An Anterior-Posterior Gradient of Refractive Index in the Ameba and Its Significance in Ameboid Movement

Sustained locomotion in Amoeba proteus and Amoeba dubia results in the establishment of a measureable gradient of refractive index along the anterior-posterior axis of the cell, provided thickness of the specimens is kept constant by even compression under a selected coverglass supported by quartz beams of uniform diameter. The tail region of the ameba develops a higher refractive index, indicative of from 6 to 40 per cent more organic matter (expressed as protein) there than present in the front. This gradient fades on cessation of movement. The average protein concentrations in the crystal-free tails and fronts of 15 A. proteus were 3.9 and 3.4 per cent, respectively. In individual experiments, the tail-front difference ranged from one to eight times the accuracy of the method. Since the gradient of refractive index was shown not to result from extraction of water from the tail by the contractile vacuole, it was interpreted as displacement of water toward the anterior part of the cell during movement. It is suggested that contraction of the ectoplasm drives forward a "tide" of syncretic fluid, the anterior border of which is visible as the hyaline cap, which contains less than 1 per cent protein. The movement of the granular endoplasm into the hyaline cap would then complete the cycle by imbibition of the fluid tide. The theoretical positions of Pantin and of Dellinger have been combined in the proposal that ameba cytoplasm consists of a network of a contractile phase which is able to expel (by syneresis) a highly mobile fluid phase. Some other possible interpretations are discussed.