The first attempt that was made to explain ameboid movement in conformity with the demands of modern experimental science, that is, on the basis of physical factors, was made by Berthold (’86). By means of simple experiments with inert fluids (oils, alcohol, water, ether) which were modeled after an experiment described by the physicist Paalzow (’58), Berthold concluded that locomotion in ameboid organisms is due to the physical attraction of the anterior end to the substratum. The ameba was supposed to behave like a drop of fluid which moved towards the point where the tension of the ameba’s surface was decreased by contact with the substratum. The ameba did not push out pseudopods according to Berthold, but they were pulled out because of a difference in surface tension between them and the substratum. But pseudopods which were extended into the water and out of contact with a solid substratum, were said to be extended by a contractile effort of the posterior region of the ameba.
Bütschli (’92, p. 187) pointed out that it was highly improbable that pseudopods in contact with a solid substratum were projected in a fundamentally different way from that in which free pseudopods were extended, as explained by Berthold. Bütschli assumed that all ameboid movement was due to the same fundamental cause. He postulated surface tension as the active agent, as Berthold had done for the extension of pseudopods in contact with a solid substrate; but Bütschli assumed that the decrease in surface tension at the anterior end of the ameba was brought about by the bursting of protoplasmic droplets of a more fluid consistency on the surface of the ameba, the consistency of which was less fluid, thus bringing about a decrease of surface tension and consequent forward streaming of the endoplasm. The necessary migration of the more fluid droplets to the surface was determined by internal conditions. The direction in which an ameba moves was assumed to depend therefore not upon the physical character of the substrate, as suggested by Berthold, but upon such internal changes as control the movement of the more liquid part of the internal protoplasm to the outer surface.
Rhumbler (’98) wrote extensively on the subject of ameboid movement, especially from the point of view of the feeding habits of amebas. He concluded that the flow of protoplasm, while engulfing a food object, was a direct result of the lowering of the surface tension of the protoplasm by contact with the food object (p. 207), thus causing its envelopment. Numerous other writers of the time, including Quincke (’88), Verworn (’89, ’92), Blochmann (’94), Bernstein (’00) and Jensen (’02), agreed in a general way with Rhumbler’s position that surface tension changes are the cause of locomotion in ameba.
In 1904 the general subject of ameban behavior was extensively studied by Jennings, and from his observations he concluded that surface tension cannot account for many of the reactions observed. Other factors, he held, must be at work, such as contractility, which, acting in the posterior region, causes the endoplasm to flow forward. But Jennings found it impossible to explain on the same basis the extension of free pseudopods, and the creeping of a pseudopod, or of the whole ameba, over a solid substratum.
From further observations Rhumbler (’05, ’10) came to modify his earlier views as stated above. The rapid advances in the study of the chemistry of colloids doubtless suggested to Rhumbler that the change from endoplasm to ectoplasm resembled the change from a sol to a gel state, and that in this process of gelation lay the source of energy manifested in ameboid movement. In thus calling attention to, and emphasizing the colloidal nature of, the conversion of endoplasm into ectoplasm and vice versa, the problem of ameboid movement came to be discussed from an entirely new angle. Certain phases of Rhumbler’s theory are developed and elaborated by Hyman (’17) who agrees in general with Rhumbler’s conclusions.
In a series of papers on feeding and other reactions of ameba, Schaeffer (’12, ’16, ’17) concluded that Rhumbler’s general statement, wherein he says that changes in behavior are directly deducible from the action of stimuli in effecting liquefaction or gelation of the ectoplasm, does not hold in many cases of feeding, and that the mechanism controlling locomotion and feeding is not external, as maintained by Rhumbler, but internal.
CHAPTER III
The General Features of Endoplasmic Streaming
The streaming of the endoplasm is the most conspicuous feature of ameboid movement. It is even more noticeable than the movement of the pseudopods themselves, because of its greater speed and because it occurs in all parts of the ameba. Its importance in movement is essential, for no continued locomotion can be observed unless accompanied by streaming. It may be profitable therefore to enquire into the general features of streaming, and to observe some of the necessary consequences streaming imposes upon such an animal as the ameba.
Let us take as an example an Amoeba proteus (Pallas, ’66, emend. Leidy, ’79, emend. Schaeffer, ’16) in characteristic movement (see [Figure 11], p. 37). The main streams of endoplasm are in the same direction as that in which the ameba moves. In the withdrawing pseudopods the current is, of course, toward the main mass of the ameba. The endoplasmic stream is continuous from the posterior end to the tips of the advancing pseudopods. The retracting pseudopods flow into the main stream as tributaries. If, as often happens, the ameba is without pseudopods, there is then a single stream arising in the posterior end and flowing to the anterior end. In such a case it is readily observed how absolutely dependent locomotion is upon endoplasmic streaming.
It often happens, such as when the ameba is receiving a strong stimulus, that streaming is arrested and brought to a stop for a few seconds, more or less. Presently however the endoplasm begins to flow as before. At what point, in such a case, is the first movement of endoplasm detectible? Is it at the free end of the pseudopod, at its middle region, at its base, or at the posterior end of the ameba? Bütschli (’80, p. 116) observed that in a withdrawing pseudopod the streaming begins at the free end of the pseudopod; but his (’92, p. 201) later explanation of ameboid movement seems to require that the endoplasm must begin to move at the base of the withdrawing pseudopod. Jennings (’04, p. 157) observed that in a withdrawing pseudopod the current of endoplasm begins at the base of the pseudopod.