We have been able to make these periodic precipitates by the reaction of a great number of chemical substances, giving a bewildering variety of form and structure. Some of these recall the form of various organisms, and especially of insects, as may be seen in Fig. 18.
All the phenomena of life are periodic. The movement of heart and lungs, sleep and waking, all nervous phenomena, have a regular periodicity. It is possible that the study of these purely physical phenomena of periodic precipitation may give us the key to the causation of rhythm and periodicity in living beings.
Besides this periodic precipitation there appear to be other chemical reactions which are periodic. Professor Bredig of Heidelberg has lately described a curious phenomenon, the {76} periodic catalysis of peroxide of hydrogen by mercury. He thus describes his experiment: "We place in a perfectly clean test tube a few cubic centimetres of perfectly pure mercury. Upon this we pour 10 c.c. of a 10 per cent. solution of hydrogen peroxide. The mercury speedily becomes covered with a thin, brilliant bronze-coloured pellicle which reflects light. Then little by little catalysis of the hydrogen peroxide begins, with liberation of oxygen. After some time, from five to twenty minutes, the liberation of gas at the surface of the mercury ceases, the cloud formed by the gas bubbles disappears, and the bronze mirror at the surface of the mercury lights up with the glint of silver. There is a pause of one or more seconds, and then the catalytic action begins afresh, commencing at the edges of the mirror.
The cloud is again formed and again disappears. This beautiful and surprising rhythmic phenomenon may continue at regular intervals for an hour or more."
[Ill.u.s.tration: FIG. 18.--Articulate form produced by periodic precipitation.]
A slight alkalinity of the liquid is necessary to start the phenomenon.
This explains the r.e.t.a.r.dation at the beginning {77} of the experiment, since the rhythmic catalysis cannot begin until the hydrogen peroxide has dissolved a little of the gla.s.s so as to render it slightly alkaline. The catalytic process may, however, be set going at once by adding a trace of pota.s.sium acetate to the solution.
We may even obtain a curve giving an automatic record of the periodicity of this catalytic action. For this purpose the oxygen given off is led to a manometer, which registers on a revolving drum the periodic variation in pressure. The curve thus obtained presents a remarkable resemblance to a tracing of the pulse. The frequency and character of the undulatory curve is modified by physical and chemical influences. Like circulation or respiration, periodic catalysis has its poisons, and exhibits signs of fatigue, and of paralysis by cold.
The rhythmic catalysis of Bredig produces an electrical current of action between the mercury and the water just like that produced by the rhythmic contraction of the heart, and this current may be registered in a similar way by means of the Einthoven galvanometer. Thus the heart-beat may be but an instance of rhythmic catalysis, since both produce the same phenomena, movement, chemical action, and periodic currents. In the chapter on physiogenesis we shall return to the study of this question and consider another rhythmic phenomenon which is the result of osmotic growth.
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CHAPTER VII
COHESION AND CRYSTALLIZATION
Chemical affinity is the force which holds together the different atoms in a molecule. Cohesion is the force which holds together molecules which are chemically similar. Although physical science distinguishes three states of matter, solid, liquid, and gaseous, yet here as elsewhere there are no sharp dividing lines, but rather an absolute continuity. We have in fact many intermediate states; between liquids and gases there are the various conditions of vapour, and between liquids and solids we get viscous, gelatinous, and paste-like conditions. The only real difference between solids, liquids, and gases is the intensity of the force of cohesion, which is considerable in solids, feeble in liquids, and absent in gases.
A living organism is the arena in which are brought into play the opposing forces of cohesion and disintegration. The study of cohesion is therefore a vital one for the biologist, and especially cohesion under the conditions which obtain in living beings, viz. in liquids of heterogeneous const.i.tution. The forces of cohesion brought into play under these conditions may be beautifully ill.u.s.trated by a simple experiment. We take a plate of gla.s.s, well cleaned and absolutely horizontal. On it we pour a layer of salt water, and in the middle we carefully drop a spot of Indian ink. The drop at once begins to diffuse, and we obtain a circular figure, like the monopolar field of diffusion already described, the rays of diffusion radiating from the centre in all directions.
[Ill.u.s.tration: FIG. 19.--Muriform cohesion figure formed by a drop of Indian ink in a solution of salt.]
If we keep the plate carefully protected from all disturbing influences, after some ten to twenty minutes we shall see the coloured particles returning on their path, and the centre of {79} the drop becoming more and more black. Each line of force becomes segmented into granules, which gradually increase in size, and approach nearer to one another and to the centre of the drop, until it a.s.sumes the mulberry appearance shown in the photograph (Fig. 19).
[Ill.u.s.tration: FIG. 20.--Seven similar drops of Indian ink diffusing in a salt solution. Two minutes after introducing the drops.]
If we sow a number of drops of Indian ink in regular order on the surface of a salt solution, we obtain most beautiful patterns formed by the mutual repulsion of the drops. Figs. 20, 21, and 22 represent the successive aspects of seven drops of Indian ink thus sown on a layer of salt solution, and kept undisturbed long enough to allow of their evolution. Fig. 20 shows the aspect after two minutes, when the diffusion is almost complete. In Fig. 21, photographed after fifteen {80} minutes, the colouring matter has almost entirely reunited to form separate granulations; whilst in Fig. 22, taken after thirty minutes, these granulations are rearranged to form an agglomeration around the centre of each drop.
[Ill.u.s.tration: FIG. 21.--The same drops 15 minutes later, showing the granulation appearance.]
The following experiment, which is more difficult, will show the cohesive attraction of one drop for another. A plate of gla.s.s is adjusted absolutely horizontal, and covered as before with a layer of salt solution. On this we sow a number of drops of the same salt solution coloured with Indian ink.
The drops must be of exactly the same concentration as the salt medium, so as to avoid any difference of osmotic pressure between the drops and the medium, otherwise the drops would not remain intact but would diffuse into the solution. Since under these conditions the liquid of the medium around the drops is perfectly symmetrical and h.o.m.ogeneous, it cannot exercise any influence on the liquid of the drops.
[Ill.u.s.tration: FIG. 22.--The same drops after 30 minutes. The granulations have agglomerated at the centre of the drops.]
It is otherwise, however, with the colouring matter of the {81} drops. The particles of Indian ink may be seen pa.s.sing from one drop to another, the coloured circles become elongated towards one another, touch, and finally unite. If, as in Fig. 23, the drops are of different size, the larger one will have a preponderating attractive action and eat up the smaller drops.
In the figure, six small drops are placed around a large one, and the smaller drops have begun to be deformed and to move towards the larger drop. This central drop is also deformed, and has a.s.sumed a more or less hexagonal form, under the influence of the attraction of the six smaller ones. It may be noticed that the least prominent angle of the hexagon is opposite the small drop which is farthest away from it, whilst one of the smaller drops has already begun to be swallowed up by the large one. This cohesion phenomenon is very slow in its action, but after an hour or two the central drop will be found to have {82} completely absorbed the six smaller ones, and only one large drop will remain.
[Ill.u.s.tration: FIG. 23.--Attraction between coloured drops in an isotonic solution.]
_Incubation._--In the living organism we frequently find conditions similar to those realized in this experiment, viz. very slow movements of diffusion in liquids containing particles in suspension. In such cases the consequences must be the same, viz. granulation and segmentation. Consider for a moment the incubation of an egg. The heat of incubation determines a certain amount of evaporation through the sh.e.l.l, with a concentration of the liquid near the surface. As a consequence of this superficial concentration we get segmentation of the vitellus, with the production of a morula.
_Artificial Parthenogenesis._--The experimental parthenogenesis of Loeb and Delage consists in plunging the egg into a liquid other than sea water, and returning it again to its original medium. This operation will necessarily determine slow movements of diffusion in the egg, which will give rise to segmentation. It may be objected that segmentation is also produced by a solution which is isotonic with sea water. Such a solution would not indeed produce an exchange of water with the egg, but it would set up an exchange of electrolytes, since there would be a difference of their osmotic pressure in the egg and in the new isotonic medium. The extremely slow movements of diffusion thus produced would be very favourable to the action of the cohesive force on the particles in suspension, and hence to the segmentation of the egg.
[Ill.u.s.tration: FIG. 24.--A circle of eight drops of Indian ink 30 minutes after they have been sown in a salt solution. The drops have undergone diffusion and subsequent cohesion, resulting in a reticulate structure.]
Few physical phenomena give us a deeper insight into the phenomena of life than those which we here contemplate. There is still another experiment which is even more convincing. On the surface of our horizontal salt solution we sow a number of drops of a more concentrated salt solution at equal distances around the circ.u.mference of a circle. Movements of diffusion are thus set up in the interior of the circle, and after a time, when this diffusion has become so slow as to be almost imperceptible, a furrow begins to appear in the coloured ma.s.s. Then a second and third appear, and others crossing the former break up the ma.s.s {83} into segments. Finally the segmentation becomes complete, and the preparation presents a muriform appearance, looking in fact something like a mulberry (Fig. 24). If the preparation is preserved for several hours longer, we may see the cells formed by segmentation unite around the circ.u.mference so as to form a hollow bag corresponding to a gastrula, as shown in Fig. 25.
[Ill.u.s.tration: FIG. 25.--The same preparation several hours later, showing a cellular gastrula-like structure.]
These preparations are extremely sensitive to external influences, which renders the demonstration of cohesion phenomena difficult. I have nevertheless on several occasions been able to project the experiment on the screen during a lecture. The segmentation is influenced by very slight currents of diffusion, and I have many preparations showing the segmentation regularly distributed in various ways along radial diffusion lines. We may in this way produce many varieties of structure lamellar, vacuolate, or cellular, in fact {84} all the tissue structures which are met with in living organisms. All these structures are retractile, the retraction going on very slowly for a long time, as if the force of cohesion continued to act in the web of the structure even after its formation was complete. The phenomenon is a purely physical synthetic reproduction of the phenomenon of coagulation, the cohesion figure being in fact a retractile clot.
[Ill.u.s.tration: FIG. 26.--Field of crystallization of sodium chloride (magnified 60 diameters).]
_Crystallization._--When we evaporate a solution of a crystalloid it becomes more concentrated, slow movements of diffusion are set up, and at a given moment agglomeration occurs, the agglomerates taking the form of crystals. Thus crystallization may be regarded as a particular case of conglomeration by cohesion, differing only in the regularity of the arrangement of the molecules, which gives the geometrical form of the crystal. Hence we can easily understand how the presence of a crystalline fragment may facilitate the process of crystallization. Consider a liquid in which extremely slow movements of diffusion are taking place. If the liquid is perfectly h.o.m.ogeneous there will be no centre of attraction to which the molecules may become attached. {85}
[Ill.u.s.tration: FIG. 27.--Field of crystallization around a crystal of sodium chloride in process of formation.]
If, however, a crystal or other heterogeneous structure is present, it forms a centre of cohesion which will attach any molecules that are brought by diffusion into its sphere of attraction. We have succeeded in photographing the arrangement of the molecules of a liquid around a crystal in the act of formation (Fig. 26). For this purpose we add to the solution traces of some colloidal substance, such as gelatine or gum, so as to delay the crystallization. It may thus be shown that the molecules of the surrounding liquid are already arranged in crystalline order for some distance from the crystal, forming a sort of field of crystallization. The arrangement of this regular field varies in different cases, and is more or less complicated according to circ.u.mstances. One of the most frequent forms is that shown in Fig. 27, which is the field around a crystal of sodium chloride. In the centre {86} of the crystal is a square with well-marked outline. At each corner of this square there is a straight line at right angles to the diagonal, which will form the sides of the crystal in process of formation. From the middle of each side arise yet other perpendiculars, which in their turn bear other cross lines, each new line being set at right angles to its predecessor. A later stage of crystallization is shown in Fig. 27, where the two squares one inside the other at an angle of 45 are clearly indicated.
[Ill.u.s.tration: FIG. 28.--Three crystals of sodium chloride in process of formation, each in the centre of a field of crystallization.]
Every crystallizable substance gives a different characteristic field of crystallization. In 1903, at the Congress of Angers, I terminated my address by these words: "The field of crystallization may serve to determine the character of a substance in solution." I have subsequently received from Carbonell y Soles of Barcelona an interesting work on this subject, which he contributed to the International Congress of Medicine at Madrid in 1903, ent.i.tled _Applicacion de la crystalogenia experimental a la investigacion toxicologica de cas alcalodes_. {87}
Six years ago I received from Australia an exceedingly beautiful photograph of a thin pellicle found in a rain gauge. My correspondent supposed that this strange figure might have been produced under the influence of an electric or magnetic field. I was able to a.s.sure him by return of post that the figure was the result of the crystallization of copper sulphate in a colloidal medium. In return I received a letter verifying this fact, and saying that there were copper works in the neighbourhood, and the air was filled with the dust of copper sulphate.
Living beings are but solutions of colloids and crystalloids, and their tissues are built up by the aggregation of these solutes. We have already seen how the forces of crystallization are modified in colloid solutions.
This force of crystallization must play an important role in the metamorphoses of the living organism, and influence their morphology. It may therefore be of interest to investigate some of the numberless forms of crystallization in colloidal solutions.
[Ill.u.s.tration: FIG. 29.--Crystallization of sodium chloride in a colloidal solution, giving a plant-like form.]
[Ill.u.s.tration: FIG. 30.--Form produced by the crystallization of chloride of ammonium in a colloidal solution.]
Figs. 29 and 30 represent the forms produced by chloride of sodium and chloride of ammonium respectively, in solutions of gelatine of different degrees of concentration. Their resemblance to vegetable growth is so remarkable that several observers on first seeing them have called them "Fern-crystals."
I should like here to recall to your notice the work of an English observer, Dr. E. Montgomery of St. Thomas"s {88} Hospital, which was published as long ago as 1865. This work was recently brought to my notice by the kindness of Professor Baumler of Freiburg. He says: "Crystals are not strangers in the organic world. Many organic compounds are able to a.s.sume crystalline forms under certain conditions. Rainey has shown that many sh.e.l.ls consist of globular crystals _i.e._ of mineral substances made to crystallize by the influence of viscid material." In this connection I may also mention the interesting work of Otto Lehmann of Karlsruhe on liquid crystals.
In conclusion, we may recall the words of Schwann himself, the originator of the cell theory: "The formation of the elementary shapes of an organism is but a crystallization of substances capable of imbibition. The organism is but an aggregate of such imbibing crystals."
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CHAPTER VIII
KARYOKINESIS
In 1873, Hermann Fol, writing of the eggs of Geryonia, thus describes the phenomenon of karyokinesis: "On either side of the residue of the nucleus there appears a concentration of plasma, thus forming two perfectly regular star-like figures, whose rays are straight lines of granulations. There are other curved rays which pa.s.s from one star or centre of attraction to the other. The whole figure is extraordinarily distinct, recalling in a striking manner the arrangement of iron filings surrounding the poles of a magnet. Sachs" theory is that the division of the nucleus is caused by centres of attraction, and I agree with him, not on theoretical grounds, but because I have actually seen these centres of attraction."
Since the discovery of Hermann Fol, a great number of explanations have been given, all of them theoretical, to account for the figures and phenomena of karyokinesis. Many of these so-called explanations are mechanical, while others invoke the aid of magnetism or electricity to account for the resemblance of the figures of karyokinesis to the magnetic or electric phantom or spectre. Among the authors who have dealt with this question we may mention Hartog of Cork, Gallardo of Buenos Ayres, and Rhumbler of Gottingen.