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In the early days of the cell-theory, more than seventy years ago, Goodsir was wont to speak of cells as “centres of growth” or “centres of nutrition,” and to consider them as essentially “centres of force.” He looked forward to a time when the forces connected with the cell should be particularly investigated: when, that is to say, minute anatomy should be studied in its dynamical aspect. “When this branch of enquiry,” he says “shall have been opened up, we shall expect to have a science of organic forces, having direct relation to anatomy, the science of organic forms198.” And likewise, long afterwards, Giard contemplated a science of morphodynamique—but still looked upon it as forming so guarded and hidden a “territoire scientifique, que la plupart des naturalistes de nos jours ne le verront que comme Moïse vit la terre promise, seulement de loin et sans pouvoir y entrer199.”

To the external forms of cells, and to the forces which produce and modify these forms, we shall pay attention in a later chapter. But there are forms and con­fi­gur­a­tions of matter within the cell, which also deserve to be studied with due regard to the forces, known or unknown, of whose resultant they are the visible expression.

In the long interval since Goodsir’s day, the visible structure, the conformation and configuration, of the cell, has been studied far more abundantly than the purely dynamic problems that are associated therewith. The overwhelming progress of microscopic observation has multiplied our knowledge of cellular and intracellular structure; and to the multitude of visible structures it {157} has been often easier to attribute virtues than to ascribe intelligible functions or modes of action. But here and there nevertheless, throughout the whole literature of the subject, we find recognition of the inevitable fact that dynamical problems lie behind the morphological problems of the cell.

Bütschli pointed out forty years ago, with emphatic clearness, the failure of morphological methods, and the need for physical methods, if we were to penetrate deeper into the essential nature of the cell200. And such men as Loeb and Whitman, Driesch and Roux, and not a few besides, have pursued the same train of thought and similar methods of enquiry.

Whitman201, for instance, puts the case in a nutshell when, in speaking of the so-called “caryokinetic” phenomena of nuclear division, he reminds us that the leading idea in the term “caryokinesis” is motion—“motion viewed as an exponent of forces residing in, or acting upon, the nucleus. It regards the nucleus as a seat of energy, which displays itself in phenomena of motion202.”

In short it would seem evident that, except in relation to a dynamical in­ves­ti­ga­tion, the mere study of cell structure has but little value of its own. That a given cell, an ovum for instance, contains this or that visible substance or structure, germinal vesicle or germinal spot, chromatin or achromatin, chromosomes or centrosomes, obviously gives no explanation of the activities of the cell. And in all such hypotheses as that of “pangenesis,” in all the theories which attribute specific properties to micellae, {158} idioplasts, ids, or other constituent particles of protoplasm or of the cell, we are apt to fall into the error of attributing to matter what is due to energy and is manifested in force: or, more strictly speaking, of attributing to material particles individually what is due to the energy of their collocation.

The tendency is a very natural one, as knowledge of structure increases, to ascribe particular virtues to the material structures themselves, and the error is one into which the disciple is likely to fall, but of which we need not suspect the master-mind. The dynamical aspect of the case was in all probability kept well in view by those who, like Goodsir himself, first attacked the problem of the cell and originated our conceptions of its nature and functions.

But if we speak, as Weismann and others speak, of an “hereditary substance,” a substance which is split off from the parent-body, and which hands on to the new generation the char­ac­teris­tics of the old, we can only justify our mode of speech by the assumption that that particular portion of matter is the essential vehicle of a particular charge or distribution of energy, in which is involved the capability of producing motion, or of doing “work.”

For, as Newton said, to tell us that a thing “is endowed with an occult specific quality, by which it acts and produces manifest effects, is to tell us nothing; but to derive two or three general principles of motion203 from phenomena would be a very great step in philosophy, though the causes of these principles were not yet discovered.” The things which we see in the cell are less important than the actions which we recognise in the cell; and these latter we must especially scrutinize, in the hope of discovering how far they may be attributed to the simple and well-known physical forces, and how far they be relevant or irrelevant to the phenomena which we associate with, and deem essential to, the manifestation of life. It may be that in this way we shall in time draw nigh to the recognition of a specific and ultimate residuum. {159}

And lacking, as we still do lack, direct knowledge of the actual forces inherent in the cell, we may yet learn something of their distribution, if not also of their nature, from the outward and inward configuration of the cell, and from the changes taking place in this configuration; that is to say from the movements of matter, the kinetic phenomena, which the forces in action set up.

The fact that the germ-cell develops into a very complex structure, is no absolute proof that the cell itself is structurally a very complicated mechanism: nor yet, though this is somewhat less obvious, is it sufficient to prove that the forces at work, or latent, within it are especially numerous and complex. If we blow into a bowl of soapsuds and raise a great mass of many-hued and variously shaped bubbles, if we explode a rocket and watch the regular and beautiful configuration of its falling streamers, if we consider the wonders of a limestone cavern which a filtering stream has filled with stalactites, we soon perceive that in all these cases we have begun with an initial system of very slight complexity, whose structure in no way foreshadowed the result, and whose comparatively simple intrinsic forces only play their part by complex interaction with the equally simple forces of the surrounding medium. In an earlier age, men sought for the visible embryo, even for the homunculus, within the reproductive cells; and to this day, we scrutinize these cells for visible structure, unable to free ourselves from that old doctrine of “pre-formation204.”

Moreover, the microscope seemed to substantiate the idea (which we may trace back to Leibniz205 and to Hobbes206), that there is no limit to the mechanical complexity which we may postulate in an organism, and no limit, therefore, to the hypotheses which we may rest thereon.

But no microscopical examination of a stick of sealing-wax, no study of the material of which it is composed, can enlighten {160} us as to its electrical manifestations or properties. Matter of itself has no power to do, to make, or to become: it is in energy that all these potentialities reside, energy invisibly associated with the material system, and in interaction with the energies of the surrounding universe.

That “function presupposes structure” has been declared an accepted axiom of biology. Who it was that so formulated the aphorism I do not know; but as regards the structure of the cell it harks back to Brücke, with whose demand for a mechanism, or organisation, within the cell histologists have ever since been attempting to comply207. But unless we mean to include thereby invisible, and merely chemical or molecular, structure, we come at once on dangerous ground. For we have seen, in a former chapter, that some minute “organisms” are already known of such all but infinitesimal magnitudes that everything which the morphologist is accustomed to conceive as “structure” has become physically impossible; and moreover recent research tends generally to reduce, rather than to extend, our conceptions of the visible structure necessarily inherent in living protoplasm. The microscopic structure which, in the last resort or in the simplest cases, it seems to shew, is that of a more or less viscous colloid, or rather mixture of colloids, and nothing more. Now, as Clerk Maxwell puts it, in discussing this very problem, “one material system can differ from another only in the configuration and motion which it has at a given instant208.” If we cannot assume differences in structure, we must assume differences in motion, that is to say, in energy. And if we cannot do this, then indeed we are thrown back upon modes of reasoning unauthorised in physical science, and shall find ourselves constrained to assume, or to “admit, that the properties of a germ are not those of a purely material system.” {161}

But we are by no means necessarily in this dilemma. For though we come perilously near to it when we contemplate the lowest orders of magnitude to which life has been attributed, yet in the case of the ordinary cell, or ordinary egg or germ which is going to develop into a complex organism, if we have no reason to assume or to believe that it comprises an intricate “mechanism,” we may be quite sure, both on direct and indirect evidence, that, like the powder in our rocket, it is very heterogeneous in its structure. It is a mixture of substances of various kinds, more or less fluid, more or less mobile, influenced in various ways by chemical, electrical, osmotic, and other forces, and in their admixture separated by a multitude of surfaces, or boundaries, at which these, or certain of these forces are made manifest.

Indeed, such an arrangement as this is already enough to constitute a “mechanism”; for we must be very careful not to let our physical or physiological concept of mechanism be narrowed to an interpretation of the term derived from the delicate and complicated contrivances of human skill. From the physical point of view, we understand by a “mechanism” whatsoever checks or controls, and guides into determinate paths, the workings of energy; in other words, whatsoever leads in the degradation of energy to its manifestation in some determinate form of work, at a stage short of that ultimate degradation which lapses in uniformly diffused heat. This, as Warburg has well explained, is the general effect or function of the physiological machine, and in particular of that part of it which we call “cell-structure209.” The normal muscle-cell is something which turns energy, derived from oxidation, into work; it is a mechanism which arrests and utilises the chemical energy of oxidation in its downward course; but the same cell when injured or disintegrated, loses its “usefulness,” and sets free a greatly increased proportion of its energy in the form of heat.

But very great and wonderful things are done after this manner by means of a mechanism (whether natural or artificial) of extreme simplicity. A pool of water, by virtue of its surface, {162} is an admirable mechanism for the making of waves; with a lump of ice in it, it becomes an efficient and self-contained mechanism for the making of currents. The great cosmic mechanisms are stupendous in their simplicity; and, in point of fact, every great or little aggregate of heterogeneous matter (not identical in “phase”) involves, ipso facto, the essentials of a mechanism. Even a non-living colloid, from its intrinsic heterogeneity, is in this sense a mechanism, and one in which energy is manifested in the movement and ceaseless rearrangement of the constituent particles. For this reason Graham (if I remember rightly) speaks somewhere or other of the colloid state as “the dynamic state of matter”; or in the same philosopher’s phrase (of which Mr. Hardy210 has lately reminded us), it possesses “energia211.”

Let us turn then to consider, briefly and dia­gram­ma­ti­cally, the structure of the cell, a fertilised germ-cell or ovum for instance, not in any vain attempt to correlate this structure with the structure or properties of the resulting and yet distant organism; but merely to see how far, by the study of its form and its changing internal configuration, we may throw light on certain forces which are for the time being at work within it.

We may say at once that we can scarcely hope to learn more of these forces, in the first instance, than a few facts regarding their direction and magnitude; the nature and specific identity of the force or forces is a very different matter. This latter problem is likely to be very difficult of elucidation, for the reason, among others, that very different forces are often very much alike in their outward and visible manifestations. So it has come to pass that we have a multitude of discordant hypotheses as to the nature of the forces acting within the cell, and producing, in cell division, the “caryokinetic” figures of which we are about to speak. One student may, like Rhumbler, choose to account for them by an hypothesis of mechanical traction, acting on a reticular web of protoplasm212; another, like Leduc, may shew us how in {163} many of their most striking features they may be admirably simulated by the diffusion of salts in a colloid medium; others again, like Gallardo213 and Hartog, and Rhumbler (in his earlier papers)214, insist on their resemblance to the phenomena of electricity and magnetism215; while Hartog believes that the force in question is only analogous to these, and has a specific identity of its own216. All these conflicting views are of secondary importance, so long as we seek only to account for certain con­fi­gur­a­tions which reveal the direction, rather than the nature, of a force. One and the same system of lines of force may appear in a field of magnetic or of electrical energy, of the osmotic energy of diffusion, of the gravitational energy of a flowing stream. In short, we may expect to learn something of the pure or abstract dynamics, long before we can deal with the special physics of the cell. For indeed (as Maillard has suggested), just as uniform expansion about a single centre, to whatsoever physical cause it may be due will lead to the configuration of a sphere, so will any two centres or foci of potential (of whatsoever kind) lead to the con­fi­gur­a­tions with which Faraday made us familiar under the name of “lines of force217”; and this is as much as to say that the phenomenon, {164} though physical in the concrete, is in the abstract purely math­e­mat­i­cal, and in its very essence is neither more nor less than a property of three-dimensional space.

But as a matter of fact, in this instance, that is to say in trying to explain the leading phenomena of the caryokinetic division of the cell, we shall soon perceive that any explanation which is based, like Rhumbler’s, on mere mechanical traction, is obviously inadequate, and we shall find ourselves limited to the hypothesis of some polarised and polarising force, such as we deal with, for instance, in the phenomena of magnetism or electricity.

Let us speak first of the cell itself, as it appears in a state of rest, and let us proceed afterwards to study the more active phenomena which accompany its division.

Our typical cell is a spherical body; that is to say, the uniform surface-tension at its boundary is balanced by the outward resistance of uniform forces within. But at times the surface-tension may be a fluctuating quantity, as when it produces the rhythmical contractions or “Ransom’s waves” on the surface of a trout’s egg; or again, while the egg is in contact with other bodies, the surface-tension may be locally unequal and variable, giving rise to an amoeboid figure, as in the egg of Hydra218.

Within the ovum is a nucleus or germinal vesicle, also spherical, and consisting as a rule of portions of “chromatin,” aggregated together within a more fluid drop. The fact has often been commented upon that, in cells generally, there is no correlation of form (though there apparently is of size) between the nucleus and the “cytoplasm,” or main body of the cell. So Whitman219 remarks that “except during the process of division the nucleus seldom departs from its typical spherical form. It divides and sub-divides, ever returning to the same round or oval form. … How different with the cell. It preserves the spherical form as rarely as the nucleus departs from it. Variation in form marks the beginning and the end of every important chapter in its {165} history.” On simple dynamical grounds, the contrast is easily explained. So long as the fluid substance of the nucleus is qualitatively different from, and incapable of mixing with, the fluid or semi-fluid protoplasm which surrounds it, we shall expect it to be, as it almost always is, of spherical form. For, on the one hand, it is bounded by a liquid film, whose surface-tension is uniform; and on the other, it is immersed in a medium which transmits on all sides a uniform fluid pressure220. For a similar reason the contractile vacuole of a Protozoon is spherical in form: it is just a “drop” of fluid, bounded by a uniform surface-tension and through whose boundary-film diffusion is taking place. But here, owning to the small difference between the fluid constituting, and that surrounding, the drop, the surface-tension equi­lib­rium is unstable; it is apt to vanish, and the rounded outline of the drop, like a burst bubble, disappears in a moment221. The case of the spherical nucleus is closely akin to the spherical form of the yolk within the bird’s egg222. But if the substance of the cell acquire a greater solidity, as for instance in a muscle {166} cell, or by reason of mucous accumulations in an epithelium cell, then the laws of fluid pressure no longer apply, the external pressure on the nucleus tends to become unsymmetrical, and its shape is modified accordingly. “Amoeboid” movements may be set up in the nucleus by anything which disturbs the symmetry of its own surface-tension. And the cases, as in many Rhizopods, where “nuclear material” is scattered in small portions throughout the cell instead of being aggregated in a single nucleus, are probably capable of very simple explanation by supposing that the “phase difference” (as the chemists say) between the nuclear and the protoplasmic substance is comparatively slight, and the surface-tension which tends to keep them separate is correspondingly small223.

It has been shewn that ordinary nuclei, isolated in a living or fresh state, easily flow together; and this fact is enough to suggest that they are aggregations of a particular substance rather than bodies deserving the name of particular organs. It is by reason of the same tendency to confluence or aggregation of particles that the ordinary nucleus is itself formed, until the imposition of a new force leads to its disruption.

Apart from that invisible or ultra-microscopic heterogeneity which is inseparable from our notion of a “colloid,” there is a visible heterogeneity of structure within both the nucleus and the outer protoplasm. The former, for instance, contains a rounded nucleolus or “germinal spot,” certain conspicuous granules or strands of the peculiar substance called chromatin, and a coarse meshwork of a protoplasmic material known as “linin” or achromatin; the outer protoplasm, or cytoplasm, is generally believed to consist throughout of a sponge-work, or rather alveolar meshwork, of more and less fluid substances; and lastly, there are generally to be detected one or more very minute bodies, usually in the cytoplasm, sometimes within the nucleus, known as the centrosome or centrosomes.

The morphologist is accustomed to speak of a “polarity” of {167} the cell, meaning thereby a symmetry of visible structure about a particular axis. For instance, whenever we can recognise in a cell both a nucleus and a centrosome, we may consider a line drawn through the two as the morphological axis of polarity; in an epithelium cell, it is obvious that the cell is morphologically symmetrical about a median axis passing from its free surface to its attached base. Again, by an extension of the term “polarity,” as is customary in dynamics, we may have a “radial” polarity, between centre and periphery; and lastly, we may have several apparently independent centres of polarity within the single cell. Only in cells of quite irregular, or amoeboid form, do we fail to recognise a definite and symmetrical “polarity.” The morphological “polarity” is accompanied by, and is but the outward expression (or part of it) of a true dynamical polarity, or distribution of forces; and the “lines of force” are rendered visible by concatenation of particles of matter, such as come under the influence of the forces in action.

When the lines of force stream inwards from the periphery towards a point in the interior of the cell, the particles susceptible of attraction either crowd towards the surface of the cell, or, when retarded by friction, are seen forming lines or “fibrillae” which radiate outwards from the centre and constitute a so-called “aster.” In the cells of columnar or ciliated epithelium, where the sides of the cell are symmetrically disposed to their neighbours but the free and attached surfaces are very diverse from one another in their external relations, it is these latter surfaces which constitute the opposite poles; and in accordance with the parallel lines of force so set up, we very frequently see parallel lines of granules which have ranged themselves perpendicularly to the free surface of the cell (cf. fig. 97).

A simple manifestation of “polarity” may be well illustrated by the phenomenon of diffusion, where we may conceive, and may automatically reproduce, a “field of force,” with its poles and visible lines of equipotential, very much as in Faraday’s conception of the field of force of a magnetic system. Thus, in one of Leduc’s experiments224, if we spread a layer of salt solution over a level {168} plate of glass, and let fall into the middle of it a drop of indian ink, or of blood, we shall find the coloured particles travelling outwards from the central “pole of concentration” along the lines of diffusive force, and so mapping out for us a “monopolar field” of diffusion: and if we set two such drops side by side, their lines of diffusion will oppose, and repel, one another. Or, instead of the uniform layer of salt solution, we may place at a little distance from one another a grain of salt and a drop of blood, representing two opposite poles: and so obtain a picture of a “bipolar field” of diffusion. In either case, we obtain results closely analogous to the “morphological,” but really dynamical, polarity of the organic cell. But in all probability, the dynamical polarity, or asymmetry of the cell is a very complicated phenomenon: for the obvious reason that, in any system, one asymmetry will tend to beget another. A chemical asymmetry will induce an inequality of surface-tension, which will lead directly to a modification of form; the chemical asymmetry may in turn be due to a process of electrolysis in a polarised electrical field; and again the chemical heterogeneity may be intensified into a chemical “polarity,” by the tendency of certain substances to seek a locus of greater or less surface-energy. We need not attempt to grapple with a subject so complicated, and leading to so many problems which lie beyond the sphere of interest of the morphologist. But yet the morphologist, in his study of the cell, cannot quite evade these important issues; and we shall return to them again when we have dealt somewhat with the form of the cell, and have taken account of some of the simpler phenomena of surface-tension.

We are now ready, and in some measure prepared, to study the numerous and complex phenomena which usually accompany the division of the cell, for instance of the fertilised egg.

Division of the cell is essentially accompanied, and preceded, by a change from radial or monopolar to a definitely bipolar polarity.

In the hitherto quiescent, or apparently quiescent cell, we perceive certain movements, which correspond precisely to what must accompany and result from a “polarisation” of forces within the {169} cell: of forces which, whatever may be their specific nature, at least are capable of polarisation, and of producing consequent attraction or repulsion between charged particles of matter. The opposing forces which were distributed in equi­lib­rium throughout the substance of the cell become focussed at two “centrosomes,” which may or may not be already distinguished as visible portions of matter; in the egg, one of these is always near to, and the other remote from, the “animal pole” of the egg, which pole is visibly as well as chemically different from the other, and is the region in which the more rapid and conspicuous developmental changes will presently begin. Between the two centrosomes, a spindle-shaped

Fig. 41. Caryokinetic figure in a dividing cell (or blastomere) of the Trout’s egg. (After Prenant, from a preparation by Prof. P. Bouin.)

figure appears, whose striking resemblance to the lines of force made visible by iron-filings between the poles of a magnet, was at once recognised by Hermann Fol, when in 1873 he witnessed for the first time the phenomenon in question. On the farther side of the centrosomes are seen star-like figures, or “asters,” in which we can without difficulty recognise the broken lines of force which run externally to those stronger lines which lie nearer to the polar axis and which constitute the “spindle.” The lines of force are rendered visible or “material,” just as in the experiment of the iron-filings, by the fact that, in the heterogeneous substance of the cell, certain portions of matter are more “permeable” to the acting force than the rest, become themselves polarised after the {170} fashion of a magnetic or “paramagnetic” body, arrange themselves in an orderly way between the two poles of the field of force, cling to one another as it were in threads225, and are only prevented by the friction of the surrounding medium from approaching and congregating around the adjacent poles.

As the field of force strengthens, the more will the lines of force be drawn in towards the interpolar axis, and the less evident will be those remoter lines which constitute the terminal, or extrapolar, asters: a clear space, free from materialised lines of force, may thus tend to be set up on either side of the spindle, the so-called “Bütschli space” of the histologists226. On the other hand, the lines of force constituting the spindle will be less concentrated if they find a path of less resistance at the periphery of the cell: as happens, in our experiment of the iron-filings, when we encircle the field of force with an iron ring. On this principle, the differences observed between cells in which the spindle is well developed and the asters small, and others in which the spindle is weak and the asters enormously developed, can be easily explained by variations in the potential of the field, the large, conspicuous asters being probably correlated with a marked permeability of the surface of the cell.

The visible field of force, though often called the “nuclear spindle,” is formed outside of, but usually near to, the nucleus. Let us look a little more closely into the structure of this body, and into the changes which it presently undergoes.

Within its spherical outline (Fig. 42), it contains an “alveolar” {171} meshwork (often described, from its appearance in optical section, as a “reticulum”), consisting of more solid substances, with more fluid matter filling up the interalveolar meshes. This phenomenon is nothing else than what we call in ordinary language, a “froth” or a “foam.” It is a surface-tension phenomenon, due to the interacting surface-tensions of two intermixed fluids, not very different in density, as they strive to separate. Of precisely the same kind (as Bütschli was the first to shew) are the minute alveolar networks which are to be discerned in the cytoplasm of the cell227, and which we now know to be not inherent in the nature of protoplasm, or of living matter in general, but to be due to various causes, natural as well as artificial. The microscopic honeycomb structure of cast metal under various conditions of cooling, even on a grand scale the columnar structure of basaltic rock, is an example of the same surface-tension phenomenon. {172}

Fig. 42.

Fig. 43.

But here we touch the brink of a subject so important that we must not pass it by without a word, and yet so contentious that we must not enter into its details. The question involved is simply whether the great mass of recorded observations and accepted beliefs with regard to the visible structure of protoplasm and of the cell constitute a fair picture of the actual living cell, or be based on appearances which are incident to death itself and to the artificial treatment which the microscopist is accustomed to apply. The great bulk of histological work is done by methods which involve the sudden killing of the cell or organism by strong reagents, the assumption being that death is so rapid that the visible phenomena exhibited during life are retained or “fixed” in our preparations. While this assumption is reasonable and justified as regards the general outward form of small organisms or of individual cells, enough has been done of late years to shew that the case is totally different in the case of the minute internal networks, granules, etc., which represent the alleged structure of protoplasm. For, as Hardy puts it, “It is notorious that the various fixing reagents are coagulants of organic colloids, and that they produce precipitates which have a certain figure or structure, … and that the figure varies, other things being equal, according to the reagent used.” So it comes to pass that some writers228 have altogether denied the existence in the living cell-protoplasm of a network or alveolar “foam”; others229 have cast doubts on the main tenets of recent histology regarding nuclear structure; and Hardy, discussing the structure of certain gland-cells, declares that “there is no evidence that the structure discoverable in the cell-substance of these cells after fixation has any counterpart in the cell when living.” “A large part of it” he goes on to say “is an artefact. The profound difference in the minute structure of a secretory cell of a mucous gland according to the reagent which is used to fix it would, it seems to me, almost suffice to establish this statement in the absence of other evidence.”

Nevertheless, histological study proceeds, especially on the part of the morphologists, with but little change in theory or in method, in spite of these and many other warnings. That certain visible structures, nucleus, vacuoles, “attraction-spheres” or centrosomes, etc., are actually present in the living cell, we know for certain; and to this class belong the great majority of structures (including the nuclear “spindle” itself) with which we are at present concerned. That many other alleged structures are artificial has also been placed beyond a doubt; but where to draw the dividing line we often do not know230. {173}

The following is a brief epitome of the visible changes undergone by a typical cell, leading up to the act of segmentation, and constituting the phenomenon of mitosis or caryokinetic division. In the egg of a sea-urchin, we see with almost diagrammatic completeness what is set forth here231.

Fig. 44.

Fig. 45.

 1. The chromatin, which to begin with was distributed in granules on the otherwise achromatic reticulum (Fig. 42), concentrates to form a skein or spireme, which may be a continuous thread from the first (Figs. 43, 44), or from the first segmented. In any case it divides transversely sooner or later into a number of chromosomes (Fig. 45), which as a rule have the shape of little rods, straight or curved, often bent into a V, but which may also be ovoid, or round, or even annular. Certain deeply staining masses, the nucleoli, which may be present in the resting nucleus, do not take part in the process of chromosome formation; they are either cast out of the nucleus and are dissolved in the cytoplasm, or fade away in situ.

 2. Meanwhile, the deeply staining granule (here extra-nuclear), known as the centrosome, has divided in two. The two resulting granules travel to opposite poles of the nucleus, and {174} there each becomes surrounded by a system of radiating lines, the asters; immediately around the centrosome is a clear space, the centrosphere (Figs. 43–45). Between the two centrosomes with their asters stretches a bundle of achromatic fibres, the spindle.

 3. The surface-film bounding the nucleus has broken down, the definite nuclear boundaries are lost, and the spindle now stretches through the nuclear material, in which lie the chromosomes (Figs. 45, 46). These chromosomes now arrange themselves midway between the poles of the spindle, where they form what is called the equatorial plate (Fig. 47). Fig. 46.Fig. 47.

 4. Each chromosome splits longitudinally into two: usually at this stage—but it is to be noticed that the splitting may have taken place so early as the spireme stage (Fig. 48).

 5. The halves of the split chromosomes now separate from one another, and travel in opposite directions towards the two poles (Fig. 49). As they move, it becomes apparent that the spindle consists of a median bundle of “fibres,” the central spindle, running from pole to pole, and a more superficial sheath of “mantle-fibres,” to which the chromosomes seem to be attached, and by which they seem to be drawn towards the asters.

 6. The daughter chromosomes, arranged now in two groups, become closely crowded in a mass near the centre of each aster {175} (Fig. 50). They fuse together and form once more an alveolar reticulum and may occasionally at this stage form another spireme. Fig. 48.Fig. 49.A boundary or surface wall is now developed round each reconstructed nuclear mass, and the spindle-fibres disappear (Fig. 51). The centrosome remains, as a rule, outside the nucleus. Fig. 50.Fig. 51.

 7. On the central spindle, in the position of the equatorial plate, there has appeared during the migration of the chromosomes, a “cell-plate” of deeply staining thickenings (Figs. 50, 51). This is more conspicuous in plant-cells. {176}

 8. A constriction has meanwhile appeared in the cytoplasm, and the cell divides through the equatorial plane. In plant-cells the line of this division is foreshadowed by the “cell-plate,” which extends from the spindle across the entire cell, and splits into two layers, between which appears the membrane by which the daughter cells are cleft asunder. In animal cells the cell-plate does not attain such dimensions, and no cell-wall is formed.

The whole, or very nearly the whole of these nuclear phenomena may be brought into relation with that polarisation of forces, in the cell as a whole, whose field is made manifest by the “spindle” and “asters” of which we have already spoken: certain particular phenomena, directly attributable to surface-tension and diffusion, taking place in more or less obvious and inevitable dependence upon the polar system*.

* The reference numbers in the following account refer to the paragraphs and figures of the preceding summary of visible nuclear phenomena.

At the same time, in attempting to explain the phenomena, we cannot say too clearly, or too often, that all that we are meanwhile justified in doing is to try to shew that such and such actions lie within the range of known physical actions and phenomena, or that known physical phenomena produce effects similar to them. We want to feel sure that the whole phenomenon is not sui generis, but is somehow or other capable of being referred to dynamical laws, and to the general principles of physical science. But when we speak of some particular force or mode of action, using it as an illustrative hypothesis, we must stop far short of the implication that this or that force is necessarily the very one which is actually at work within the living cell; and certainly we need not attempt the formidable task of trying to reconcile, or to choose between, the various hypotheses which have already been enunciated, or the several assumptions on which they depend.

Any region of space within which action is manifested is a field of force; and a simple example is a bipolar field, in which the action is symmetrical with reference to the line joining two points, or poles, and also with reference to the “equatorial” plane equidistant from both. We have such a “field of force” in {177} the neighbourhood of the centrosome of the ripe cell or ovum, when it is about to divide; and by the time the centrosome has divided, the field is definitely a bipolar one.

The quality of a medium filling the field of force may be uniform, or it may vary from point to point. In particular, it may depend upon the magnitude of the field; and the quality of one medium may differ from that of another. Such variation of quality, within one medium, or from one medium to another, is capable of diagrammatic representation by a variation of the direction or the strength of the field (other conditions being the same) from the state manifested in some uniform medium taken as a standard. The medium is said to be permeable to the force, in greater or less degree than the standard medium, according as the variation of the density of the lines of force from the standard case, under otherwise identical conditions, is in excess or defect. A body placed in the medium will tend to move towards regions of greater or less force according as its permeability is greater or less than that of the surrounding medium232. In the common experiment of placing iron-filings between the two poles of a magnetic field, the filings have a very high permeability; and not only do they themselves become polarised so as to attract one another, but they tend to be attracted from the weaker to the stronger parts of the field, and as we have seen, were it not for friction or some other resistance, they would soon gather together around the nearest pole. But if we repeat the same experiment with such a metal as bismuth, which is very little permeable to the magnetic force, then the conditions are reversed, and the particles, being repelled from the stronger to the weaker parts of the field, tend to take up their position as far from the poles as possible. The particles have become polarised, but in a sense opposite to that of the surrounding, or adjacent, field.

Now, in the field of force whose opposite poles are marked by {178} the centrosomes the nucleus appears to act as a more or less permeable body, as a body more permeable than the surrounding medium, that is to say the “cytoplasm” of the cell. It is accordingly attracted by, and drawn into, the field of force, and tries, as it were, to set itself between the poles and as far as possible from both of them. In other words, the centrosome-foci will be apparently drawn over its surface, until the nucleus as a whole is involved within the field of force, which is visibly marked out by the “spindle” (par. 3, Figs. 44, 45).

If the field of force be electrical, or act in a fashion analogous to an electrical field, the charged nucleus will have its surface-tensions diminished233: with the double result that the inner alveolar meshwork will be broken up (par. 1), and that the spherical boundary of the whole nucleus will disappear (par. 2). The break-up of the alveoli (by thinning and rupture of their partition walls) leads to the formation of a net, and the further break-up of the net may lead to the unravelling of a thread or “spireme” (Figs. 43, 44).

Here there comes into play a fundamental principle which, in so far as we require to understand it, can be explained in simple words. The effect (and we might even say the object) of drawing the more permeable body in between the poles, is to obtain an “easier path” by which the lines of force may travel; but it is obvious that a longer route through the more permeable body may at length be found less advantageous than a shorter route through the less permeable medium. That is to say, the more permeable body will only tend to be drawn in to the field of force until a point is reached where (so to speak) the way round and the way through are equally advantageous. We should accordingly expect that (on our hypothesis) there would be found cases in which the nucleus was wholly, and others in which it was only partially, and in greater or less degree, drawn in to the field between the centrosomes. This is precisely what is found to occur in actual fact. Figs. 44 and 45 represent two so-called “types,” of a phase which follows that represented in Fig. 43. According to the usual descriptions (and in particular to Professor {179} E. B. Wilson’s234), we are told that, in such a case as Fig. 44, the “primary spindle” disappears and the centrosomes diverge to opposite poles of the nucleus; such a condition being found in many plant-cells, and in the cleavage-stages of many eggs. In Fig. 45, on the other hand, the primary spindle persists, and subsequently comes to form the main or “central” spindle; while at the same time we see the fading away of the nuclear membrane, the breaking up of the spireme into separate chromosomes, and an ingrowth into the nuclear area of the “astral rays,”—all as in Fig. 46, which represents the next succeeding phase of Fig. 45. This condition, of Fig. 46, occurs in a variety of cases; it is well seen in the epidermal cells of the salamander, and is also on the whole char­ac­ter­is­tic of the mode of formation of the “polar bodies.” It is clear and obvious that the two “types” correspond to mere differences of degree, and are such as would naturally be brought about by differences in the relative permeabilities of the nuclear mass and of the surrounding cytoplasm, or even by differences in the magnitude of the former body.

But now an important change takes place, or rather an important difference appears; for, whereas the nucleus as a whole tended to be drawn in to the stronger parts of the field, when it comes to break up we find, on the contrary, that its contained spireme-thread or separate chromosomes tend to be repelled to the weaker parts. Whatever this difference may be due to—whether, for instance, to actual differences of permeability, or possibly to differences in “surface-charge,”—the fact is that the chromatin substance now behaves after the fashion of a “diamagnetic” body, and is repelled from the stronger to the weaker parts of the field. In other words, its particles, lying in the inter-polar field, tend to travel towards the equatorial plane thereof (Figs. 47, 48), and further tend to move outwards towards the periphery of that plane, towards what the histologist calls the “mantle-fibres,” or outermost of the lines of force of which the spindle is made up (par. 5, Fig. 47). And if this comparatively non-permeable chromatin substance come to consist of separate portions, more or less elongated in form, these portions, or separate “chromosomes,” will adjust themselves longitudinally, {180} in a peripheral equatorial circle (Figs. 48, 49). This is precisely what actually takes place. Moreover, before the breaking up of the nucleus, long before the chromatin material has broken up into separate chromosomes, and at the very time when it is being fashioned into a “spireme,” this body already lies in a polar field, and must already have a tendency to set itself in the equatorial plane thereof. But the long, continuous spireme thread is unable, so long as the nucleus retains its spherical boundary wall, to adjust itself in a simple equatorial annulus; in striving to do so, it must tend to coil and “kink” itself, and in so doing (if all this be so), it must tend to assume the char­ac­ter­is­tic convolutions of the “spireme.”