In the leaves of dicotyledons two modes of phloem loading have been identified (Turgeon and Wimmers, 1988; van Bel et al., 1992van Bel et al., , 1994. These mechanisms of phloem loading, symplasmic or apoplasmic, seem to be associated with minor vein configuration (Gamalei, 1985(Gamalei, , 1989) and carbohydrate metabolism (Gamalei, 1985;Turgeon et al., 1993;Flora and Madore, 1996). CCs in minor veins of apoplasmically phloem-loading species (further referred to as apoplasmic species) are termed TCs and have virtually no plasmodesmata at the interface with the mesophyll domains (Gamalei, 1989). The TCs possess cell wall protrusions, varying in surface area with the transit of photosynthate, and unfragmented vacuoles (Gamalei, 1989; Wimmers and Turgeon, 1991;Gamalei et al., 1992). CCs in minor veins of symplasmically phloem-loading species (further referred to as symplasmic species) are termed ICs and are connected with the mesophyll symplast via numerous plasmodesmata (Gamalei, 1989). The ICs usually contain vesicular networks or heavily fragmented vacuoles and have no cell wall protrusions (Gamalei, 1989).The most persuasive evidence for two phloem-loading mechanisms is the consistent coincidence between physiological behavior and minor vein configuration. The diverse structure-functional indications in favor of two modes of phloem loading are numerous and were thoroughly reviewed by van Bel (1996). More recent evidence supports the existence of principally different systems of phloem loading (Flora and Madore, 1996;Kingston-Smith and Pollock, 1996). The question arises whether the different ways of carbohydrate processing in the phloem-loading zone continue to exist along the phloem trajectory, of which ultrastructure and plasmodesmal connectivity of the CCs in the transport phloem may be indicative. Hence, the present electronmicroscopic investigation was focused on the ultrastructure of the CCs in the transport phloem of two species that were described to be symplasmic, squash (Cucurbita maxima L.;Gamalei, 1991) and Lythrum salicaria L. (van Bel et al., 1994), and two species that were considered to be apoplasmic, broad bean (Vicia faba L.; Gamalei, 1991) and Zinnia elegans L. (Y.V. Gamalei and A.V. Sjutkina, unpublished results), to obtain an impression of the functional continuity between collection and transport phloem in apoplasmic and symplasmic loaders.
MATERIALS AND METHODSPhloem specimens were cut from the stems of four species, broad bean (Vicia faba L. cv Witkiem major [Nunhems Zaden bv, Haelen, The Netherlands]), Lythrum salicaria L., Zinnia elegans L. (bv Cruydthoeck, Groningen, The Netherlands), and squash (Cucurbita maxima L. cv Golden Deli-1 A part of this study was subsidized by NWO (Dutch Organization for Scientific Research).