost layer must always be relatively the most vigorously developed; for when the concentration of the solution is only so strong, that all that must be deposited in a certain time, can be deposited in the outermost layer, it is all applied to the increase of this layer.

Such, then, would be the phenomena under which substances capable of imbibition would probably crystallize, if they did so at all. I say probably, for our incomplete knowledge of crystallization and the faculty of imbibition, does not as yet admit of our saying anything positively _a priori_. It is, however, obvious that these are the principal phenomena attending the formation of cells. They consist always of substance capable of imbibition; the first part formed is a small corpuscle, not angular (nucleolus), around this a lamina is deposited (nucleus), which advances rapidly in its growth, until a second lamina (cell) is formed around it. This second now grows more quickly and expands into a vesicle, as indeed often happens with the first layer. In some rarer instances only one layer is formed; in others, again, there are three. The only other difference in the formation of cells is, that the separate layers do not consist of the same chemical substance, while a common crystal is always composed of one material. In instituting a comparison, therefore, between the formation of cells and crystallization, the above-mentioned differences in form, structure, and mode of growth fall altogether to the ground. If crystals were formed from the same substance as cells, they would probably, in these respects, be subject to the same conditions as the cells. Meanwhile the metabolic phenomena, which are entirely absent in crystals, still indicate essential distinctions.

Should this important difference between the mode of formation of cells and crystals lead us to deny all intimate connexion of the two processes, the comparison of the two may serve at least to give a clear representation of the cell-life. The following may be conceived to be the state of the matter: the material of which the cells are composed is capable of producing chemical changes in the substance with which it is in contact, just as the well-known preparation of platinum converts alcohol into acetic acid. This power is possessed by every part of the cell. Now, if the cytoblastema be so changed by a cell already formed, that a substance is produced which cannot become attached to that cell, it immediately crystallizes as the central nucleolus of a new cell. And then this converts the cytoblastema in the same manner. A portion of that which is converted may remain in the cytoblastema in solution, or may crystallize as the commencement of new cells; another portion, the cell-substance, crystallizes around the central corpuscle. The cell-substance is either soluble in the cytoblastema, and crystallizes from it, so soon as the latter becomes saturated with it; or else it is insoluble, and crystallizes at the time of its formation, according to the laws of crystallization of bodies capable of imbibition mentioned above, forming in this manner one or more layers around the central corpuscle, and so on. If we conceive the above to represent the mode of formation of cells, we regard the plastic power of the cells as identical with the power by which crystals grow. According to the foregoing description of the crystallization of bodies capable of imbibition, the most important plastic phenomena of the cells are certainly satisfactorily explained. But let us see if this comparison agrees with all the characteristics of the plastic power of the cells.

The attractive power of the cells does not always operate symmetrically; the deposition of new molecules may be more vigorous in particular spots, and thus produce a change in the form of the cell. This is quite analogous to what happens in crystals; for although in them an angle is never altered, there may be much more material deposited on some surfaces than on others; and thus, for instance, a quadrilateral prism may be formed out of a cube. In this case new layers are deposited on one, or on two opposite sides of a cube. Now, if one layer in cells represent a number of layers in a common crystal, it may be easily perceived that instead of several new layers being formed on two opposite surfaces of a cell, the one layer would grow more at those spots, and thus a round cell would be elongated into a fibre; and so with the other changes of form. Division of the cells can have no analogue in common crystals, because that which is once deposited is incapable of any further change. But this phenomenon may be made to accord with the representation of crystals capable of imbibition.... And if we ascribe to a layer of a crystal capable of imbibition the power of producing chemical changes in organic substances, we can very well understand also the origin of secondary deposits on its inner surface as they occur in cells. For if, in accordance with the laws of crystallization, the lamina has become expanded into a vesicle, and its cavity has become filled by imbibition with a solution of organic substance, there may be materials formed by means of the converting influence of the lamina, which cannot any longer be held in solution. These may, then, either crystallize within the vesicle, as new crystals capable of imbibition under the form of cells; or if they are allied to the substance of the vesicle, they may so crystallize as to form part of the system of the vesicle itself: the latter may occur in two ways, the new matters may be applied to the increase of the vesicle, or they may form new layers on its inner surface from the same cause which led to the first formation of the vesicle itself as a layer. In the cells of plants these secondary deposits have a spiral arrangement. This is a very important fact, though the laws of crystallization do not seem to account for the absolute necessity of it. If, however, it could be mathematically proved from the laws of the crystallization of inorganic bodies, that under the altered circumstances in which bodies capable of imbibition are placed, these deposits must be arranged in spiral forms, it might be asserted without hesitation that the plastic power of cells and the fundamental powers of crystals are identical.

We come now, however, to some peculiarities in the plastic power of cells, to which we might, at first sight, scarcely expect to find anything analogous in crystals. The attractive power of the cells manifests a certain degree of election in its operation; it does not attract every substance present in the cytoblastema, but only particular ones; and here a muscle-cell, there a fat-cell, is generated from the same fluid, the blood. Yet crystals afford us an example of a precisely similar phenomenon, and one which has already been frequently adduced as analogous to assimilation. If a crystal of nitre be placed in a solution of nitre and sulphate of soda, only the nitre crystallizes; when a crystal of sulphate of soda is put in, only the sulphate of soda crystallizes. Here, therefore, there occurs just the same selection of the substance to be attracted.

We observed another law attending the development of the plastic phenomena in the cells, viz. that a more concentrated solution is requisite for the first formation of a cell than for its growth when already formed, a law upon which the difference between organized and unorganized tissues is based. In ordinary crystallization the solution must be more than saturated for the process to begin. But when it is over, there remains a mother lye, according to Thénard, which is no longer saturated at the same temperature. This phenomenon accords precisely with the cells; it shows that a more concentrated solution is requisite for the commencement of crystallization than for the increase of a crystal already formed. The fact has indeed been disputed by Thomson; but if, in the undisputed experiment quoted above, the crystal of sulphate of soda attracts the dissolved sulphate of soda rather than the dissolved nitre, and _vice versa_, the crystal of nitre attracts the dissolved nitre more than the dissolved sulphate of soda, it follows that a crystal does attract a salt held in solution, because the experiment proves that there are degrees of this attraction. But if there be such an attraction exerted by a crystal, then the introduction of a crystal into a solution of a salt, affords an efficient cause for the deposition of this salt, which does not exist when no crystal is introduced. The solution must therefore be more concentrated in the latter case than in the former, though the difference be so slight as not to be demonstrable by experiment. It would not, however, be superfluous to repeat the experiments. In the instance of crystals capable of imbibition, this difference may be considerably augmented, since the attraction of molecules may increase perhaps considerably by the penetrating of the solution between those already deposited.

We see then how all the plastic phenomena in the cells may be compared with phenomena which, in accordance with the ordinary laws of crystallization, would probably appear if bodies capable of imbibition could be brought to crystallize. So long as the object of such a comparison were merely to render the representation of the process by which cells are formed more clear, there could not be much urged against it; it involves nothing hypothetical, since it contains no explanation; no assertion is made that the fundamental power of the cells really has something in common with the power by which crystals are formed. We have, indeed, compared the growth of organisms with crystallization, in so far as in both cases solid substances are deposited from a fluid, but we have not therefore asserted the identity of the fundamental powers. So far we have not advanced beyond the data, beyond a certain simple mode of representing the facts.

The question is, however, whether the exact accordance of the phenomena would not authorize us to go further. If the formation and growth of the elementary particles of organisms have nothing more in common with crystallization than merely the deposition of solid substances from out of a fluid, there is certainly no reason for assuming any more intimate connexion of the two processes. But we have seen, first, that the laws which regulate the deposition of the molecules forming the elementary particles of organisms are the same for all elementary parts; that there is a common principle in the development of all elementary parts, namely, that of the formation of cells; it was then shown that the power which induced the attachment of the new molecules did not reside in the entire organism, but in the separated elementary particles (this we called the plastic power of the cells); lastly, it was shown that the laws, according to which the new molecules combine to form cells, are (so far as our incomplete knowledge of the laws of crystallization admits of our anticipating their probability) the same as those by which substances capable of imbibition would crystallize. Now the cells do, in fact, consist only of material capable of imbibition; should we not then be justified in putting forth the proposition, that the formation of the elementary parts of organisms is nothing but a crystallization of substance, capable of imbibition, and the organism nothing but an aggregate of such crystals capable of imbibition?

To advance so important a point as absolutely true, would certainly need the clearest proof; but it cannot be said that even the premises which have been set forth have in all points the requisite force. For too little is still known of the cause of crystallization to predict with safety (as was attempted above) what would follow if a substance capable of imbibition were to crystallize. And if these premises were allowed, there are two other points which must be proved in order to establish the proposition in question: 1. That the metabolic phenomena of the cells, which have not been referred to in the foregoing argument, are as much the necessary consequence of the faculty of imbibition, or of some other peculiarity of the substance of cells, as the plastic phenomena are. 2. That if a number of crystals capable of imbibition are formed, they must combine according to certain laws so as to form a systematic whole, similar to an organism. Both these points must be clearly proved, in order to establish the truth of the foregoing view. But it is otherwise if this view be adduced merely as an hypothesis, which may serve as a guide for new investigations. In such case the inferences are sufficiently probable to justify such an hypothesis, if only the two points just mentioned can be shown to accord with it.

With reference to the first of these points, it would certainly be impossible, in our ignorance as to the cause of chemical phenomena in general, to prove that a crystal capable of imbibition must produce chemical changes in substances surrounding it; but then we could not infer, from the manner in which spongy platinum is formed, that it would act so peculiarly upon oxygen and hydrogen. But in order to render this view tenable as a possible hypothesis, it is only necessary to see that it may be a consequence. It cannot be denied that it may: there are several reasons for it, though they certainly are but weak. For instance, since all cells possess this metabolic power, it is more likely to depend on a certain position of the molecules, which in all probability is essentially the same in all cells, than on the chemical combination of the molecules, which is very different in different cells. The presence, too, of different substances on the inner and outer surface of the cell-membrane in some measure implies that a certain direction of the axes of the atoms may be essential to the metabolic phenomena of the cells. I think, therefore, that the cause of the metabolic phenomena resides in that definite mode of arrangement of the molecules which occurs in crystals, combined with the capacity which the solution has to penetrate between these regularly deposited molecules (by means of which, presuming the molecules to possess polarity, a sort of galvanic pile will be formed), and that the same phenomena would be observed in an ordinary crystal, if it could be rendered capable of imbibition. And then perhaps the differences of quality in the metabolic phenomena depend upon their chemical composition.

In order to render tenable the hypothesis contained in the second point, it is merely necessary to show that crystals capable of imbibition can unite with one another according to certain laws. If at their first formation all crystals were isolated, if they held no relation whatever to each other, the view would leave entirely unexplained how the elementary parts of organisms, that is, the crystals in question, become united to form a whole. It is therefore necessary to show that crystals do unite with each other according to certain laws, in order to perceive, at least, the possibility of their uniting also to form an organism, without the need of any further combining power. But there are many crystals in which a union of this kind, according to certain laws, is indisputable; indeed they often form a whole, so like an organism in its entire form, that groups of crystals are known in common life by the names of flowers, trees, etc. I need only refer to the ice-flowers on the windows, or to the lead-tree, etc. In such instances a number of crystals arrange themselves in groups around others, which form an axis. If we consider the contact of each crystal with the surrounding fluid to be an indispensable condition to the growth of crystals which are not capable of imbibition, but that those which are capable of imbibition, in which the solution can penetrate whole layers of crystals, do not require this condition, we perceive that the similarity between organisms and these aggregations of crystals is as great as could be expected with such difference of substance. As most cells require for the production of their metabolic phenomena, not only their peculiar nutrient fluid, but also the access of oxygen and the power of exhaling carbonic acid, or _vice versa_; so, on the other hand, organisms in which there is no circulation of respiratory fluid, or in which at least it is not sufficient, must be developed in such a way as to present as extensive a surface as possible to the atmospheric air. This is the condition of plants, which require for their growth that the individual cells should come into contact with the surrounding medium in a similar manner, if not in the same degree as occurs in a crystal tree, and in them indeed the cells unite into a whole organism in a form much resembling a crystal tree. But in animals the circulation renders the contact of the individual cells with the surrounding medium superfluous, and they may have more compact forms, even though the laws by which the cells arrange themselves are essentially the same.

The view then that organisms are nothing but the form under which substances capable of imbibition crystallize, appears to be compatible with the most important phenomena of organic life, and may be so far admitted, that it is a possible hypothesis; or attempt towards an explanation of these phenomena. It involves very much that is uncertain and paradoxical, but I have developed it in detail, because it may serve as a guide for new investigations. For even if no relation between crystallization and the growth of organisms be admitted in principle, this view has the advantage of affording a distinct representation of the organic processes; an indispensable requisite for the institution of new inquiries in a systematic manner, or for testing by the discovery of new facts a mode of explanation which harmonizes with phenomena already known.

FOOTNOTES:

[Footnote 33: Translated from _Mikroskopische Untersuchungen über die Wachstum der Tiere und der Pflanzen_ (Berlin, 1839) by Henry Smith in the _Publications of the Sydenham Society_ (1847).]

XXXI

HERMANN VON HELMHOLTZ

1821-1894

_Hermann von Helmholtz, born at Potsdam, Prussia, August 31, 1821, studied medicine at the University of Berlin, from which he received his degree in 1842. He then entered the German Army as surgeon and in 1847 published his paper on “The Conservation of Energy,” which summarized historically the development of the idea. In 1849 he was appointed professor of physiology and general pathology at Königsberg. In 1855 he was called to Bonn, and in 1858 was elected to the chair of physiology at Heidelberg._

_In 1851 he invented the ophthalmoscope and later at Heidelberg he continued his researches in the subject of sight, and also cleared up the problem of the mechanical causes of sound. In 1871 he was appointed professor of physics at the University of Berlin, where he remained until his death, September 8, 1894._

THE CONSERVATION OF ENERGY[34]

A new conquest of very general interest has been recently made by natural philosophy. In the following pages I will endeavour to give a notion of the nature of this conquest. It has reference to a new and universal natural law, which rules the action of natural forces in their mutual relations towards each other, and is as influential on our theoretic views of natural processes as it is important in their technical applications.