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The phloem unloading pathway remains unclear in fleshy fruits accumulating a high level of soluble sugars. A structural investigation in apple fruit (Malus domestica Borkh. cv Golden Delicious) showed that the sieve element-companion cell complex of the sepal bundles feeding the fruit flesh is symplasmically isolated over fruit development. 14 C-autoradiography indicated that the phloem of the sepal bundles was functional for unloading. Confocal laser scanning microscopy imaging of carboxyfluorescein unloading showed that the dye remained confined to the phloem strands of the sepal bundles from the basal to the apical region of the fruit. A 52-kD putative monosaccharide transporter was immunolocalized predominantly in the plasma membrane of both the sieve elements and parenchyma cells and its amount increased during fruit development. A 90-kD plasma membrane H 1 -ATPase was also localized in the plasma membrane of the sieve element-companion cell complex. Studies of [ 14 C]sorbitol unloading suggested that an energy-driven monosaccharide transporter may be functional in phloem unloading. These data provide clear evidence for an apoplasmic phloem unloading pathway in apple fruit and give information on the structural and molecular features involved in this process.The partitioning of sugars in economically important sink organs such as fruits or seeds is governed by several complex physiological processes, including photosynthetic rate, phloem loading in the source leaf, long-distance translocation in the phloem, phloem unloading in sink organs, postphloem transport, and metabolism of imported sugars in sink cells (Oparka, 1990;Patrick, 1997). It is now well accepted that phloem unloading plays a key role in the partitioning of photoassimilate (Fisher and Oparka, 1996;Patrick, 1997;Viola et al., 2001). The process of phloem unloading has been studied extensively over the last 20 years (for review, see Oparka, 1990;Patrick, 1997;Schulz, 1998) but still remains poorly understood. Elucidation of the cellular pathway of phloem unloading is central to this process, because, to a large extent, the unloading path determines the key transport events responsible for assimilate movement from the sieve elements (SEs) to the recipient sink cells (Fisher and Oparka, 1996;Patrick, 1997). A symplasmic phloem unloading pathway predominates in most sink tissues such as vegetative apices (Oparka et al., 1995;Patrick, 1997;Imlau et al., 1999), sink leaves (Roberts et al., 1997;Imlau et al., 1999;Haupt et al., 2001), and potato tubers that represent a typical terminal vegetative storage sink (Oparka and Santa Cruz, 2000;Viola et al., 2001). Symplasmic unloading is also efficient in the maternal tissues of developing seeds that represent a class of terminal reproductive storage sinks (Ellis et al., 1992;Patrick et al., 1995;Wang et al., 1995a;Patrick, 1997), although transfer of solutes to the apoplasm may occur at some point along the postphloem pathway (Patrick, 1997). In some cases, symplasmic unloading also occurs in elongating z...

Bud burst is a decisive process in plant architecture that requires light in Rosa sp. This light effect was correlated with stimulation of sugar transport and metabolism in favor of bud outgrowth. We investigated whether sugars could act as signaling entities in the light-mediated regulation of vacuolar invertases and bud burst. Full-length cDNAs encoding two vacuolar invertases (RhVI1 and RhVI2) were isolated from buds. Unlike RhVI2, RhVI1 was preferentially expressed in bursting buds, and was up-regulated in buds of beheaded plants exposed to light. To assess the importance of sugars in this process, the expression of RhVI1 and RhVI2 and the total vacuolar invertase activity were further characterized in buds cultured in vitro on 100 mM sucrose or mannitol under light or in darkness for 48 h. Unlike mannitol, sucrose promoted the stimulatory effect of light on both RhVI1 expression and vacuolar invertase activity. This up-regulation of RhVI1 was rapid (after 6 h incubation) and was induced by as little as 10 mM sucrose or fructose. No effect of glucose was found. Interestingly, both 30 mM palatinose (a non-metabolizable sucrose analog) and 5 mM psicose (a non-metabolizable fructose analog) promoted the light-induced expression of RhVI1 and total vacuolar invertase activity. Sucrose, fructose, palatinose and psicose all promoted bursting of in vitro cultured buds under light. These findings indicate that soluble sugars contribute to the light effect on bud burst and vacuolar invertases, and can function as signaling entities.

Bud burst in certain species is conditioned by the luminous environment. With roses, the requirement for light is absolute, and darkness totally inhibits bud burst. Few studies have looked into understanding the action of light on the physiological bud burst processes. Here, we show the impact of light on certain components of glucidic metabolism during bud burst. Measurements were taken on decapitated plants of Rosa hybrida L. 'Radrazz' exposed either to darkness, white, blue or R light. Results show that a mobilization of bud and the carrying stem sucrose reserves only takes place in light and accompanies the bud burst. Furthermore, the activity of the RhVI vacuolar acid invertase which contributes to the breakdown of sucrose in the buds, as well as the transcription of the RhVI gene, is reduced in darkness, although it is strongly stimulated by light. The same analysis concerning the RhNAD-SDH gene, coding an NADdependent sorbitol dehydrogenase, shows, on the contrary, a strong induction of its transcription in darkness that could reflect the use of survival mechanisms in this condition.