From the behaviour of wort cultures of 12 yeasts and of washed cells from these cultures, it is concluded that: (a) yeasts differ in their abilities to flocculate in the presence of maltose or mannose, and in the absence of calcium ions; (b) during growth young cells actually lose their power to flocculate and are not merely prevented from doing so by substances in wort; (c) the tendency to form floes containing branching chains of cells is a phenomenon distinct from true flocculation.
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Allison A. Eddy (1955) studied this question.