GENERAL DISCUSSION Cloning Trehalose Synthase Gene And Its Expression In Arabidopsis thaliana

followed by a cluster analysis based on the characters suggests that 33 of the transgenic plants resistant to drought. From leaf water recovery test, it was observed 20 of leaves recover after 16h keeping on RT. Hence it might be suggested that there is possibility about 30 to obtain transgenic plant with improved drought response by introducing trehalose synthase gene from Thermobifida fusca . It was suggested that the improved response toward stress of transgenic plant bearing trehalose metabolizing genes was due to the increase of soluble carbohydrate as result of the increase of photosynthetic capacity Garg et al 2002. While Jang et al 2003 suggested, trehalose give global protection against abiotic stresses, yet, they did not suggest the mechanism. While Avonce et al. 2004 suggested that AtTPS1 is regulator of glucose, abcisic acid, and stress signaling. In this experiment, the capability of some TfTreS expressing lines withstand drought might also has been implicated with the increase of T6P. Arabidopsis expressing AtTPS1 is sugar and ABA insensitive, and T6P regulate the expression of ABI4. ABI4 is down regulated by T6P when glucose is presence and is up regulated in the absence of metabolizable sugar. T6P affect growth and development Schluepmann et al 2004, mutant of AtTPS1 is embryo lethal. Hence, trehalose metabolism is indispensable for growth Easmond et la. 2002; Schluepmann et al. 2004. This effect on growth was also shown on M. smegmatis Woodruff et al 2007. The presence of T6P, however, leads to induction of APL3 hence starch is accumulated in chloroplast and growth is arrested. Feeding trehalose to seedlings increased T6P level Schluepmann et al. 2004. Expressing TfTreS that increase trehalose level, might increase increase T6P level on plant in some extend, as starch accumulation was proved occurred to some lines Figure 33. The increase, of T6P however, did not retard growth that might be due to the reverse TfTreS activity that still provides sugar for energy source. Furthermore, the presence of the sugar down regulates ABI4, hence ABA production is reduced, and growth retardation is rendered Avonce et al. 2004. In more general about the linking of trehalose and ABA is suggested by Brodmann 2006. He revealed that application of 25 mM trehalosevalidamycin incresease or decrease the expression of 2277 genes, and more than a third of them are regulated by ABA, linking trehalose to ABA or ABA signaling. Interestigly, on ABA deficient mutant abi-1 the application of trehalose and validamycin affect ABA inducible genes, indicating that the induction of Trehalosevalisdamycin A is independent from ABA. However, some the induced genes and ABA induced genes were sppressed in mutant abi2-1, suggesting that signaling cascade triggered by trehalosevalidamycin affect the ABA sigbaling pathway downstream of the ABA binding site but upstream of ABI2 activity. He suggested further that those genes that are regulated by trehalosevalidamycin but not by ABA are those involve in disease resistant and secondary metabolites, hence trehalose act also as an elicitor in plant. Figure 33. Starch staining of seedlings grown on trehalose or ½ MS. TfTreS expressing leave grown on 12 MS A and its magnification C or on trehalose B and its magnification D. Leave of sensitive as Wt grown on 12MS E and its magnification G or on trehalose F, and its agnification H. Some TfTreS expressing plants accumulated starch I. It is very recently reported new role of trehalose that controls the dynamic mRNA of heat shock protein Hsf1 by activating transcriptional activity. This acceleration and the presence of trehalose enhance phosphorylation of the protein Collin and C A B D E D C F G H C D I J I G H Nelson 2007. This report support previous suggestions that trehalose prevent protein denaturation and agregation of denaturated protein as heat shock protein involves in molecular chaperon that refold protein back to natural structure. TfTreS plants resistance to trehalose open a question about the possibility of the used of TfTreS as selectable marker on trehalose. As earlier discussed there are several non toxic selectable marker has been developed, to meet the public concern about bio safety of the use antibioticherbicide resistance genes. Method to engineer marker free transgenic plants is also developing, however, analog to Darbani et al. 2007 suggesting that one of the methods to produce free marker transgenic plant is by using non toxic selectable marker. Selection TfTreS on trehalose as well as on a combination of trehalose and validamycin A, may obtain marker free transgenic plants. The combination is stronger selection system, since validamycin A is a trehalase inhibitor that keeps trehalose remain un-degraded and increases trehalose level within tissue Godijn et al. 1997; Muller et al. 2001. As the selection system gave transgenic lines, hence, both selection system can be used with considerable amount of transgenic plants obtained. The combination, however, gave fewer lines than when trehalose alone is used as the selection agent. None of the wild type plants survived either on trehalose alone 100 mM or on the combination 10 µ M validamycin A and 50 mM or more trehalose. These lead to the conclusion that TreS can be used as a selectable marker. The use of the gene as an alternative selectable marker might be favorable to the transgenic obtained for some reasons. First, about 30 of transgenic plants might withstand stresses, 2 the product of the gene, trehalose, is biologically friendly and has been used widely for many aspects of human life and 3 the selection agent is harmless sugar to the environment. The validamycin A as the waste after selection is biodegradable with half life less than 5h in soil Sticher et al. 1997.

VIII. CONCLUSION AND RECOMMENDATION

Gene encoding for trehalose synthase form Thermobifida fusca and from Mycobacterium tuberculosis have been successfully cloned within cloning vector pGemT and constructed within plant expression cassette. The genes were introduced to Arabidospsis thaliana as plant model to see the effect the gene on plant responses to stresses, especially to drought. TreS MtTreS from M. tuberculosis , however, has not been successfully introduced into the plant. TreS from T. fusca TfTreS was functionally active in Arabidopsis thaliana and was inherited to the next generations. About 17 of the transgenic plants contained high level of trehalose ranging from 42-65 µ M gr -1 FW. While the enzyme activity of TfTreS expressing lines was significantly higher than the control plants. Drought either tested in leaves using drought parameters or in planta by withholding water suggests that about one third 33 of TfTreS expressing plants withstand drought. The capability of TRES to convert maltose to trehalose but still provide metabolizable sugar allowed TfTreS expressing plants survive on high trehalose level, as well as trehalose in combination with validamycin A, ensuring the construct and its selection system could be used as an alternatives among useful non toxic selectable markers. It might be To see the influence of TreS in plant responses to stresses, it is necessary to reassess the transgenic plants against some environmental stresses with involvement of higher number of plants with wider aspects of observation to obtain a better feature about the presence of the gene toward stresses. List of References Alarico, S., Empadinhas, N., Simoes, C., Silva, Z., Henne, A., Mingote, A., Santos, H., da Costa, M. S. 2005. Distribution of Genes for Synthesis of Trehalose and Mannosylglycerate in Thermus spp. and Direct Correlation of These Genes with Halotolerance. Appl. Environ. Microbiol. 71: 2460-2466. Almeida AM, villalobos E, Ara’ujo Aa, Leyman B, Van Dijk P, Alfaro-Cardoso L, Fevereiro PS, Torn’e JM and Santos DM. 2005. Transformation of tobacco with an Arabidopsis thaliana gene involved in trehalose biosynthesis increases tolerance to several abiotic stresses. Euphytica 2005 146: 165–176 Elbein A D, Pan YT, Pastuszak I, and Carroll D 2003. New insights on trehalose: a multifunctional molecule. Glycobiology . 13: 14-27. Alpert P 2006. Constraints of tolerance: why are desiccation-tolerant organisms so small or rare?. J. Exp. Biol. 2099: 1575 – 1584 Anonym www.agronomy.psu.edu Anonym www.nature.com Anonym www.sbcu.ac.uk. Anonym www.inspection.gc.ca Avonce N, leyman B, Mascorro-Gallardo JO, Van Dijk P, Thevelein JM and Iturriaga G 2004. The Arabidopsis Trehalose-6-P Synthase AtTPS1 Gene Is a Regulator of Glucose, Absisic Acid, and Stress Signaling. Plant Physiol. 136: 1649-3659. Avonce N, Mendoza-Vargas A, Morett and Iturriaga G 2006. Insight on the evolution of trehalose biosynthesis. BMC Evol. Biol. 6109: 1471-2148 Balmer Y, Vensel WH, Cai N, Manieri W, Schurmann P, Hurkman WJ, and Buchanan BB 2006. A complete ferredoxinthioredoxin system regulates fundamental processes in amyloplasts. PNAS. 1038: 2988 - 2993. Blazquez MA, Stucka R, Feldmann H and Ganzedo C 1994. Trehalose-6-P synthase is Dispensible for Growth and Glucose but not for Spore Germination in Schizosaccharimyces pombe. J. Bacteriol. 176 13: 3895-3902. Bayles BB, Taylor JW and Bartel AT 1937. Rate of Water Loss in Wheat Cultivars and Resistance to Artificial Drought. J. Am. Soc. Agron. 29: 40-52 Benourouj N, Lee DH, and Goldnerg AL 2001. Trehalose Accumulation During Cellular Stress Protect Cell and cellular Protein From Damage by Oxygen Radicals. J. Biol. Chem. 276 26: 24261-242267. Blum A online.Towards Standard Assays of Drought Resistance in Crop Plants. CIMMYT. Workshop on Molecular Approaches for the genetic Improvement of Cereals for Stable Production in Water-Limited Environments. Download date: june 2007 Bonini, B.M., vanVaeck, C., Larsson, C., Gustafsson, L., Ma, P., Winderickx, J., vanDijck, P., and Thevelein, J.M. 2000 Expression of Escherichia coli otsA in a Saccharomyces cerevesiae tps1 mutant restores trehalose-6-P levels and partially restores growth and fermentation with glucose and control of glucose influx into glycolysis. Biochem. J. 350, 261–268.