Diffusion and bulk flow in phloem loading - a theoretical analysis of the polymer trap mechanism in plants

Plants create sugar in the mesophyll cells of their leaves by photosynthesis. This sugar, mostly sucrose, has to be loaded via the bundle sheath into the phloem vascular system (the sieve elements), where it is distributed to growing parts of the plant. We analyze the feasibility of a particular loading mechanism, active symplasmic loading, also called the polymer trap mechanism, where sucrose is transformed into heavier sugars, such as raffinose and stachyose, in the intermediary-type companion cells bordering the sieve elements in the minor veins of the phloem. Keeping the heavier sugars from diffusing back requires that the plasmodesmata connecting the bundle sheath with the intermediary cell act as extremely precise filters, which are able to distinguish between molecules that differ by less than 20% in size. In our modeling, we take into account the coupled water and sugar movement across the relevant interfaces, without explicitly considering the chemical reactions transforming the sucrose into the heavier sugars. Based on the available data for plasmodesmata geometry, sugar concentrations and flux rates, we conclude that this mechanism can in principle function. We find that the water flow through the plasmodesmata, which has not been quantified before, contributes only 10-20% to the sucrose flux into the intermediary cells, while the main part is transported by diffusion. On the other hand, the subsequent sugar translocation into the sieve elements would very likely be carried predominantly by bulk water flow through the plasmodesmata. Thus, in contrast to apoplasmic loaders, all the necessary water for phloem translocation would be supplied in this way with no need for additional water uptake across the plasma membranes of the phloem.Comment: 29 pages with 5 figure

Energy harvesting through gas dynamics in the free molecular flow regime between structured surfaces at different temperatures

For a gas confined between surfaces held at different temperatures the velocity distribution shows a significant deviation from the Maxwell distribution when the mean free path of the molecules is comparable to or larger than the channel dimensions. If one of the surfaces is suitably structured, this non-equilibrium distribution can be exploited for momentum transfer in tangential direction between the two surfaces. This opens up the possibility to extract work from the system which operates as a heat engine. Since both surfaces are held at constant temperatures, the mode of momentum transfer is different from thermal creep flow that has gained more attention so far. This situation is studied in the limit of free-molecular flow for the case that an unstructured surface is allowed to move tangentially with respect to a structured surface. Parameter studies are conducted, and configurations with maximum thermodynamic efficiency are identified. Overall, it is shown that significant efficiencies can be obtained by tangential momentum transfer between structured surfaces

Swimming and feeding of mixotrophic biflagellates

Many unicellular flagellates are mixotrophic and access resources through both photosynthesis and prey capture. Their fitness depends on those processes as well as on swimming and predator avoidance. How does the flagellar arrangement and beat pattern of the flagellate affect swimming speed, predation risk due to flow-sensing predators, and prey capture? Here, we describe measured flows around two species of mixotrophic, biflagellated haptophytes with qualitatively different flagellar arrangements and beat patterns. We model the near cell flows using two symmetrically arranged point forces with variable position next to a no-slip sphere. Utilizing the observations and the model we find that puller force arrangements favour feeding, whereas equatorial force arrangements favour fast and quiet swimming. We determine the capture rates of both passive and motile prey, and we show that the flow facilitates transport of captured prey along the haptonema structure. We argue that prey capture alone cannot fulfil the energy needs of the observed species, and that the mixotrophic life strategy is essential for survival