(J.M., P.G.) Sucrose (Suc) transporters belong to a large gene family. The physiological role of SUT1 proteins has been intensively investigated in higher plants, whereas that of SUT4 proteins is so far unknown. All three known Suc transporters from potato (Solanum tuberosum), SUT1, SUT2, and SUT4, are colocalized and their RNA levels not only follow a diurnal rhythm, but also oscillate in constant light. Here, we examined the physiological effects of transgenic potato plants on RNA interference (RNAi)-inactivated StSUT4 expression. The phenotype of StSUT4-RNAi plants includes early flowering, higher tuber production, and reduced sensitivity toward light enriched in far-red wavelength (i.e. in canopy shade). Inhibition of StSUT4 led to tuber production of the strict photoperiodic potato subsp. andigena even under noninductive long-day conditions. Accumulation of soluble sugars and Suc efflux from leaves of transgenic plants are modified in StSUT4-RNAi plants, leading to modified Suc levels in sink organs. StSUT4 expression of wild-type plants is induced by gibberellins and ethephon, and external supply of gibberellic acid leads to even more pronounced differences between wild-type and StSUT4-RNAi plants regarding tuber yield and internode elongation, indicating a reciprocal regulation of StSUT4 and gibberellins.

Green plants are Earth's primary solar energy collectors. They harvest the energy of the Sun by converting light energy into chemical energy stored in the bonds of sugar molecules. A multitude of carefully orchestrated transport processes are needed to move water and minerals from the soil to sites of photosynthesis and to distribute energy-rich sugars throughout the plant body to support metabolism and growth. The long-distance transport happens in the plants' vascular system, where water and solutes are moved along the entire length of the plant. In this review, the current understanding of the mechanism and the quantitative description of these flows are discussed, connecting theory and experiments as far as possible. The article begins with an overview of low-Reynolds-number transport processes, followed by an introduction to the anatomy and physiology of vascular transport in the phloem and xylem. Next, sugar transport in the phloem is explored with attention given to experimental results as well as the fluid mechanics of osmotically driven flows. Then water transport in the xylem is discussed with a focus on embolism dynamics, conduit optimization, and couplings between water and sugar transport. Finally, remarks are given on some of the open questions of this research field.

The plant sucrose transporter SUT1 from Solanum tuberosum revealed a dramatic redox-dependent increase in sucrose transport activity when heterologously expressed in Saccharomyces cerevisiae. Plant plasma membrane vesicles do not show any change in proton flux across the plasma membrane in the presence of redox reagents, indicating a SUT1-specific effect of redox reagents. Redox-dependent sucrose transport activity was confirmed electrophysiologically in Xenopus laevis oocytes with SUT1 from maize (Zea mays). Localization studies of green fluorescent protein fusion constructs showed that an oxidative environment increased the targeting of SUT1 to the plasma membrane where the protein concentrates in 200-to 300-nm raft-like microdomains. Using plant plasma membranes, St SUT1 can be detected in the detergent-resistant membrane fraction. Importantly, in yeast and in plants, oxidative reagents induced a shift in the monomer to dimer equilibrium of the St SUT1 protein and increased the fraction of dimer. Biochemical methods confirmed the capacity of SUT1 to form a dimer in plants and yeast cells in a redox-dependent manner. Blue native PAGE, chemical cross-linking, and immunoprecipitation, as well as the analysis of transgenic plants with reduced expression of St SUT1, confirmed the dimerization of St SUT1 and Sl SUT1 (from Solanum lycopersicum) in planta. The ability to form homodimers in plant cells was analyzed by the split yellow fluorescent protein technique in transiently transformed tobacco (Nicotiana tabacum) leaves and protoplasts. Oligomerization seems to be cell type specific since under native-like conditions, a phloem-specific reduction of the dimeric form of the St SUT1 protein was detectable in SUT1 antisense plants, whereas constitutively inhibited antisense plants showed reduction only of the monomeric form. The role of redox control of sucrose transport in plants is discussed.

Suc transporters (SUTs) play a key role in the allocation and partitioning of photosynthetically fixed carbon in plants. While a function could be assigned to many members of the SUT family, almost no information is available on their regulation. Here, the transcriptional regulation of SUTs in response to various environmental stimuli in the leaves of five dicots (Arabidopsis Extensive data on expression of SUTs in relation to changes of environmental conditions were obtained through a global analysis of 168 transcriptomics data sets. Results were validated by quantitative PCR measurements and extended by the measurement of photosynthesis rate and phloem sugar content to draw insight on the correlation of SUT expression and sugar export from leaves. For the apoplasmic phloem loaders, a clear difference in transcriptional regulation in response to different environmental stimuli was observed. The consistent patterns of SUT expression under abiotic stress indicates which types of SUTs are involved in the regulation of leaf sugar status and in stress signaling. Furthermore, it is shown that down-regulation of phloem loading is likely to be caused by transcriptional regulation of SUTs, while up-regulation depends on post-transcriptional regulation. In poplar, expression of PtaSUT4 was found to consistently respond to environmental stimuli, suggesting a significant role in the regulation of sugar export from leaves in this passive symplasmic phloem loader.

Since Münch in the 1920s proposed that sugar transport in the phloem vascular system is driven by osmotic pressure gradients, his hypothesis has been strongly supported by evidence from herbaceous angiosperms. Experimental constraints made it difficult to test this proposal in large trees, where the distance between source and sink might prove incompatible with the hypothesis. Recently, the theoretical optimization of the Münch mechanism was shown to lead to surprisingly simple predictions for the dimensions of the phloem sieve elements in relation to that of fast growing angiosperms. These results can be obtained in a very transparent way using a simple coupled resistor model. To test the universality of the Münch mechanism, we compiled anatomical data for 32 angiosperm and 38 gymnosperm trees with heights spanning 0.1-50 m. The species studied showed a remarkable correlation with the scaling predictions. The compiled data allowed calculating stem sieve element conductivity and predicting phloem sap flow velocity. The central finding of this work is that all vascular plants seem to have evolved efficient osmotic pumping units, despite their huge disparity in size and morphology. This contribution extends the physical understanding of phloem transport, and will facilitate detailed comparison between theory and field experiments.