Plant Physiology Preview
Published on December 23, 2005; 10.1104/pp.105.068312
Received July 15, 2005
Returned for revision October 31, 2005
Accepted November 3, 2005
Phloem Loading in Two Scrophulariaceae Species. What Can Drive Symplastic Flow via Plasmodesmata?
Olga V. Voitsekhovskaja *, Olga A. Koroleva , Denis R. Batashev , Christian Knop , A. Deri Tomos , Yuri V. Gamalei , Hans-Walter Heldt , and Gertrud Lohaus
Albrecht-von-Haller-Institute for Plant Sciences, Plant Biochemistry, 37077 Goettingen, Germany; Komarov Botanical Institute, Russian Academy of Sciences, Plant Ecological Physiology, 197376 St. Petersburg, Russia
School of Biological Sciences, University of Wales, Bangor, Gwynedd, Wales LL57 2UW, United Kingdom
Komarov Botanical Institute, Russian Academy of Sciences, Plant Ecological Physiology, 197376 St. Petersburg, Russia
Albrecht-von-Haller-Institute for Plant Sciences, Plant Biochemistry, 37077 Goettingen, Germany
* Corresponding author; email: ovoitse{at}yandex.ru.
To determine the driving forces for symplastic sugar flux between mesophyll and phloem, gradients of sugar concentrations and osmotic pressure were studied in leaf tissues of two Scrophulariaceae species, Alonsoa meridionalis and Asarina barclaiana. A. meridionalis has a typical symplastic configuration of minor-vein phloem, i.e. intermediary companion cells with highly developed plasmodesmal connections to bundle-sheath cells. In A. barclaiana, two types of companion cells, modified intermediary cells and transfer cells, were found in minor-vein phloem, giving this species the potential to have a complex phloem-loading mode. We identified all phloem-transported carbohydrates in both species and analyzed the levels of carbohydrates in chloroplasts, vacuoles, and cytoplasm of mesophyll cells by nonaqueous fractionation. Osmotic pressure was measured in single epidermal and mesophyll cells and in whole leaves and compared with calculated values for phloem sap. In A. meridionalis, a 2-fold concentration gradient for sucrose between mesophyll and phloem was found. In A. barclaiana, the major transported carbohydrates, sucrose and antirrhinoside, were present in the phloem in 22- and 6-fold higher concentrations, respectively, than in the cytoplasm of mesophyll cells. The data show that diffusion of sugars along their concentration gradients is unlikely to be the major mechanism for symplastic phloem loading if this were to occur in these species. We conclude that in both A. meridionalis and A. barclaiana, apoplastic phloem loading is an indispensable mechanism and that symplastic entrance of solutes into the phloem may occur by mass flow. The conditions favoring symplastic mass flow into the phloem are discussed.
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