When the surface tension of a drop of fluid is lowered by bringing into contact with it some other substance that possesses this power, the surface rushes away at great speed in all directions from the point where the tension is lowered, because usually the tension is reduced very considerably. In this surface movement it is found that new surface is made where the tension is lowered and old surface is destroyed, that is, pulled into the interior over a large part of the surface opposite to where the tension is lowered. The speed of the surface movement is most rapid near the point where the tension is lowered and becomes gradually slower as the opposite side of the drop is approached, where there is no movement. This variation in speed of the moving surface seems to be due largely to the small area in which the tension is lowered as compared with the whole surface of the drop.
In the ameba the conditions are reversed. The surface layer moves toward a point with increasing speed, instead of away from a point. In both the ameba and the drop the greatest speed is attained near the small area where the change in surface tension occurs.
The behavior of large and heavy particles on the surface of a drop of fluid and on an ameba are similar. A heavy particle as of sand, or a small glass rod, laid on the ameba, is not moved by the surface layer. It forms an island of surface matter around which the moving surface layer flows. Precisely the same thing happens in surface layer movements in inanimate fluids.
Again in point of thinness there is no disagreement so far as microscopic observation goes. Neither the surface film on an ameba nor the surface film on a fluid can be directly observed microscopically to be different from the fluid below it. The surface layer is, as is generally believed, of molecular dimensions, and its thickness is beyond the limits of vision. Unless some special means is discovered therefore for making visible the surface film, such as a process of staining, it may be impossible to ascertain its ultimate structure directly, for it overlies a mass of heterogeneous fluid whose composition is constantly changing.
It seems to follow from what is observed of the surface tension layers of the fluids of physics that such layers must be of the same constitution as the body of the fluid over which the layer is formed, although, as is well known, the proportion of the ingredients in the surface layer is different from that in the body of the fluid. Now since the resemblance between the surface layer of an ameba and a surface layer on a drop of fluid has thus far been found to be complete, it is pertinent at this point to discuss Gruber’s (’12) suggestion that the movement of particles forward on an ameba is due to the forward movement of an inert layer of mucus or gelatinous material secreted by the ameba.
To begin with, observation does not support Gruber’s suggestion. No such layer can be seen. Such a layer, since it is shown to persist for several minutes at least, should remain after an ameba bursts, under experimental conditions, but no such remains can be seen. Its existence should be demonstrable by the use of dyes, but the evidence is negative. Indeed there is not any direct evidence that can be brought in support of the suggestion that this surface layer is gelatinous in composition. Moreover, as we have seen, the layer on the ameba that carries particles forward seems to be destroyed at the anterior end, for in what other way would particles remain at the anterior end after being brought there? But the supposition that a gelatinous layer might be drawn into the interior at the anterior end is also negatived by observation, for no very small particles clinging to or imbedded in the surface substance are ever drawn into the ameba, as would almost certainly be the case if the substance composing the layer were gelatinous. And as to supposing that this layer, if gelatinous, might behave essentially as a surface tension layer and therefore be drawn in at the anterior end of the ameba, this is contrary to the experience of physics; for the physical nature of the ameba would make it impossible for the ameba to have a surface layer of gelatinous matter. There do not seem to exist any grounds therefore for supposing that the outermost layer of an ameba, the layer that carries particles as described in the preceding pages, can consist of an inert substance as Gruber suggests.[4]
From these considerations, then it appears that all the evidence available, both direct and indirect, points to the conclusion that the behavior of the surface layer on the ameba resembles in general and in detail the behavior of a surface tension layer in an inert drop of fluid, and that we must regard the surface layer on the ameba as a true surface tension layer. This layer is therefore a dynamic layer, containing free energy, and capable of performing work. It is physiologically distinct from ectoplasm, as ectoplasm is distinct physiologically from endoplasm. But the distinctive properties which the surface layer possesses are functions of its position. These properties clearly indicate that its constitution is protoplasmic, corresponding to the fluid parts of the internal protoplasm.
The surface layer of the ameba is probably identical with what is commonly called the plasma membrane or semi-permeable membrane as postulated by Overton (’07). The peculiar structure supposed to be possessed by plasma membranes are held to be due chiefly to surface forces. The fact that the surface layer of the ameba is continually being destroyed and re-created during locomotion does not support the view that the plasma membrane is of inert composition, as for example, lipoidal, as has been suggested. The observations, on the contrary, confirm Höber’s (’11) view that the plasma membranes generally are living structures. But it may be regarded as certain that if lipoids are present in the protoplasm of the ameba, these substances, according to the principle of Willard Gibbs, will be found in higher concentration in the surface film than in the body of the ameba.
Perhaps the most important question that arises in connection with the surface layer of the ameba is: What causes it to move in the manner described? But we can do little more than ask the question. It has been seen that the surface film moves toward an area of increased tension rather than from an area where the tension has been lowered. However, since we are completely in the dark respecting the composition of the surface layer or of the fluid parts of the ameba, it is exceedingly hazardous to venture an explanation. If the surface layer should have its tension lowered by a concentration of lipoids in it, we would be faced by the necessity of explaining their removal at the anterior end. If we turn to electrical causes we meet again with great difficulties. An ameba moves with the electric current, when a current is passed through the water. The surface layer under these conditions behaves normally, as may be inferred from Jennings’ (’04) figure on page 198. That is, the current controls the direction of the movement of the ameba, with the current leaving the ameba at the point of highest surface tension. This is contrary to the action of the mercuric capillary electrometer, in which the mercury column also moves with the current, but because of lowered surface tension where the current leaves the mercury. The conditions surrounding these cases are so different however, that very little can be gained by setting them in contrast to each other.