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- ISBN: 9781405145381 | 1405145382
- Cover: Hardcover
- Copyright: 4/2/2007
Regulatory light signals are detected by an array of specialised, information-transducing photoreceptors, including the phytochrome, cryptochrome and phototropin families of chromoproteins. In recent years the application of genetic, biochemical and molecular studies has led to the identification and characterisation not only of the photoreceptors and their genes, but also of many of the components that act downstream of photoreceptor activation. It is evident that the photoreceptors operate through interactions with one another and with other signalling systems, so forming complex response networks. This volume provides the reader with state-of-the-art accounts of our current understanding of the major classes of higher plant regulatory photoreceptors and the signal transduction networks that comprise plant developmental photobiology. Consideration is also given to the ways in which our knowledge of plant photoreceptors and their signalling networks can be exploited to improve the quality and productivity of commercially-grown plants.
Professor Garry C. Whitelam, Head of Department, School of Biological Sciences, University of Leicester, UK
Dr Karen J. Halliday, School of Biological Sciences, The University of Edinburgh, UK
Dr Karen J. Halliday, School of Biological Sciences, The University of Edinburgh, UK
Contributors | p. xiii |
Preface | p. xv |
Photoreceptors | p. 1 |
Phytochromes | p. 3 |
Introduction | p. 3 |
Historical aspects | p. 3 |
Properties of phyA in vivo | p. 4 |
Properties in yeast cells | p. 4 |
In vivo properties of phytochromes | p. 5 |
In vivo spectroscopy | p. 5 |
Intracellular localisation of phytochromes | p. 7 |
Classical methods | p. 7 |
Spectroscopic methods | p. 7 |
Cell biological methods | p. 8 |
Immunocytochemical methods | p. 8 |
Novel methods | p. 8 |
Intracellular localisation of phyB in dark and light | p. 9 |
Intracellular localisation of phyA in dark and light | p. 11 |
Intracellular localisation of phyC, phyD and phyE in dark and light | p. 12 |
Phytochrome/PIF3 co-localisation and nuclear speckles | p. 12 |
Regulation of intracellular localisation of phytochromes | p. 13 |
Acknowledgements | p. 14 |
References | p. 15 |
Cryptochromes | p. 17 |
Introduction | p. 17 |
Cryptochrome genes and their evolution | p. 17 |
Cryptochrome domains, chromophores and structure | p. 20 |
Domain structure of the cryptochromes | p. 20 |
Cryptochrome chromophores | p. 21 |
Photolyase and cryptochrome structure | p. 22 |
Photolyase structure and reaction mechanism | p. 22 |
Cryptochrome structure | p. 23 |
Cryptochrome biochemistry and spectroscopy | p. 25 |
Phosphorylation | p. 25 |
Nucleotide-binding and kinase activity | p. 25 |
DNA-binding activity | p. 26 |
Electron transfer | p. 27 |
Expression and biological activity of cryptochromes | p. 28 |
Expression and light regulation of cryptochromes in planta | p. 28 |
Cellular localization | p. 31 |
Growth responses controlled by cryptochromes | p. 31 |
Regulation of gene expression through cryptochromes | p. 33 |
Cryptochrome signalling | p. 35 |
Dimerization and output domains | p. 36 |
Cryptochrome partners | p. 38 |
Interaction with COP1 | p. 39 |
Interaction with zeitlupe/ADAGIO 1 | p. 39 |
Interaction with phytochromes | p. 40 |
Further downstream components | p. 41 |
Summary | p. 43 |
Acknowledgements | p. 43 |
References | p. 43 |
Phototropins and other LOV-containing proteins | p. 49 |
Introduction | p. 49 |
Phototropins and their biological functions | p. 50 |
Physiological roles in higher plants | p. 50 |
Physiological roles in lower plants | p. 52 |
Phototropin structure, localization and activity | p. 53 |
Phototropin structure and localization | p. 53 |
Phototropin autophosphorylation | p. 55 |
Light sensing by the LOV domains | p. 57 |
LOV-domain photochemistry | p. 57 |
LOV-domain structure | p. 59 |
Functional roles of LOV1 and LOV2 | p. 60 |
Light-induced protein movements | p. 62 |
Phototropin signaling | p. 63 |
Phototropin-interacting proteins | p. 63 |
Downstream signaling targets | p. 65 |
Other LOV-containing proteins | p. 67 |
LOV-containing proteins in Arabidopsis | p. 67 |
LOV-containing proteins in fungi | p. 70 |
LOV-containing proteins in bacteria | p. 72 |
Conclusions and future perspectives | p. 73 |
Acknowledgements | p. 73 |
References | p. 74 |
Photoreceptor signal transduction | p. 79 |
Phytochrome-interacting factors | p. 81 |
Introduction | p. 81 |
Methodology | p. 81 |
Initial identification of PIFs | p. 81 |
Subsequent assay and characterization of the interaction | p. 82 |
Reverse genetic assessment of functional relevance to phy signalling | p. 83 |
Phy-interactors | p. 84 |
PIF3 | p. 84 |
Other bHLH transcription factors and the active phyB binding domain | p. 90 |
Nucleoside diphosphate kinase 2 | p. 93 |
Phytochrome kinase substrate 1 | p. 94 |
Type 5 protein phosphatase | p. 94 |
Protein phosphatase 2A | p. 95 |
Early flowering 3 | p. 95 |
Far-red elongated hypocotyl 1 | p. 96 |
Pre-selected interaction targets | p. 96 |
Arabidopsis response regulator 4 | p. 96 |
Zeitlupe | p. 97 |
Cryptochrome 1 and 2 | p. 97 |
AUX/IAA proteins | p. 98 |
COP1 | p. 98 |
Perspective | p. 99 |
Acknowledgements | p. 100 |
References | p. 101 |
Phosphorylation/dephosphorylation in photoreceptor signalling | p. 106 |
Introduction | p. 106 |
The photoreceptors autophosphorylate, but the classical activation loop is not involved | p. 107 |
Phosphatases in photoreceptor signalling | p. 108 |
Phytochromes | p. 109 |
Phosphorylation of phytochrome | p. 109 |
Phytochrome kinase substrate 1 | p. 110 |
Nucleoside diphosphate kinase 2 | p. 111 |
FYPP | p. 112 |
PAPP5 | p. 113 |
Cryptochromes | p. 114 |
Cryptochrome phosphorylation | p. 114 |
Phosphorylation of the C-terminal end is necessary for signal transduction | p. 115 |
Phototropins | p. 116 |
Is phosphorylation/dephosphorylation important for downstream events? | p. 118 |
HY5 | p. 118 |
Long hypocotyl in far-red light | p. 119 |
Circadian clock-associated and late elongated hypocotyl | p. 119 |
EID1 | p. 120 |
Aux/IAA | p. 120 |
PP7 | p. 121 |
Downstream of phototropin | p. 121 |
Conclusions | p. 122 |
References | p. 123 |
The role of ubiquitin/proteasome-mediated proteolysis in photoreceptor action | p. 128 |
Introduction | p. 128 |
Overview of the ubiquitin/proteasome system | p. 129 |
Ubiquitin conjugation and deconjugation pathways | p. 129 |
Diversity of E3 Ub ligases | p. 130 |
26S proteasome | p. 131 |
Role of COP/DET/FUS proteins in photoreceptor-mediated signal transduction and ubiquitin/proteasome-mediated proteolysis | p. 132 |
COP/DET/FUS proteins integrate divergent photoreceptor signaling pathways and downstream gene expression | p. 132 |
COP1 | p. 134 |
Light regulation of COP1 localization | p. 135 |
COP1 acts as an E3 Ub ligase | p. 136 |
Interaction between photoreceptors and COP1 | p. 137 |
CDD complex | p. 137 |
COP10 is an E2 Ub-conjugating enzyme variant | p. 137 |
COP10 forms a complex with DET1 and DDB1 | p. 138 |
COP9 signalosome | p. 139 |
Composition and structure of the COP9 signalosome | p. 139 |
Biochemical activities of the C0P9 signalosome | p. 140 |
Regulation of cullin-containing E3 Ub ligases by the C0P9 signalosome | p. 140 |
SPA protein family | p. 142 |
Other connection points between light signaling and selective proteolysis | p. 143 |
F-box proteins that are involved in light signaling | p. 143 |
Other light signaling pathway components that are targets of proteolysis | p. 144 |
Phytochrome A | p. 144 |
Cryptochrome 2 | p. 144 |
Phytochrome-interacting factor 3 and far-red elongated hypocotyl 1 | p. 145 |
Concluding remarks | p. 146 |
References | p. 147 |
UV-B perception and signal transduction | p. 155 |
Introduction | p. 155 |
UV-B in the environment | p. 156 |
Plant responses to UV-B | p. 157 |
UV-B perception and signal transduction | p. 159 |
Non-photomorphogenic UV-B signalling | p. 160 |
Damage/stress signalling | p. 160 |
Overlap with defence/wound signalling | p. 162 |
Photomorphogenic UV-B signalling | p. 163 |
UV-B perception | p. 164 |
Signal transduction | p. 166 |
Genetic approach | p. 167 |
Screens for UV-B signalling mutants | p. 167 |
UVR8 | p. 170 |
HY5 | p. 173 |
Other transcription factors involved in UV-B responses | p. 174 |
Concluding remarks | p. 175 |
Acknowledgements | p. 176 |
References | p. 176 |
Physiological responses | p. 183 |
Photocontrol of flowering | p. 185 |
Introduction | p. 185 |
Internal cues | p. 187 |
The autonomous pathway | p. 187 |
The gibberellin-dependent pathway | p. 189 |
External cues | p. 189 |
Photoperiodism | p. 189 |
Long days or short nights? | p. 190 |
The circadian clock | p. 191 |
The coincidence model | p. 196 |
Flowering time mutants of Arabidopsis | p. 197 |
Application to other species | p. 199 |
Site of perception of photoperiodic stimulus | p. 200 |
Shade avoidance | p. 201 |
Vernalisation | p. 203 |
Convergence of the flowering pathways | p. 203 |
Conclusion | p. 205 |
References | p. 205 |
Red:far-red ratio perception and shade avoidance | p. 211 |
Introduction | p. 211 |
Natural light environment | p. 211 |
Shade avoidance syndrome | p. 215 |
Phytochrome regulation of shade avoidance | p. 217 |
The role of phytochromes B, D and E in R:FR ratio signalling | p. 218 |
The role of phyA in R:FR ratio signalling | p. 220 |
The roles of other signals in shade avoidance | p. 221 |
PAR and B signals | p. 221 |
Hormone signals | p. 222 |
Signalling in shade avoidance | p. 224 |
ATHB-2 | p. 224 |
PLL1 | p. 224 |
R:FR ratio and flowering | p. 226 |
The adaptive value of shade avoidance | p. 227 |
Conclusions | p. 228 |
References | p. 229 |
Photoreceptor interactions with other signals | p. 235 |
Introduction | p. 235 |
Light-clock connections | p. 235 |
The clock | p. 235 |
Photoentrainment | p. 237 |
Light control of flowering time | p. 239 |
Light-hormone connections | p. 240 |
Light and auxin signal integration | p. 240 |
Light regulation of auxin biosynthesis and transport | p. 241 |
Signalling components shared by light and auxin | p. 242 |
Light regulation of the GH3 gene family | p. 242 |
Role of Aux/IAAs and proteolysis in light and auxin signalling | p. 243 |
The tropisms | p. 244 |
Light and auxin control of shoot phototropism | p. 245 |
Phytochrome and cryptochrome modification of shoot phototropism | p. 246 |
Root phototropism | p. 247 |
Gravitropism | p. 247 |
Light regulation of gravitropism | p. 248 |
Light and GA signal integration | p. 249 |
Phytochrome regulation of GA biosynthesis and homeostasis | p. 249 |
Light regulation of GA signalling | p. 250 |
The thermosensory pathways | p. 251 |
Summary | p. 255 |
References | p. 256 |
Applied aspects of photomorphogenesis | p. 265 |
Photoreceptor biotechnology | p. 267 |
Introduction and background | p. 267 |
Approaches to modification of photomorphogenic responses in crop plants | p. 268 |
Dwarfing plants using photoreceptors | p. 268 |
The shade-avoidance response | p. 269 |
Control of gene expression and shade avoidance | p. 270 |
Alteration of the timing of flowering | p. 271 |
Taxonomic differences and similarities in higher plants | p. 272 |
Modification of photomorphogenesis using genetic transformation - the state of the art | p. 273 |
Plants transgenic for phytochromes | p. 273 |
Modification of other photoreceptors | p. 276 |
Overexpression of signaling components | p. 278 |
Modification of photomorphogenesis by utilizing genetic diversity | p. 278 |
Natural variation in photomorphogenesis | p. 278 |
Photoreceptors and photomorphogenic genes as targets for selection in crops | p. 280 |
Photoreceptor biotechnology ex planta | p. 281 |
Using phytochrome to control gene expression | p. 281 |
Phytochromes as fluorescent probes | p. 282 |
Other potential uses of photoreceptors | p. 284 |
Future directions in photoreceptor biotechnology | p. 284 |
Acknowledgements | p. 285 |
References | p. 285 |
Light-quality manipulation by horticulture industry | p. 290 |
Introduction | p. 290 |
Regions of light spectrum important for plant growth and development | p. 290 |
Plant responses to quality of light | p. 292 |
Light manipulation by horticulture industry | p. 293 |
Electric light sources | p. 294 |
Spectral filters | p. 295 |
Greenhouse covers | p. 295 |
Photoselective nets | p. 300 |
Coloured plastic mulch | p. 305 |
Future prospects | p. 307 |
References | p. 307 |
Index | p. 313 |
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