Summary
1. This review aims to survey the process of translocation of solutes in the phloem, including the experimental observations of the process, hypothetical mechanisms with their consequences, and the compatibility of these mechanisms with the experimental information.
2. Some properties of the sieve elements are summarized. The characteristic constituent of the sieve elements is a fibrillar protein, P‐protein, of 60–120 A. filaments, whose function and distribution in intact sieve elements are still the subject of debate.
3. Apart from the very high levels of sucrose (0.3–0.9 m) and of specific amino acids and amides (10–100 mm), the contents of the sieve elements are characterized by close regulation of the ionic content; thus K (20–85 mM) and Mg (2.3–23 mM) are very high relative to Na (0.06–0.3 mM) and Ca (0.25–0.5 mM) respectively; the pH is also very high.
4. Convective movement (mass flow) is demanded by the very high rates of mass transfer. The longitudinal sucrose flux is about 2.5 times 106 pmoles cm.‐2 sec.‐1 in petioles, and several times higher in fruits or trees; this is about 105 times any reasonable transmembrane flux, and demands very large loading areas for each file of sieve elements. It also renders unlikely any mechanism demanding an associated trans‐membrane flux of any solute which approaches within several orders of magnitude of the sucrose flow.
5. The evidence from tracer measurements (of 14C or of heat) favour a mass flow of some kind in the sieve tube, with only restricted exchange between the flowing stream and other sucrose pools in the phloem (or out of it). It is not consistent with ready equilibration with a large stationary reservoir of sucrose, or with reverse flows. There is close correspondence between the input and output kinetics of a length of the trans‐location path, or of build‐up curves at different distances; hence lateral exchange from the moving stream is relatively minor.
6. Tracer measurements show that loading into the translocation stream is relatively slow, and is the main determining factor in the time course of appearance of tracer down the stem, or in the profile of radioactivity against distance in the stem. This applies not only to the initial steep front of radioactivity in the stem, but also to the error function profiles found at longer times in some plants; those do not arise as has been suggested, by exchange in a two‐way system of transcellular strands, but are a reflexion of the loading kinetics.
7. The evidence for or against bidirectional movement is equivocal. In conditions in which there is a strong source/sink gradient imposed, the movement of both labelled carbon and heat is consistent with a one‐way system, and is difficult to reconcile with two‐way movement. However, in the absence of any strong gradient there is evidence for bidirectional movement. It is suggested that the pattern of flow, as well as the direction and rate of flow, may be controlled by the source/sink relations along the path.
8. Electro‐osmosis as a mechanism for transl...