Razumovskaya A.V. Cytology of the minor-vein phloem in 320 species from the subclass Asteridae suggests a high diversity of phloem-loading modes. Frontiers in Plant Science. 2013, V. 4, Article 312.

Batashev et al Minor vein phloem in Asteridae and phloem loading in leaves of Sonchus oleraceus (Asteraceae), a species with m inor vein transfer cells. Am. /. Bot. 7 8 ,1 5 4 9 -1 5 5 9 . doi: 10.2307/2444980 Gamalei, Yu. V. (1974). Symplastic con­ nections in the leaf m inor veins of Fraxinus (to the question about sugar transport in leaves). Bot. Zh. 59, 980-988. Gamalei, Yu. V. (1989). Structure and function of leaf m inor veins in trees and herbs. Trees 3, 9 6 -11 0 . doi: 10.1007/BF01021073 Gamalei, Yu. V. (1990). Leaf Phloem. Leningrad: Nauka. Gamalei, Yu. V. (1991). Phloem load­ ing and its development related to plant evolution from trees to herbs. Trees 5, 5 0-6 4 . doi: 10.1007/ BF00225335 Gamalei, Yu. V. (2000). Comparative anatom y and physiology of ter­ minal bundles and surrounding parenchym a in leaves of Dicots. Bot. Zh. 85, 3 4 -4 9 . Gamalei, Yu. V. (2007). Evolution of carbohydrate nutrition in the Scrophulariaeae family. Bot. Zh. 92, 1793-1809. Gamalei, Yu. V., Batashev, D. R., and Pakhomova, М . V. (2008). Organization of the m inor vein phloem in the Rubiaceae family in relation with its phylogeny. Bot. Zh. 9 3 ,1 8 4 6 -1 8 6 2 . Gamalei, Yu. V., Pakhomova, М . V., Sjutkina, A. V., and Voitsekhovskaja, О. V. (2000). Compartm entation of assimilate fluxes in leaves. I. Ultrastructural responses of m es- ophyll and com panion cells to the alteration of assimilate export. Plant Biol 2, 9 8 -1 0 6. doi: 10.1055/ s-2000-9151 Gil, L., Yaron, I., Shalitin, D., Sauer, N., Turgeon, R., and Wolf, S. (2011). Sucrose transporter plays a role in phloem loading in CMV-infected m elon plants that are defined as symplastic loaders. Plant J. 66, 3 66-374. doi: 10.111 l/j.l3 6 5 -3 1 3 X .2 0 1 1. 04498.x Goggin, F. L., Medville, R., and Turgeon, R. (2001). Phloem loading in the tulip tree. M echanism and evolutionary implications. Plant Physiol. 125, 8 9 1-8 9 9. doi: 10.1104/pp. 125. 2.891 Gunning, В. E. S., and Pate, J. S. (1969). ‘Transfer cells’. Plant cells with wall ingrowths, specialized in relation to short distance transport of solutes - their occurrence, structure, and development. Protoplasma 68, 107-133. doi: 10.1007/ B F01247900 Haritatos, E., Medville, R., and Turgeon, R. (2000). M inor vein structure and sugar trans­ port in Arabidopsis thaliana. Planta 211, 105-111. doi: 10.1007/s004250000268 H offm ann-Thom a, G., van Bel, A. J. E., and Ehlers, K. (2001). Ultrastructure of m inor-vein phloem ans assimilate export in summ er and winter leaves of the symplasmically loading evergreens Ajuga reptans L., Aucuba japon- ica Thunb., and H edera helix L. Planta 212, 2 3 1 -2 4 2 . doi: 10.1007/ S004250000382 Kempers, R., Amm erlaan, A., and van Bel, A. J. E. (1998). Symplasmic constriction and ultrastructural features of the sieve elem ent/com panion cell com plex in the transport phloem of apoplasmically and symplas­ mically phloem-loading species. Plant Physiol. 116, 27 1 -27 8 . doi: 10.1104/pp.l 16.1.271 Kursanov, A. L. (1984). Assimilate Transport in Plants. Amsterdam : Elsevier. Lundberg, J. (2009). “Asteraceae and relationships within Asterale,” in Systematics, Evolution, and Biogeography o f Compositae, eds V. A. Funk, A. Susanna, T. F. Stuessy, and R. J. Bayer (Vienna: IAPT), 157-169. M adore, M . A., and Grodzinski, B. (1984). Effect of oxygen concen­ tration on C-photoassimilate trans­ port from leaves of Salvia splendens L. Plant Physiol. 76, 782-786. doi: 10.1104/pp.76.3.782 M adore, M . A., Oross, J. W ., and Lucas, W . J. (1986). Symplastic transport in Ipom ea tricolor source leaves: dem onstration of functional symplastic connections from m es- ophyll to m inor veins by a novel dye-tracer m ethod. Plant Physiol. 82, 4 3 2 ^ 4 2 . doi: 10.1104/pp.82. 2.432 M cCauley, М . М ., and Evert, R. F. (1989). M inor veins of the potato (Solanum tuberosum L.) leaf: ultra­ structure and plasmodesmatal fre­ quency. Bot. Gaz. 150, 3 51-368. doi: 10.1086/337781 M oore, B. D., Palmquist, D. E., and Seemann, J. R. (1997). Influence of plant growth at high CO 2 concentrations on leaf content of ribulose-1, 5-biphosphate carboxy­ lase/oxygenase and intracellular distribution of solublecarbohy- drates in tobacco, snapdragon, and parsley. Plant Physiol. 115, 24 1-2 4 8. M oore, M . J., Soltis, P. S., Bell, C. D., Gordon Burleigh, J., and Soltis, D. E. (2010). Phylogenetic anal­ ysis of 83 plastid genes further resolves the early diversification of eudicots. Proc. Natl. Acad. Sci. U.S.A. 107, 46 2 3-4 6 28 . doi: 10.1073/pnas.0907801107 Offler, С. E., McCurdy, D. W ., Patrick, J. W ., and Talbot, M . J. (2003). Transfer cells: cells specialized for a special purpose. Ann. Rev. Plant Biol. 54, 4 3 1 -4 5 4. doi: 10. 1146/annurev.arplant.54.031902. 134812 Pate, J. S., and Gunning, В. E. S. (1969). Vascular transfer cells in angiosperm leaves, a taxo­ nom ic and morphological survey. Protoplasma 68, 135-156. doi: 10.1007/BF01247901 Peterson, R. L., and Yeung, E. C. (1975). Ontogeny of phloem trans­ fer cells in H ieracium floribundum. Can. J. Bot. 53, 274 5 -2 7 5 8 . doi: 10.1139/b75-303 Reidel, E. J., Rennie, E. A ., Amiard, V., Cheng, L., and Turgeon, R. (2009). Phloem loading strate­ gies in three plant species that transport sugar alcohols. Plant Physiol. 149, 1601-1608. doi: 10.1104/p p .l08.134791 Rennie, E. A., and Turgeon, R. (2009). A comprehensive picture of phloem loading strategies. Proc. Natl. Acad. Sci. U.S.A. 106, 14162-14167. doi: 10.1073/pnas.0902279106 Roberts, A. G., Santa Cruz, S., Roberts, I. М ., Prior, D. A. М ., Turgeon, R., and Oparka, K. J. (1997). Phloem unloading in sink leaves of Nicotiana benthamiana: com ­ parison of a fluorescent solute with a fluorescent virus. Plant Cell 9, 1381-1396. Russin, W . A ., and Evert, R. F. (1985). Studies on the leaf of Populus deltoids (Salicaceae): ultrastructure, plasmodesmatal fre­ quency, and solute concentrations. Am. J. Bot. 72, 1232-1247. doi: 10.2307/2443404 Schmitz, K., Cuypers, B., and Moll, M. (1987). Pathway of assimilate trans­ fer between mesophyll cells and m inor veins in leaves of Cucumis melo L. Planta 171, 19-29. doi: 10.1007/B F00395064 Tangl, E. (1879). Ueber offene kom- munikationen zwischen den zellen des endosperms einiger samen. Jahrb. Wiss. Bot. 12, 170-190. Tešitel, J., Plavcova, L., and Cam eron, D. D. (2010). Heterotrophic carbon gain by the root hemiparasites, Rhinanthus m inor and Euphrasia rostkoviana (Orobanchaceae). Planta 231, 1137-1144. doi: 10.1007/s00425-010-l 114-0 Turgeon, R., Beebe, D. U., and Gowan, E. (1993). The interm ediary cell: minor-vein anatom y and raffi- nose oligosaccharide synthesis in the Scrophulariaceae. Planta 191, 44 6 -4 5 6 . doi: 10.1007/B F00195746 Turgeon, R., and Medville, R. (2004). Phloem loading. A reevalua­ tion of the relationship between plasmodesmatal frequencies and loading strategies. Plant Physiol. 136, 3 7 95-3803. doi: 10.1104/pp. 104.042036 Turgeon, R., Medville, R., and Nixon, К. C. (2001). The evolution of m inor vein phloem and phloem loading. Am. J. Bot. 88, 1331-1339. doi: 10.2307/3558441 Turgeon, R., Webb, J. A., and Evert, R. F. (1975). Ultrastructure of m inor veins in Cucurbita pepо leaves. Protoplasma 83, 2 1 7-2 3 1. doi: 10.1007/BF01282555 Tyree, М . T. (1970). The symplast concept. A general theory of symplastic transport according to the thermodynam ics of irre­ versible processes. J. Theor. Biol. 26, 181-214. doi: 10.1016/S0022-5193 (70)80012-1 van Bel, A. J. E., and Gamalei, Yu. V. (1992). Ecophysiology of phloem loading in source leaves. Plant Cell Environ. 15, 26 5 -2 7 0 . doi: 10.1111/j.l365- 3040.19 92.tb00973.x van Bel, A. J. E., Gamalei, Yu. V., Amm erlaan, A., and Bik, L. P. M . (1992). Dissimilar phloem load­ ing in leaves with symplasmic of apoplastic m inor-vein configura­ tions. Planta 186, 51 8 -5 2 5 . doi: 10.1007/B F00198031 van Bel, A. J. E., and Hafke, J. B. (2005). “Physiochemical determinants of phloem transport,” in Vascular Transport in Plants , eds N . M . Holbrook and M . A. Zwieniecki (Am sterdam : Elsevier Academic Press), 19^14. Voitsekhovskaja, О. V. (2002). On the Role o fSugar Compartmentation and Stachyose Synthesis in Symplastic Phloem Loading. Dissertation, Cuvillier Verlag, Goettingen. Voitsekhovskaja, О. V., Koroleva, O. A ., Batashev, D. R., Knop, C., Tomos, A. D., Gamalei, Yu. V., et al. (2006). Phloem loading in two Scrophulariaceae species. W hat can drive symplastic flow via plasmodesmata? Plant Physiol. 140, 3 8 3 -3 9 5 . doi: 10.1104/pp.l05. 068312 Voitsekhovskaja, О. V., Rudashevskaya, E. L., Demchenko, K. N., Pakhomova, М . V., Batashev, D. R., Gamalei, Yu. V., et al. (2009). Evidence for functional www.frontiersin.org August 2013 | Volume 4 | Article 312 | 13