Many plants accumulate organic osmolytes in response to the imposition of environmental stresses that cause cellular dehydration. Although an adaptive role for these compounds in mediating osmotic adjustment and protecting subcellular structure has become a central dogma in stress physiology, the evidence in favour of this hypothesis is largely correlative. Transgenic plants engineered to accumulate proline, mannitol, fructans, trehalose, glycine betaine or ononitol exhibit marginal improvements in salt and/or drought tolerance. While these studies do not dismiss causative relationships between osmolyte levels and stress tolerance, the absolute osmolyte concentrations in these plants are unlikely to mediate osmotic adjustment. Metabolic benefits of osmolyte accumulation may augment the classically accepted roles of these compounds. In re-assessing the functional significance of compatible solute accumulation, it is suggested that proline and glycine betaine synthesis may buffer cellular redox potential. Disturbances in hexose sensing in transgenic plants engineered to produce trehalose, fructans or mannitol may be an important contributory factor to the stress-tolerant phenotypes observed. Associated effects on photoassimilate allocation between root and shoot tissues may also be involved. Whether or not osmolyte transport between subcellular compartments or different organs represents a bottleneck that limits stress tolerance at the whole-plant level is presently unclear. None the less, if osmolyte metabolism impinges on hexose or redox signalling, then it may be important in long-range signal transmission throughout the plant.Key-words: betaine; cold stress; drought; fructans; mannitol; osmolytes; proline; salinity; sugar signalling; trehalose. Abbreviations INTRODUCTIONEnvironmental stress is the major factor limiting plant productivity . Abiotic stresses which cause depletion of cellular water (drought, high soil salinity and temperature extremes) are responsible for the greatest agricultural losses. Upon exposure to these prevalent stresses, many plants accumulate organic osmolytes, most commonly polyhydroxylic compounds (saccharides and polyhydric alcohols) and zwitterionic alkylamines (amino acids and quaternary ammonium compounds).Several recent reviews discuss osmolyte accumulation in plants (Ingram & Bartels 1996;Serrano 1996). It is generally accepted that the increase in cellular osmolarity which results from the accumulation of non-toxic (thus 'compatible') osmotically active solutes is accompanied by the influx of water into, or at least a reduced efflux from, cells, thus providing the turgor necessary for cell expansion. None the less, a conclusive demonstration that osmotic adjustment contributes to fitness in stressful environments has yet to be achieved (Munns 1993). Since all subcellular structures must exist in an aqueous environment, tolerance to dehydration also depends on the ability of cells to maintain membrane integrity and prevent protein denaturation. Hypotheses that attribut...
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Selectable marker genes are required to ensure the efficient genetic modification of crops. Economic incentives and safety concerns have prompted the development of several strategies (site-specific recombination, homologous recombination, transposition, and co-transformation) to eliminate these genes from the genome after they have fulfilled their purpose. Recently, chemically inducible site-specific recombinase systems have emerged as valuable tools for efficiently regulating the excision of transgenes when their expression is no longer required. The implementation of these strategies in crops and their further improvement will help to expedite widespread public acceptance of agricultural biotechnology
SummaryPhytochrome A (phyA) plays a primary role in initiating seedling de-etiolation and is the only plant photoreceptor known to be activated by far-red light (FR). The signaling intermediate FHY1 appears to either participate directly in relaying the phyA signal or to positively regulate a critical signaling event(s) downstream of phyA activation. Here we identify a homolog of FHY1 named FHL (FHY1-like) as a novel signaling factor essential for complete responsiveness to phyA. FHL possesses functional nuclear localization and nuclear export signals. Lines in which FHL function was abolished by insertional mutagenesis or attenuated by RNAimediated suppression displayed a weaker hyposensitivity to continuous FR than fhy1 null mutants and most reported phyA signaling mutants. However, hypocotyl elongation assays indicated that suppression of FHL expression in fhy1-3 caused an insensitivity of hypocotyl elongation to FR and blue light (B) indistinguishable from that seen in phyA. Real-time PCR indicates that in FR, FHY1 transcripts are approximately 15-fold more abundant than FHL transcripts. Although both FHY1 and FHL are capable of homo-and hetero-interaction via their C-termini, the ability of FHL overexpression to restore wild-type (WT) morphological and molecular phenotypes to fhy1-3 seedlings suggests that the extreme insensitivity to FR associated with suppression of FHL expression in fhy1-3 cannot be accounted for by a critical role for FHY1-FHL heterodimers in phyA signal transmission. Rather, we suggest that the relative abundances of FHY1 and FHL in WT plants account for the differences in the severity of fhy1 and fhl mutations. As for FHY1, FHL transcript accumulation is dependent on FHY3 and is decreased after exposure to FR, R or B light. These findings reiterate the prevalence of partial degeneracy in plant signaling networks that regulate responses crucial to survival.
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