Ameboid movement

by Asa A. Schaeffer

From numerous observations directed toward this point, it appears that the conditions under which streaming is resumed after a pause, whether in the same or in the reverse direction, are of great variety. The shape, size, slenderness, and the position on the ameba of the pseudopod, as well as the strength and character of the stimulus, are among the factors capable of changing in whole or in part the flow of endoplasm in a pseudopod. In an ameba that has been moving along a homogeneous flat surface, as nearly unstimulated as may be, the endoplasm first begins to flow out of the lower half of the retracting pseudopod, if the pseudopod is more or less uniformly conical in shape and rather slender. In such a case it may be said that the retracting pseudopod was withdrawn “by the ameba,” and that it did not itself receive an external stimulus producing retraction. If, however, the tip of a pseudopod as described receives a strong negative stimulus, the endoplasm frequently flows back from the tip while it is still flowing into the pseudopod at the base. But very soon thereafter, in such an event, the streaming becomes unified and the pseudopod is withdrawn. In broad pseudopods about to be withdrawn, the endoplasm may begin to move anywhere along its length. This is undoubtedly due to the continuous local changes in the walls of the pseudopod, which are characteristic of this species of ameba (see p. 20).

In an ameba which has been brought to a standstill, as by a sudden flash of light, the first sign of recurring streaming is in the anterior half, whether the original direction of streaming is resumed or reversed. If the direction is reversed, the active pseudopods retract for a considerable distance before a new one is projected. The endoplasmic stream in a slender withdrawing pseudopod may not reach to the tip for from several seconds to a minute, if the tip is slightly positively stimulated. One may then observe ectoplasm streaming toward the tip and toward the base, in the respective regions, at the same time, with considerable fluctuation back and forth of the neutral zone separating the two streams. The fate of such a pseudopod depends on its size, on its position on the ameba, and the strength of the stimulus affecting it and the rest of the ameba. That is, if the pseudopod is small or on the posterior half of the ameba, or only slightly stimulated, it will be retracted; but if it is large, or on the anterior end of the ameba, or more strongly stimulated than the rest of the ameba, it may again become active.

The fact that protoplasm is practically incompressible makes it clear that if streaming can be observed to begin after a pause at some point after it begins at others, the ectoplasmic walls of the ameba must give way in the region where streaming begins. Since it has been established by observation that the ectoplasm may give way at any point, it follows that one of the principal factors affecting streaming is the elasticity and liquefiability of the ectoplasm.

The streaming in an ameba is coordinated. The direction in which the endoplasm flows in the several pseudopods, when there are no stimuli received externally that produce visible changes in behavior, gives one the impression that there is a “centre” controlling movement. The several pseudopods do not act at all capriciously. The ameba seems to move the pseudopods, not the pseudopods the ameba. If this impression of coordination is correct, it is of the first importance in a study of ameboid movement. Further on, this point will be taken up at length in connection with the character of the path an externally unstimulated ameba describes (p. 109); but there are certain observations which aid in the analysis of the problem of coördination from the point of view of the pseudopod, instead of that of the ameba as a whole, and to these observations we may now direct our attention.

The mass of endoplasm within a pseudopod moves practically always in one direction. In any cross-section of a pseudopod that is more or less cylindrical in shape, the endoplasm in the center moves most rapidly, that near to it less rapidly, while that near the ectoplasm moves very slowly. One never observes a forward stream on one side of the pseudopod and a backward stream on the other. Nor does one observe parallel streams of endoplasm flowing in opposite directions within the same

Figure 1. Illustrating the various directions of endoplasmic streaming in growing and retracting pseudopods. a, two oppositely directed streams in a pseudopod, one directed toward the base and the other toward the tip of the pseudopod, with a neutral zone between. b, two streams flowing toward each other. Cases c to r are self explanatory. s, rotational currents observed occasionally in various species of amebas. t, “fountain currents,” sometimes observed in Amoeba blattae, and rarely in other forms. u and v represent cases of streaming which have not been observed and which probably do not occur. w, similar to v, but with a wide neutral zone between the streams, represents an actual observed case. m and r probably occur only very rarely; no such cases have been seen, but there seems to be no reason why they do not sometimes occur. Excepting m, u, r and v, all these figures were drawn from observed cases of streaming.

ectoplasmic tube, in an ameba of several pseudopods, excepting where there is a wide zone of stationary endoplasm between the streams (Figure 1, v, w). But in “fountain currents,” such as Rhumbler (’98, p. 190) described and figured for Amoeba blattae Bütschli, and which may readily be observed in most species of amebas if immersed in a solution of gelatin thick enough to keep the amebas from sinking, there is a central stream of endoplasm flowing forward, and a peripheral stream of ectoplasm flowing backward, with a thin neutral zone between ([Figure 29], d). As we shall see further on, however, these fountain currents are in principle the same as the currents observed in ordinary locomotion, the apparent difference being due to the fact that there is no locomotion. It is true, then, that within the same pseudopod at any cross section the endoplasm always streams in one direction, and the streaming is unified.

When new pseudopods are formed, or when old ones are retracted, and especially when both these phenomena occur at the same time and close together on a part of an older pseudopod, some of the details of coordination in streaming are readily made out. In [Figure 1] are shown a number of observed cases of pseudopod formation and retraction, with the direction of endoplasmic streams indicated at a given instant. For the purpose of illustration, several (presumably) possible but unobserved cases, m and r, are sketched, and also two cases, u and v, which have not been observed and which probably do not occur. The general conclusion to be drawn from these observations is that, while the endoplasm in the body of an ameba as a whole may be streaming in several different directions at any given instant, that is almost never the case with an individual pseudopod, especially if the pseudopod is of small or medium size and not too flat or otherwise irregular in shape. The pseudopod is therefore the unit of coordinated protoplasmic streaming.