Physiology and Molecular Biology of Stress ... - KHAM PHA MOI

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Metabolic Engineering for Stress Tolerance 261 Table 2. Select “milestone” studies on metabolic engineering for stress tolerant plants Mechanism Stress tolerance References Fatty acid unsaturation Chilling Murata et. al. (1992) Antioxidation Oxidative stress, drought Gupta et. al. (1993a & 1993b) Compatible solute synthesis Salinity, cold and freezing Tarczynski et. al. (1993) Heat shock proteins and Heat Lee et. al. (1995) Chaperones LEA proteins Salinity, Drought Xu et. al. (1996) Glutathione and amino Oxidative stress, Roxas et. al. (1997) acid metabolism drought, salinity Transcriptional control Cold, drought Jaglo-Ottosen et. al. of stress pathways (1998) Sodium vacuolar transport Salinity Apse et. al (1999) High concentrations of compatible solutes are non-toxic to the cells. So transgenic approaches can be used to facilitate their accumulation in crop plants for long-term osmotic stress tolerance (Chen and Murata, 2002; Serraj and Sinclair, 2002). Natural accumulation of osmoprotectants varies in plants i.e. 5-50 ìmol g -1 fresh weight (~6-60 mM on a plant water basis) and it often increases several-fold during exposure to osmotic stress (Rhodes and Hanson, 1993; Bohnert et. al., 1995). In plant cells, osmoprotectants are normally restricted to the cytosol, chloroplasts, and other cytoplasmic compartments. Many crop plants do not naturally synthesize high levels of osmoprotectants. Therefore, it was hoped that crop stress tolerance could be improved by introducing one or many genes implicated in the synthesis of a specific osmoprotectant. Hence, metabolic engineering of synthetic pathways to osmoprotectants has been vigorously pursued during the last decade. Some examples for metabolic engineering of plants for osmoprotectant synthesis are listed in Table 3 and are discussed below. 3.1.1. Mannitol The sugar alcohol mannitol is synthesized in numerous species of plants and its accumulation increases under dehydration stress (Patonnier et al., 1999). In celery, a natural accumulator of mannitol, its synthesis from mannose-6-phosphate is catalyzed by mannose-6-phosphate reductase (M6PR). Under stress, there is down-regulation of
262 B. Rathinasabapathi and R. Kaur Figure 2 . Structures of some osmoprotectants found in stress-tolerant organisms mannitol’s utilization (Williamson et al., 1995). In the first demonstration of engineering for mannitol overproduction in plants, the gene for M6PR was not used. Instead, a bacterial gene for the reversible mannitol-1-phosphate dehydrogenase (mtlD), which catalyzes the conversion of fructose-6-phosphate to mannitol-1-phosphate was employed. This led to mannitol synthesis in transgenic tobacco, resulting in an osmotolerance phenotype against salinity (Tarczynski et. al., 1993; Karakas et. al., 1997). In independent experiments, transgenic tobacco with increased mannitol in the chloroplasts was shown to exhibit increased resistance to methyl viologen-induced oxidative stress (Shen et. al., 1997). Transgenic wheat expressing the bacterial mtlD gene also exhibited tolerance to water stress and salinity (Abebe et. al., 2003), despite the lack of osmotically significant quantities of mannitol in these transgenic plants. This suggested that mannitol’s effects are from protective mechanisms other than osmotic adjustment or via mannitol’s osmoprotective role in the meristematic regions (Abebe et. al., 2003). Arabidopsis, a non-mannitol producer, was transformed with the celery leaf M6PR gene under the control of the CaMV 35S promoter. Transgenic plants accumulated from 0.5 to 6 ìmoles g -1 fresh weight mannitol. Salt tolerance in transgenic plants was demonstrated in soil irrigated with 300 mM NaCl in the nutrient solution (Zhifang and Loescher, 2003). Pathways to other sugar alcohols have also been successfully installed in transgenic plants and stress tolerance phenotypes have been observed (Sheveleva et. al., 1997).
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PHYSIOLOGY AND MOLECULAR BIOLOGY OF
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A C.I.P. Catalogue record for this
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About the Editors K.V. Madhava Rao
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LIST OF CONTRIBUTORS K. AKASHI Grad
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List of Contributors xiii NAVINDER
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PREFACE Increasing agricultural pro
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2 K.V. Madhava Rao Abiotic stresses
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4 K.V. Madhava Rao SOME O THE PROMI
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6 K.V. Madhava Rao 2. WATER STRESS
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8 K.V. Madhava Rao 5. FREEZING STRE
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10 K.V. Madhava Rao of these pathwa
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12 K.V. Madhava Rao Bray, E.A. (199
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14 K.V. Madhava Rao Rao, K.V. Madha
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16 A. Yokota, K. Takahara and K. Ak
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41 CHAPTER 3 SALT STRESS ZORA DAJIC
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Salt Stress 43 activities (mainly i
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Salt Stress 45 In summary, mechanis
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Salt Stress 47 tolerance research i
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Salt Stress 49 need to rely on sodi
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Salt Stress 51 (Echeverria, 2000).
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Salt Stress 53 Therefore, the capac
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Salt Stress 55 Reduced plant growth
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Salt Stress 57 Table 3. Salt tolera
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Salt Stress 59 6.2. Nitrogen Fixati
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Salt Stress 61 A significant number
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Salt Stress 63 macromolecules, irre
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Salt Stress 65 8.2. Ion Homeostasis
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Salt Stress 67 1997), is speculated
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Salt Stress 69 together with the At
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Salt Stress 71 important role in si
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Salt Stress 73 Figure 5. Determinan
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Salt Stress 75 9.1.Transgenic Plant
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Salt Stress 77 tolerance from halop
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Salt Stress 79 sponse and yield (Su
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Salt Stress 81 Table 5. Possible ut
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Salt Stress 83 monitored with fluor
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Salt Stress 85 Func. Plant Biol. 29
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Salt Stress 87 Dajic, Z., Stevanovi
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Salt Stress 89 Gouia, H., Ghorbal,
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Salt Stress 91 Larcher, W. (1995).
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Salt Stress 93 Munns, R. and James,
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Salt Stress 95 Rausell, A., Kanhono
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Salt Stress 97 durum wheat crops gr
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Salt Stress 99 Yoshida, K. (2002).
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102 T.D. Sharkey and S.M. Schrader
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131 CHAPTER 5 FREEZING STRESS: SYST
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Freezing Stress 133 Whereas, in the
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Freezing Stress 135 genes at the tr
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Freezing Stress 137 with physiologi
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Freezing Stress 139 (1997). However
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Freezing Stress 141 (Barnett et al.
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Freezing Stress 143 (dehydrin) prot
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Freezing Stress 145 in cytosolic Ca
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Freezing Stress 147 Phospholiphase
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Freezing Stress 149 Accumulation of
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Freezing Stress 151 Ideker, T., Gal
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Freezing Stress 153 ellin acid on f
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Freezing Stress 155 Yoshida, S. and
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158 A.R. Reddy and A.S. Raghavendra
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188 K. Janardhan Reddy constitution
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190 K. Janardhan Reddy World nitrog
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192 K. Janardhan Reddy nitrogen def
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194 K. Janardhan Reddy endoplasmic
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196 K. Janardhan Reddy drought cond
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198 K. Janardhan Reddy Manganese-de
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200 K. Janardhan Reddy zinc deficie
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202 K. Janardhan Reddy Table 12 . E
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204 K. Janardhan Reddy Table 14. Ef
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206 K. Janardhan Reddy Table 15. Th
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208 K. Janardhan Reddy Table 17. Co
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Page 252 and 253: Heavy Metal Stress 245 7. CONCLUSIO
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Page 256 and 257: Heavy Metal Stress 249 Kägi, J.H.R
Page 258 and 259: Heavy Metal Stress 251 Murphy, A.,
Page 260 and 261: Heavy Metal Stress 253 through xyle
Page 262 and 263: 255 CHAPTER 9 METABOLIC ENGINEERING
Page 264 and 265: Metabolic Engineering for Stress To
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312 A.K. Tyagi, S. Vij and N. Saini
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Table 3. Continued... Source Resour
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336 Index Auxins, 146 Avena sativa
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338 Expressed sequence tags (ESTs),
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340 Index Magnesium, 195 Mairiena s
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342 Index Processes less sensitive
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344 Index Sunflecks, 104 Sunflower,
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