Annual Plant Reviews, Phosphorus Metabolism in Plants
Herausgegeben von Plaxton, William; Lambers, Hans
Annual Plant Reviews, Phosphorus Metabolism in Plants
Herausgegeben von Plaxton, William; Lambers, Hans
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The development of phosphorus (P)-efficient crop varieties is urgently needed to reduce agriculture s current over-reliance on expensive, environmentally destructive, non-renewable and inefficient P-containing fertilizers. The sustainable management of P in agriculture necessitates an exploitation of P-adaptive traits that will enhance the P-acquisition and P-use efficiency of crop plants. Action in this area is crucial to ensure sufficient food production for the world's ever-expanding population, and the overall economic success of agriculture in the 21st century. This informative and…mehr
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The development of phosphorus (P)-efficient crop varieties is urgently needed to reduce agriculture s current over-reliance on expensive, environmentally destructive, non-renewable and inefficient P-containing fertilizers. The sustainable management of P in agriculture necessitates an exploitation of P-adaptive traits that will enhance the P-acquisition and P-use efficiency of crop plants. Action in this area is crucial to ensure sufficient food production for the world's ever-expanding population, and the overall economic success of agriculture in the 21st century.
This informative and up-to-date volume presents pivotal research directions that will facilitate the development of effective strategies for bioengineering P-efficient crop species. The 14 chapters reflect the expertise of an international team of leading authorities in the field, who review information from current literature, develop novel hypotheses, and outline key areas for future research. By evaluating aspects of vascular plant and green algal P uptake and metabolism, this book provides insights as to how plants sense, acquire, recycle, scavenge and use P, particularly under the naturally occurring condition of soluble inorganic phosphate deficiency that characterises the vast majority of unfertilised soils, worldwide. The reader is provided with a full appreciation of the diverse information concerning plant P-starvation responses, as well as the crucial role that plant-microbe interactions play in plant P acquisition.
Annual Plant Reviews, Volume 48: Phosphorus Metabolism in Plants is an important resource for plant geneticists, biochemists and physiologists, as well as horticultural and environmental research workers, advanced students of plant science and university lecturers in related disciplines. It is an essential addition to the shelves of university and research institute libraries and agricultural and ecological institutions teaching and researching plant science.
This informative and up-to-date volume presents pivotal research directions that will facilitate the development of effective strategies for bioengineering P-efficient crop species. The 14 chapters reflect the expertise of an international team of leading authorities in the field, who review information from current literature, develop novel hypotheses, and outline key areas for future research. By evaluating aspects of vascular plant and green algal P uptake and metabolism, this book provides insights as to how plants sense, acquire, recycle, scavenge and use P, particularly under the naturally occurring condition of soluble inorganic phosphate deficiency that characterises the vast majority of unfertilised soils, worldwide. The reader is provided with a full appreciation of the diverse information concerning plant P-starvation responses, as well as the crucial role that plant-microbe interactions play in plant P acquisition.
Annual Plant Reviews, Volume 48: Phosphorus Metabolism in Plants is an important resource for plant geneticists, biochemists and physiologists, as well as horticultural and environmental research workers, advanced students of plant science and university lecturers in related disciplines. It is an essential addition to the shelves of university and research institute libraries and agricultural and ecological institutions teaching and researching plant science.
Produktdetails
- Produktdetails
- Annual Plant Reviews Vol.48
- Verlag: Wiley & Sons
- 1. Auflage
- Seitenzahl: 480
- Erscheinungstermin: 15. Juni 2015
- Englisch
- Abmessung: 236mm x 157mm x 25mm
- Gewicht: 916g
- ISBN-13: 9781118958858
- ISBN-10: 1118958853
- Artikelnr.: 42053504
- Annual Plant Reviews Vol.48
- Verlag: Wiley & Sons
- 1. Auflage
- Seitenzahl: 480
- Erscheinungstermin: 15. Juni 2015
- Englisch
- Abmessung: 236mm x 157mm x 25mm
- Gewicht: 916g
- ISBN-13: 9781118958858
- ISBN-10: 1118958853
- Artikelnr.: 42053504
About the Editors William Plaxton is currently a Full Professor and Queen's Research Chair in the Department of Biology at Queen's University, Kingston, Canada. Hans Lambers is Professor of Plant Physiological Ecology in the School of Plant Biology at the University of Western Australia, Perth, Australia.
List of Contributors xvii Preface xxiii Section I Introduction 1
Phosphorus: Back to the Roots 3 Hans Lambers and William C. Plaxton 1.1
Introduction 3 1.2 Phosphorus or phosphorous? 4 1.3 Phosphorus on a
geological time scale 6 1.4 Phosphorus as an essential, but frequently
limiting, soil nutrient for plant productivity 7 1.5 Soil phosphorus pools
9 1.6 Soil phosphorus mobility 10 1.7 Factors determining rates of
phosphorus uptake by roots 11 1.8 Phosphorus-starvation responses: does
phosphorus homeostasis exist? 13 1.9 Concluding remarks 14 Acknowledgements
15 References 15 Section II P-Sensing, Transport, and Metabolism 2 Sensing,
Signalling, and Control of Phosphate Starvation in Plants: Molecular
Players and Applications 25 Wolf-Rüdiger Scheible and Monica Rojas-Triana
2.1 Introduction 25 2.2 The plant phosphate-starvation response 26 2.3
Sensing of phosphate and other macronutrient limitations in plants 29 2.3.1
Nutrient transporters as sensors/receptors 29 2.3.2 Local Pi sensing and
signalling at the root tip by PDR2/LPR1 31 2.3.3 Phosphite, a tool to
investigate P-sensing/signalling 31 2.4 Signalling of phosphate limitation
32 2.4.1 The role of phytohormones 33 2.4.2 Systemic signalling during
P-starvation 37 2.4.3 Transcriptional regulators involved in P-signalling
and affecting P-starvation responses 39 2.4.4 The role of microRNAs and
targeted protein degradation in P-signalling 41 2.4.5 Additional regulators
of P-signalling 43 2.5 Improving plant P-acquisition and -utilization
efficiency: approaches and targets 44 2.6 Concluding remarks 48 References
49 3 'Omics' Approaches Towards Understanding Plant Phosphorus Acquisition
and Use 65 Ping Lan, Wenfeng Li and Wolfgang Schmidt 3.1 Introduction 66
3.2 Towards a transcriptomics-derived 'phosphatome' 67 3.3 Pi
deficiency-induced alterations in the proteome 77 3.4 Core PSR proteins 80
3.5 Membrane lipid remodelling: insights from the transcriptome, the
proteome, and the lipidome 83 3.6 Genome-wide histone modifications in
Pi-deficient plants 86 3.7 Conclusions and outlook 89 3.8 Acknowledgements
90 References 90 4 The Role of Post-Translational Enzyme Modifications in
the Metabolic Adaptations of Phosphorus-Deprived Plants 99 William C.
Plaxton and Michael W. Shane 4.1 Introduction 100 4.2 In the beginning
there was protein phosphorylation 101 4.3 Monoubiquitination has emerged as
a crucial PTM that interacts with phosphorylation to control the function
of diverse proteins 104 4.4 Post-translational modification of plant
phosphoenolpyruvate carboxylase by phosphorylation versus
monoubiquitination 107 4.4.1 Activation of PEP carboxylase by in-vivo
phosphorylation appears to be a universal aspect of the plant P-starvation
response 107 4.4.2 PEP carboxylase monoubiquitination: an old dog learns
new tricks 109 4.4.3 Reciprocal control of PEP carboxylase by in-vivo
monoubiquitination and phosphorylation in developing proteoid roots of
P-deficient harsh hakea 111 4.5 Glycosylation is a sweet PTM of
glycoproteins 114 4.5.1 A pair of AtPAP26 glycoforms is upregulated and
secreted by P-deprived Arabidopsis 115 4.5.2 The AtPAP26-S2 glycoform
copurifies with, and appears to interact with, a curculin-like lectin 116
4.6 Concluding remarks 117 Acknowledgements 118 References 119 5 Phosphate
Transporters 125 Yves Poirier and Ji-Yul Jung 5.1 Introduction 125 5.2 The
PHT1 transporters 126 5.2.1 PHT1 structure, activity, and expression
patterns 126 5.3 Control of PHT1 activity 130 5.3.1 Control of PHT1
transcript levels 130 5.3.2 Post-transcriptional control of PHT1 133 5.4
PHO1 and phosphate export 136 5.4.1 PHO1 structure, activity, and
expression patterns 136 5.4.2 Transcriptional control of PHO1 expression
139 5.4.3 Post-transcriptional control of PHO1 139 5.5 Phosphate
transporters of organelles 140 5.5.1 Mitochondrial phosphate transporters
140 5.5.2 Plastidial phosphate transporters 141 5.5.3 The role of PHT2 in
plastid phosphate transport 143 5.5.4 The role of PHT4 in plastid phosphate
transport 143 5.6 Phosphate transporters of other organelles 145 5.6.1
Golgi phosphate transporters 145 5.6.2 Peroxisomal phosphate transporters
146 5.6.3 Vacuolar (tonoplast) phosphate transporters 146 5.7 Concluding
remarks 146 Acknowledgements 147 References 147 6 Molecular Components that
Drive Phosphorus-Remobilisation During Leaf Senescence 159 Aaron P. Smith,
Elena B. Fontenot, Sara Zahraeifard and Sandra Feuer DiTusa 6.1
Introduction 159 6.2 Transcriptomes of senescence and phosphate-deficiency
160 6.3 Major biochemical components that mediate P-remobilisation during
leaf senescence 162 6.3.1 Nucleases 163 6.3.2 Phosphatases 166 6.3.3
Lipid-remodelling enzymes 168 6.3.4 Pi transporters 169 6.4 Regulatory and
signalling components of senescing leaves 170 6.4.1 Transcription factors
170 6.4.2 The SPX superfamily 173 6.4.3 Ubiquitination components and
miRNAs 174 6.5 Role of hormones during leaf senescence 175 6.5.1 Ethylene
and strigolactones 175 6.5.2 Abscisic acid 176 6.5.3 Cytokinins 176 6.6
Concluding remarks 176 Acknowledgements 177 References 177 7 Interactions
Between Nitrogen and Phosphorus Metabolism 187 John A. Raven 7.1
Introduction 188 7.2 Roles of N and P in plants and the extent to which
compounds containing N or P can be substituted by compounds lacking N or P
188 7.3 Variability in the N:P ratio in plants and its metabolic and
ecological significance 195 7.3.1 Fixed N:P ratios: the role of compounds
containing both N and P 195 7.3.2 Protein:RNA ratio, organism N:P ratio,
the Growth Rate Hypothesis 197 7.3.3 Organism N and P concentration as a
function of external supply of N and P 200 7.3.4 Conclusions 201 7.4
Interactions in N and P acquisition and assimilation 201 7.4.1 Structures
involved in acquisition of N and P 202 7.4.2 Secretion of enzymes and
organic anions facilitates root N and P acquisition 204 7.5 Protein
synthesis and protein degradation during P-deprivation: significance for
N-P interaction 207 7.6 General conclusions 207 Acknowledgements 208
References 208 Section III P-deprivation Responses 8 Metabolomics of Plant
Phosphorus-Starvation Response 217 Chris Jones, Jean-Hugues Hatier, Mingshu
Cao, Karl Fraser and Susanne Rasmussen 8.1 Introduction 218 8.2 Metabolomic
approaches 219 8.3 Metabolomic analysis platforms 220 8.4 Data analysis 222
8.5 Metabolomics strategies directed at dissecting responses to P
starvation 223 8.6 Opportunities for metabolomics to contribute to the
development of P-efficient crops 229 8.7 Future prospects 230
Acknowledgements 231 References 231 9 Membrane Remodelling in
Phosphorus-Deficient Plants 237 Meike Siebers, Peter Dörmann and Georg
Hölzl 9.1 Introduction 237 9.2 Membrane lipid remodelling during phosphate
deprivation 238 9.3 Monogalactosyldiacylglycerol (MGDG) 242 9.4
Digalactosyldiacylglycerol (DGDG) 243 9.5 Sulfolipid (SQDG) and
glucuronosyldiacylglycerol (GlcADG) 247 9.6 Phospholipid degradation by
phospholipase D and phosphatidate phosphatase 248 9.7 Phospholipase C (PLC)
249 9.8 Acyl hydrolases 250 9.9 Lipid trafficking under phosphate
starvation 250 9.10 Glucosylceramide, sterol glucoside, and acylated sterol
glucoside 253 9.11 The role of auxin in remodelling of membrane lipid
composition 254 9.12 Improved Pi status by symbiosis with arbuscular
mycorrhizal fungi 255 9.13 Outlook 255 References 256 10 The Role of
Intracellular and Secreted Purple Acid Phosphatases in Plant Phosphorus
Scavenging and Recycling 265 Jiang Tian and Hong Liao 10.1 Introduction 266
10.2 Bioinformatics and structural analysis of plant PAPs 266 10.2.1 PAP
bioinformatics 266 10.2.2 Structural biochemistry of plant PAPs 269 10.3
Biochemical characterisation of plant PAPs 269 10.4 Diverse subcellular
localisation of plant PAPs 271 10.5 Transcriptional and
post-transcriptional regulation of PAP expression by P availability 275
10.5.1 Complex signal transduction pathways integrate nutritional P status
with PAP expression 276 10.5.2 Post-translational PAP modification 277 10.6
Functional analysis of PAPs involved in P mobilization and utilisation 278
10.7 Perspectives 281 Acknowledgements 282 References 282 11 Metabolic
Adaptations of the Non-Mycotrophic Proteaceae to Soils With Low Phosphorus
Availability 289 Hans Lambers, Peta L. Clode, Heidi-Jayne Hawkins, Etienne
Laliberté, Rafael S. Oliveira, Paul Reddell, Michael W. Shane, Mark Stitt
and Peter Weston 11.1 Introduction 290 11.2 Phosphorus nutrition of
Proteaceae, with a focus on south-western Australia 291 11.2.1 Phosphorus
acquisition by non-mycorrhizal roots: cluster roots 291 11.2.2 Proteaceae
species that do not produce cluster roots 298 11.2.3 Phosphorus toxicity
299 11.2.4 High rates of photosynthesis despite low leaf P concentrations
300 11.2.5 Leaf longevity 307 11.2.6 Delayed greening 308 11.2.7 Efficient
and proficient P remobilisation from senescing organs 310 11.2.8 Seed
Preserves 311 11.3 Comparison of species of Proteaceae in south-western
Australia with species elsewhere 312 11.3.1 The Cape Floristic Region in
South Africa 312 11.3.2 Eastern Australia 314 11.3.3 Southern South America
316 11.3.4 Brazil 317 11.4 Perspectives 318 Acknowledgements 323 References
323 12 Algae in a Phosphorus-Limited Landscape 337 Arthur R. Grossman and
Munevver Aksoy 12.1 Introduction 338 12.2 P-deprivation responses of green
algae and vascular plants 339 12.2.1 Phosphatases 342 12.2.2 Nucleases 346
12.2.3 Pi transport 348 12.2.4 Polyphosphates 350 12.2.5 Phospholipids 351
12.3 Control of P deprivation responses 353 12.3.1 PSR1-dependent gene
expression in P-starved algae 356 12.3.2 Low-phosphate bleaching mutants
358 12.4 Future prospects 359 Acknowledgements 360 References 360 Section
IV Significance of Plant-Microbe Interactions for P-Acquisition and
Metabolism 13 Impact of Roots, Microorganisms and Microfauna on the Fate of
Soil Phosphorus in the Rhizosphere 377 Philippe Hinsinger, Laetitia
Herrmann, Didier Lesueur, Agnès Robin, Jean Trap, Kittima Waithaisong and
Claude Plassard 13.1 Introduction 378 13.2 Spatial extension of the
rhizosphere 378 13.2.1 Root architecture and growth 379 13.2.2 Root hairs
and mycorrhizas 380 13.2.3 Root growth-promoting effect of rhizosphere
biota 381 13.3 Mobilisation of inorganic P in the rhizosphere 385 13.3.1
Effect of rhizosphere pH changes 385 13.3.2 Effect of exudation of
carboxylates 387 13.4 Mobilisation of organic P in the rhizosphere 389
13.4.1 Effects of phosphatases 390 13.4.2 Effects of phytases 391 13.5
Microbial P, microbial loop, and P recycling in the rhizosphere 393 13.5.1
Abiotic processes 393 13.5.2 Biotic processes 394 13.6 Conclusions and
future prospects 397 References 398 14 Mycorrhizal Associations and
Phosphorus Acquisition: From Cells to Ecosystems 409 Sally E. Smith, Ian C.
Anderson and F. Andrew Smith 14.1 Introduction 410 14.2 Arbuscular
mycorrhizas 413 14.2.1 Establishment of the symbiosis 413 14.2.2
Specialised AM interfaces in soil and roots are critical for P uptake 413
14.2.3 The AM pathway in plant P nutrition 416 14.2.4 The
'mutualism-parasitism' continuum 417 14.2.5 Some higher-scale issues in AM
symbiosis 418 14.2.6 Significance of AM symbioses in agriculture and
horticulture 419 14.3 Ectomycorrhizas 421 14.3.1 Establishment of the
symbiosis 421 14.3.2 Roles of ectomycorrhizas in plant P nutrition 422
14.3.3 ECM phosphate transporters 423 14.3.4 Solubilisation of inorganic
phosphates by ECM fungi 425 14.3.5 Mobilisation of organic-P sources by ECM
fungi 426 14.3.6 ECM symbioses and forest tree P nutrition: future
challenges 428 14.4 Conclusions 429 References 430 Index 441
Phosphorus: Back to the Roots 3 Hans Lambers and William C. Plaxton 1.1
Introduction 3 1.2 Phosphorus or phosphorous? 4 1.3 Phosphorus on a
geological time scale 6 1.4 Phosphorus as an essential, but frequently
limiting, soil nutrient for plant productivity 7 1.5 Soil phosphorus pools
9 1.6 Soil phosphorus mobility 10 1.7 Factors determining rates of
phosphorus uptake by roots 11 1.8 Phosphorus-starvation responses: does
phosphorus homeostasis exist? 13 1.9 Concluding remarks 14 Acknowledgements
15 References 15 Section II P-Sensing, Transport, and Metabolism 2 Sensing,
Signalling, and Control of Phosphate Starvation in Plants: Molecular
Players and Applications 25 Wolf-Rüdiger Scheible and Monica Rojas-Triana
2.1 Introduction 25 2.2 The plant phosphate-starvation response 26 2.3
Sensing of phosphate and other macronutrient limitations in plants 29 2.3.1
Nutrient transporters as sensors/receptors 29 2.3.2 Local Pi sensing and
signalling at the root tip by PDR2/LPR1 31 2.3.3 Phosphite, a tool to
investigate P-sensing/signalling 31 2.4 Signalling of phosphate limitation
32 2.4.1 The role of phytohormones 33 2.4.2 Systemic signalling during
P-starvation 37 2.4.3 Transcriptional regulators involved in P-signalling
and affecting P-starvation responses 39 2.4.4 The role of microRNAs and
targeted protein degradation in P-signalling 41 2.4.5 Additional regulators
of P-signalling 43 2.5 Improving plant P-acquisition and -utilization
efficiency: approaches and targets 44 2.6 Concluding remarks 48 References
49 3 'Omics' Approaches Towards Understanding Plant Phosphorus Acquisition
and Use 65 Ping Lan, Wenfeng Li and Wolfgang Schmidt 3.1 Introduction 66
3.2 Towards a transcriptomics-derived 'phosphatome' 67 3.3 Pi
deficiency-induced alterations in the proteome 77 3.4 Core PSR proteins 80
3.5 Membrane lipid remodelling: insights from the transcriptome, the
proteome, and the lipidome 83 3.6 Genome-wide histone modifications in
Pi-deficient plants 86 3.7 Conclusions and outlook 89 3.8 Acknowledgements
90 References 90 4 The Role of Post-Translational Enzyme Modifications in
the Metabolic Adaptations of Phosphorus-Deprived Plants 99 William C.
Plaxton and Michael W. Shane 4.1 Introduction 100 4.2 In the beginning
there was protein phosphorylation 101 4.3 Monoubiquitination has emerged as
a crucial PTM that interacts with phosphorylation to control the function
of diverse proteins 104 4.4 Post-translational modification of plant
phosphoenolpyruvate carboxylase by phosphorylation versus
monoubiquitination 107 4.4.1 Activation of PEP carboxylase by in-vivo
phosphorylation appears to be a universal aspect of the plant P-starvation
response 107 4.4.2 PEP carboxylase monoubiquitination: an old dog learns
new tricks 109 4.4.3 Reciprocal control of PEP carboxylase by in-vivo
monoubiquitination and phosphorylation in developing proteoid roots of
P-deficient harsh hakea 111 4.5 Glycosylation is a sweet PTM of
glycoproteins 114 4.5.1 A pair of AtPAP26 glycoforms is upregulated and
secreted by P-deprived Arabidopsis 115 4.5.2 The AtPAP26-S2 glycoform
copurifies with, and appears to interact with, a curculin-like lectin 116
4.6 Concluding remarks 117 Acknowledgements 118 References 119 5 Phosphate
Transporters 125 Yves Poirier and Ji-Yul Jung 5.1 Introduction 125 5.2 The
PHT1 transporters 126 5.2.1 PHT1 structure, activity, and expression
patterns 126 5.3 Control of PHT1 activity 130 5.3.1 Control of PHT1
transcript levels 130 5.3.2 Post-transcriptional control of PHT1 133 5.4
PHO1 and phosphate export 136 5.4.1 PHO1 structure, activity, and
expression patterns 136 5.4.2 Transcriptional control of PHO1 expression
139 5.4.3 Post-transcriptional control of PHO1 139 5.5 Phosphate
transporters of organelles 140 5.5.1 Mitochondrial phosphate transporters
140 5.5.2 Plastidial phosphate transporters 141 5.5.3 The role of PHT2 in
plastid phosphate transport 143 5.5.4 The role of PHT4 in plastid phosphate
transport 143 5.6 Phosphate transporters of other organelles 145 5.6.1
Golgi phosphate transporters 145 5.6.2 Peroxisomal phosphate transporters
146 5.6.3 Vacuolar (tonoplast) phosphate transporters 146 5.7 Concluding
remarks 146 Acknowledgements 147 References 147 6 Molecular Components that
Drive Phosphorus-Remobilisation During Leaf Senescence 159 Aaron P. Smith,
Elena B. Fontenot, Sara Zahraeifard and Sandra Feuer DiTusa 6.1
Introduction 159 6.2 Transcriptomes of senescence and phosphate-deficiency
160 6.3 Major biochemical components that mediate P-remobilisation during
leaf senescence 162 6.3.1 Nucleases 163 6.3.2 Phosphatases 166 6.3.3
Lipid-remodelling enzymes 168 6.3.4 Pi transporters 169 6.4 Regulatory and
signalling components of senescing leaves 170 6.4.1 Transcription factors
170 6.4.2 The SPX superfamily 173 6.4.3 Ubiquitination components and
miRNAs 174 6.5 Role of hormones during leaf senescence 175 6.5.1 Ethylene
and strigolactones 175 6.5.2 Abscisic acid 176 6.5.3 Cytokinins 176 6.6
Concluding remarks 176 Acknowledgements 177 References 177 7 Interactions
Between Nitrogen and Phosphorus Metabolism 187 John A. Raven 7.1
Introduction 188 7.2 Roles of N and P in plants and the extent to which
compounds containing N or P can be substituted by compounds lacking N or P
188 7.3 Variability in the N:P ratio in plants and its metabolic and
ecological significance 195 7.3.1 Fixed N:P ratios: the role of compounds
containing both N and P 195 7.3.2 Protein:RNA ratio, organism N:P ratio,
the Growth Rate Hypothesis 197 7.3.3 Organism N and P concentration as a
function of external supply of N and P 200 7.3.4 Conclusions 201 7.4
Interactions in N and P acquisition and assimilation 201 7.4.1 Structures
involved in acquisition of N and P 202 7.4.2 Secretion of enzymes and
organic anions facilitates root N and P acquisition 204 7.5 Protein
synthesis and protein degradation during P-deprivation: significance for
N-P interaction 207 7.6 General conclusions 207 Acknowledgements 208
References 208 Section III P-deprivation Responses 8 Metabolomics of Plant
Phosphorus-Starvation Response 217 Chris Jones, Jean-Hugues Hatier, Mingshu
Cao, Karl Fraser and Susanne Rasmussen 8.1 Introduction 218 8.2 Metabolomic
approaches 219 8.3 Metabolomic analysis platforms 220 8.4 Data analysis 222
8.5 Metabolomics strategies directed at dissecting responses to P
starvation 223 8.6 Opportunities for metabolomics to contribute to the
development of P-efficient crops 229 8.7 Future prospects 230
Acknowledgements 231 References 231 9 Membrane Remodelling in
Phosphorus-Deficient Plants 237 Meike Siebers, Peter Dörmann and Georg
Hölzl 9.1 Introduction 237 9.2 Membrane lipid remodelling during phosphate
deprivation 238 9.3 Monogalactosyldiacylglycerol (MGDG) 242 9.4
Digalactosyldiacylglycerol (DGDG) 243 9.5 Sulfolipid (SQDG) and
glucuronosyldiacylglycerol (GlcADG) 247 9.6 Phospholipid degradation by
phospholipase D and phosphatidate phosphatase 248 9.7 Phospholipase C (PLC)
249 9.8 Acyl hydrolases 250 9.9 Lipid trafficking under phosphate
starvation 250 9.10 Glucosylceramide, sterol glucoside, and acylated sterol
glucoside 253 9.11 The role of auxin in remodelling of membrane lipid
composition 254 9.12 Improved Pi status by symbiosis with arbuscular
mycorrhizal fungi 255 9.13 Outlook 255 References 256 10 The Role of
Intracellular and Secreted Purple Acid Phosphatases in Plant Phosphorus
Scavenging and Recycling 265 Jiang Tian and Hong Liao 10.1 Introduction 266
10.2 Bioinformatics and structural analysis of plant PAPs 266 10.2.1 PAP
bioinformatics 266 10.2.2 Structural biochemistry of plant PAPs 269 10.3
Biochemical characterisation of plant PAPs 269 10.4 Diverse subcellular
localisation of plant PAPs 271 10.5 Transcriptional and
post-transcriptional regulation of PAP expression by P availability 275
10.5.1 Complex signal transduction pathways integrate nutritional P status
with PAP expression 276 10.5.2 Post-translational PAP modification 277 10.6
Functional analysis of PAPs involved in P mobilization and utilisation 278
10.7 Perspectives 281 Acknowledgements 282 References 282 11 Metabolic
Adaptations of the Non-Mycotrophic Proteaceae to Soils With Low Phosphorus
Availability 289 Hans Lambers, Peta L. Clode, Heidi-Jayne Hawkins, Etienne
Laliberté, Rafael S. Oliveira, Paul Reddell, Michael W. Shane, Mark Stitt
and Peter Weston 11.1 Introduction 290 11.2 Phosphorus nutrition of
Proteaceae, with a focus on south-western Australia 291 11.2.1 Phosphorus
acquisition by non-mycorrhizal roots: cluster roots 291 11.2.2 Proteaceae
species that do not produce cluster roots 298 11.2.3 Phosphorus toxicity
299 11.2.4 High rates of photosynthesis despite low leaf P concentrations
300 11.2.5 Leaf longevity 307 11.2.6 Delayed greening 308 11.2.7 Efficient
and proficient P remobilisation from senescing organs 310 11.2.8 Seed
Preserves 311 11.3 Comparison of species of Proteaceae in south-western
Australia with species elsewhere 312 11.3.1 The Cape Floristic Region in
South Africa 312 11.3.2 Eastern Australia 314 11.3.3 Southern South America
316 11.3.4 Brazil 317 11.4 Perspectives 318 Acknowledgements 323 References
323 12 Algae in a Phosphorus-Limited Landscape 337 Arthur R. Grossman and
Munevver Aksoy 12.1 Introduction 338 12.2 P-deprivation responses of green
algae and vascular plants 339 12.2.1 Phosphatases 342 12.2.2 Nucleases 346
12.2.3 Pi transport 348 12.2.4 Polyphosphates 350 12.2.5 Phospholipids 351
12.3 Control of P deprivation responses 353 12.3.1 PSR1-dependent gene
expression in P-starved algae 356 12.3.2 Low-phosphate bleaching mutants
358 12.4 Future prospects 359 Acknowledgements 360 References 360 Section
IV Significance of Plant-Microbe Interactions for P-Acquisition and
Metabolism 13 Impact of Roots, Microorganisms and Microfauna on the Fate of
Soil Phosphorus in the Rhizosphere 377 Philippe Hinsinger, Laetitia
Herrmann, Didier Lesueur, Agnès Robin, Jean Trap, Kittima Waithaisong and
Claude Plassard 13.1 Introduction 378 13.2 Spatial extension of the
rhizosphere 378 13.2.1 Root architecture and growth 379 13.2.2 Root hairs
and mycorrhizas 380 13.2.3 Root growth-promoting effect of rhizosphere
biota 381 13.3 Mobilisation of inorganic P in the rhizosphere 385 13.3.1
Effect of rhizosphere pH changes 385 13.3.2 Effect of exudation of
carboxylates 387 13.4 Mobilisation of organic P in the rhizosphere 389
13.4.1 Effects of phosphatases 390 13.4.2 Effects of phytases 391 13.5
Microbial P, microbial loop, and P recycling in the rhizosphere 393 13.5.1
Abiotic processes 393 13.5.2 Biotic processes 394 13.6 Conclusions and
future prospects 397 References 398 14 Mycorrhizal Associations and
Phosphorus Acquisition: From Cells to Ecosystems 409 Sally E. Smith, Ian C.
Anderson and F. Andrew Smith 14.1 Introduction 410 14.2 Arbuscular
mycorrhizas 413 14.2.1 Establishment of the symbiosis 413 14.2.2
Specialised AM interfaces in soil and roots are critical for P uptake 413
14.2.3 The AM pathway in plant P nutrition 416 14.2.4 The
'mutualism-parasitism' continuum 417 14.2.5 Some higher-scale issues in AM
symbiosis 418 14.2.6 Significance of AM symbioses in agriculture and
horticulture 419 14.3 Ectomycorrhizas 421 14.3.1 Establishment of the
symbiosis 421 14.3.2 Roles of ectomycorrhizas in plant P nutrition 422
14.3.3 ECM phosphate transporters 423 14.3.4 Solubilisation of inorganic
phosphates by ECM fungi 425 14.3.5 Mobilisation of organic-P sources by ECM
fungi 426 14.3.6 ECM symbioses and forest tree P nutrition: future
challenges 428 14.4 Conclusions 429 References 430 Index 441
List of Contributors xvii Preface xxiii Section I Introduction 1
Phosphorus: Back to the Roots 3 Hans Lambers and William C. Plaxton 1.1
Introduction 3 1.2 Phosphorus or phosphorous? 4 1.3 Phosphorus on a
geological time scale 6 1.4 Phosphorus as an essential, but frequently
limiting, soil nutrient for plant productivity 7 1.5 Soil phosphorus pools
9 1.6 Soil phosphorus mobility 10 1.7 Factors determining rates of
phosphorus uptake by roots 11 1.8 Phosphorus-starvation responses: does
phosphorus homeostasis exist? 13 1.9 Concluding remarks 14 Acknowledgements
15 References 15 Section II P-Sensing, Transport, and Metabolism 2 Sensing,
Signalling, and Control of Phosphate Starvation in Plants: Molecular
Players and Applications 25 Wolf-Rüdiger Scheible and Monica Rojas-Triana
2.1 Introduction 25 2.2 The plant phosphate-starvation response 26 2.3
Sensing of phosphate and other macronutrient limitations in plants 29 2.3.1
Nutrient transporters as sensors/receptors 29 2.3.2 Local Pi sensing and
signalling at the root tip by PDR2/LPR1 31 2.3.3 Phosphite, a tool to
investigate P-sensing/signalling 31 2.4 Signalling of phosphate limitation
32 2.4.1 The role of phytohormones 33 2.4.2 Systemic signalling during
P-starvation 37 2.4.3 Transcriptional regulators involved in P-signalling
and affecting P-starvation responses 39 2.4.4 The role of microRNAs and
targeted protein degradation in P-signalling 41 2.4.5 Additional regulators
of P-signalling 43 2.5 Improving plant P-acquisition and -utilization
efficiency: approaches and targets 44 2.6 Concluding remarks 48 References
49 3 'Omics' Approaches Towards Understanding Plant Phosphorus Acquisition
and Use 65 Ping Lan, Wenfeng Li and Wolfgang Schmidt 3.1 Introduction 66
3.2 Towards a transcriptomics-derived 'phosphatome' 67 3.3 Pi
deficiency-induced alterations in the proteome 77 3.4 Core PSR proteins 80
3.5 Membrane lipid remodelling: insights from the transcriptome, the
proteome, and the lipidome 83 3.6 Genome-wide histone modifications in
Pi-deficient plants 86 3.7 Conclusions and outlook 89 3.8 Acknowledgements
90 References 90 4 The Role of Post-Translational Enzyme Modifications in
the Metabolic Adaptations of Phosphorus-Deprived Plants 99 William C.
Plaxton and Michael W. Shane 4.1 Introduction 100 4.2 In the beginning
there was protein phosphorylation 101 4.3 Monoubiquitination has emerged as
a crucial PTM that interacts with phosphorylation to control the function
of diverse proteins 104 4.4 Post-translational modification of plant
phosphoenolpyruvate carboxylase by phosphorylation versus
monoubiquitination 107 4.4.1 Activation of PEP carboxylase by in-vivo
phosphorylation appears to be a universal aspect of the plant P-starvation
response 107 4.4.2 PEP carboxylase monoubiquitination: an old dog learns
new tricks 109 4.4.3 Reciprocal control of PEP carboxylase by in-vivo
monoubiquitination and phosphorylation in developing proteoid roots of
P-deficient harsh hakea 111 4.5 Glycosylation is a sweet PTM of
glycoproteins 114 4.5.1 A pair of AtPAP26 glycoforms is upregulated and
secreted by P-deprived Arabidopsis 115 4.5.2 The AtPAP26-S2 glycoform
copurifies with, and appears to interact with, a curculin-like lectin 116
4.6 Concluding remarks 117 Acknowledgements 118 References 119 5 Phosphate
Transporters 125 Yves Poirier and Ji-Yul Jung 5.1 Introduction 125 5.2 The
PHT1 transporters 126 5.2.1 PHT1 structure, activity, and expression
patterns 126 5.3 Control of PHT1 activity 130 5.3.1 Control of PHT1
transcript levels 130 5.3.2 Post-transcriptional control of PHT1 133 5.4
PHO1 and phosphate export 136 5.4.1 PHO1 structure, activity, and
expression patterns 136 5.4.2 Transcriptional control of PHO1 expression
139 5.4.3 Post-transcriptional control of PHO1 139 5.5 Phosphate
transporters of organelles 140 5.5.1 Mitochondrial phosphate transporters
140 5.5.2 Plastidial phosphate transporters 141 5.5.3 The role of PHT2 in
plastid phosphate transport 143 5.5.4 The role of PHT4 in plastid phosphate
transport 143 5.6 Phosphate transporters of other organelles 145 5.6.1
Golgi phosphate transporters 145 5.6.2 Peroxisomal phosphate transporters
146 5.6.3 Vacuolar (tonoplast) phosphate transporters 146 5.7 Concluding
remarks 146 Acknowledgements 147 References 147 6 Molecular Components that
Drive Phosphorus-Remobilisation During Leaf Senescence 159 Aaron P. Smith,
Elena B. Fontenot, Sara Zahraeifard and Sandra Feuer DiTusa 6.1
Introduction 159 6.2 Transcriptomes of senescence and phosphate-deficiency
160 6.3 Major biochemical components that mediate P-remobilisation during
leaf senescence 162 6.3.1 Nucleases 163 6.3.2 Phosphatases 166 6.3.3
Lipid-remodelling enzymes 168 6.3.4 Pi transporters 169 6.4 Regulatory and
signalling components of senescing leaves 170 6.4.1 Transcription factors
170 6.4.2 The SPX superfamily 173 6.4.3 Ubiquitination components and
miRNAs 174 6.5 Role of hormones during leaf senescence 175 6.5.1 Ethylene
and strigolactones 175 6.5.2 Abscisic acid 176 6.5.3 Cytokinins 176 6.6
Concluding remarks 176 Acknowledgements 177 References 177 7 Interactions
Between Nitrogen and Phosphorus Metabolism 187 John A. Raven 7.1
Introduction 188 7.2 Roles of N and P in plants and the extent to which
compounds containing N or P can be substituted by compounds lacking N or P
188 7.3 Variability in the N:P ratio in plants and its metabolic and
ecological significance 195 7.3.1 Fixed N:P ratios: the role of compounds
containing both N and P 195 7.3.2 Protein:RNA ratio, organism N:P ratio,
the Growth Rate Hypothesis 197 7.3.3 Organism N and P concentration as a
function of external supply of N and P 200 7.3.4 Conclusions 201 7.4
Interactions in N and P acquisition and assimilation 201 7.4.1 Structures
involved in acquisition of N and P 202 7.4.2 Secretion of enzymes and
organic anions facilitates root N and P acquisition 204 7.5 Protein
synthesis and protein degradation during P-deprivation: significance for
N-P interaction 207 7.6 General conclusions 207 Acknowledgements 208
References 208 Section III P-deprivation Responses 8 Metabolomics of Plant
Phosphorus-Starvation Response 217 Chris Jones, Jean-Hugues Hatier, Mingshu
Cao, Karl Fraser and Susanne Rasmussen 8.1 Introduction 218 8.2 Metabolomic
approaches 219 8.3 Metabolomic analysis platforms 220 8.4 Data analysis 222
8.5 Metabolomics strategies directed at dissecting responses to P
starvation 223 8.6 Opportunities for metabolomics to contribute to the
development of P-efficient crops 229 8.7 Future prospects 230
Acknowledgements 231 References 231 9 Membrane Remodelling in
Phosphorus-Deficient Plants 237 Meike Siebers, Peter Dörmann and Georg
Hölzl 9.1 Introduction 237 9.2 Membrane lipid remodelling during phosphate
deprivation 238 9.3 Monogalactosyldiacylglycerol (MGDG) 242 9.4
Digalactosyldiacylglycerol (DGDG) 243 9.5 Sulfolipid (SQDG) and
glucuronosyldiacylglycerol (GlcADG) 247 9.6 Phospholipid degradation by
phospholipase D and phosphatidate phosphatase 248 9.7 Phospholipase C (PLC)
249 9.8 Acyl hydrolases 250 9.9 Lipid trafficking under phosphate
starvation 250 9.10 Glucosylceramide, sterol glucoside, and acylated sterol
glucoside 253 9.11 The role of auxin in remodelling of membrane lipid
composition 254 9.12 Improved Pi status by symbiosis with arbuscular
mycorrhizal fungi 255 9.13 Outlook 255 References 256 10 The Role of
Intracellular and Secreted Purple Acid Phosphatases in Plant Phosphorus
Scavenging and Recycling 265 Jiang Tian and Hong Liao 10.1 Introduction 266
10.2 Bioinformatics and structural analysis of plant PAPs 266 10.2.1 PAP
bioinformatics 266 10.2.2 Structural biochemistry of plant PAPs 269 10.3
Biochemical characterisation of plant PAPs 269 10.4 Diverse subcellular
localisation of plant PAPs 271 10.5 Transcriptional and
post-transcriptional regulation of PAP expression by P availability 275
10.5.1 Complex signal transduction pathways integrate nutritional P status
with PAP expression 276 10.5.2 Post-translational PAP modification 277 10.6
Functional analysis of PAPs involved in P mobilization and utilisation 278
10.7 Perspectives 281 Acknowledgements 282 References 282 11 Metabolic
Adaptations of the Non-Mycotrophic Proteaceae to Soils With Low Phosphorus
Availability 289 Hans Lambers, Peta L. Clode, Heidi-Jayne Hawkins, Etienne
Laliberté, Rafael S. Oliveira, Paul Reddell, Michael W. Shane, Mark Stitt
and Peter Weston 11.1 Introduction 290 11.2 Phosphorus nutrition of
Proteaceae, with a focus on south-western Australia 291 11.2.1 Phosphorus
acquisition by non-mycorrhizal roots: cluster roots 291 11.2.2 Proteaceae
species that do not produce cluster roots 298 11.2.3 Phosphorus toxicity
299 11.2.4 High rates of photosynthesis despite low leaf P concentrations
300 11.2.5 Leaf longevity 307 11.2.6 Delayed greening 308 11.2.7 Efficient
and proficient P remobilisation from senescing organs 310 11.2.8 Seed
Preserves 311 11.3 Comparison of species of Proteaceae in south-western
Australia with species elsewhere 312 11.3.1 The Cape Floristic Region in
South Africa 312 11.3.2 Eastern Australia 314 11.3.3 Southern South America
316 11.3.4 Brazil 317 11.4 Perspectives 318 Acknowledgements 323 References
323 12 Algae in a Phosphorus-Limited Landscape 337 Arthur R. Grossman and
Munevver Aksoy 12.1 Introduction 338 12.2 P-deprivation responses of green
algae and vascular plants 339 12.2.1 Phosphatases 342 12.2.2 Nucleases 346
12.2.3 Pi transport 348 12.2.4 Polyphosphates 350 12.2.5 Phospholipids 351
12.3 Control of P deprivation responses 353 12.3.1 PSR1-dependent gene
expression in P-starved algae 356 12.3.2 Low-phosphate bleaching mutants
358 12.4 Future prospects 359 Acknowledgements 360 References 360 Section
IV Significance of Plant-Microbe Interactions for P-Acquisition and
Metabolism 13 Impact of Roots, Microorganisms and Microfauna on the Fate of
Soil Phosphorus in the Rhizosphere 377 Philippe Hinsinger, Laetitia
Herrmann, Didier Lesueur, Agnès Robin, Jean Trap, Kittima Waithaisong and
Claude Plassard 13.1 Introduction 378 13.2 Spatial extension of the
rhizosphere 378 13.2.1 Root architecture and growth 379 13.2.2 Root hairs
and mycorrhizas 380 13.2.3 Root growth-promoting effect of rhizosphere
biota 381 13.3 Mobilisation of inorganic P in the rhizosphere 385 13.3.1
Effect of rhizosphere pH changes 385 13.3.2 Effect of exudation of
carboxylates 387 13.4 Mobilisation of organic P in the rhizosphere 389
13.4.1 Effects of phosphatases 390 13.4.2 Effects of phytases 391 13.5
Microbial P, microbial loop, and P recycling in the rhizosphere 393 13.5.1
Abiotic processes 393 13.5.2 Biotic processes 394 13.6 Conclusions and
future prospects 397 References 398 14 Mycorrhizal Associations and
Phosphorus Acquisition: From Cells to Ecosystems 409 Sally E. Smith, Ian C.
Anderson and F. Andrew Smith 14.1 Introduction 410 14.2 Arbuscular
mycorrhizas 413 14.2.1 Establishment of the symbiosis 413 14.2.2
Specialised AM interfaces in soil and roots are critical for P uptake 413
14.2.3 The AM pathway in plant P nutrition 416 14.2.4 The
'mutualism-parasitism' continuum 417 14.2.5 Some higher-scale issues in AM
symbiosis 418 14.2.6 Significance of AM symbioses in agriculture and
horticulture 419 14.3 Ectomycorrhizas 421 14.3.1 Establishment of the
symbiosis 421 14.3.2 Roles of ectomycorrhizas in plant P nutrition 422
14.3.3 ECM phosphate transporters 423 14.3.4 Solubilisation of inorganic
phosphates by ECM fungi 425 14.3.5 Mobilisation of organic-P sources by ECM
fungi 426 14.3.6 ECM symbioses and forest tree P nutrition: future
challenges 428 14.4 Conclusions 429 References 430 Index 441
Phosphorus: Back to the Roots 3 Hans Lambers and William C. Plaxton 1.1
Introduction 3 1.2 Phosphorus or phosphorous? 4 1.3 Phosphorus on a
geological time scale 6 1.4 Phosphorus as an essential, but frequently
limiting, soil nutrient for plant productivity 7 1.5 Soil phosphorus pools
9 1.6 Soil phosphorus mobility 10 1.7 Factors determining rates of
phosphorus uptake by roots 11 1.8 Phosphorus-starvation responses: does
phosphorus homeostasis exist? 13 1.9 Concluding remarks 14 Acknowledgements
15 References 15 Section II P-Sensing, Transport, and Metabolism 2 Sensing,
Signalling, and Control of Phosphate Starvation in Plants: Molecular
Players and Applications 25 Wolf-Rüdiger Scheible and Monica Rojas-Triana
2.1 Introduction 25 2.2 The plant phosphate-starvation response 26 2.3
Sensing of phosphate and other macronutrient limitations in plants 29 2.3.1
Nutrient transporters as sensors/receptors 29 2.3.2 Local Pi sensing and
signalling at the root tip by PDR2/LPR1 31 2.3.3 Phosphite, a tool to
investigate P-sensing/signalling 31 2.4 Signalling of phosphate limitation
32 2.4.1 The role of phytohormones 33 2.4.2 Systemic signalling during
P-starvation 37 2.4.3 Transcriptional regulators involved in P-signalling
and affecting P-starvation responses 39 2.4.4 The role of microRNAs and
targeted protein degradation in P-signalling 41 2.4.5 Additional regulators
of P-signalling 43 2.5 Improving plant P-acquisition and -utilization
efficiency: approaches and targets 44 2.6 Concluding remarks 48 References
49 3 'Omics' Approaches Towards Understanding Plant Phosphorus Acquisition
and Use 65 Ping Lan, Wenfeng Li and Wolfgang Schmidt 3.1 Introduction 66
3.2 Towards a transcriptomics-derived 'phosphatome' 67 3.3 Pi
deficiency-induced alterations in the proteome 77 3.4 Core PSR proteins 80
3.5 Membrane lipid remodelling: insights from the transcriptome, the
proteome, and the lipidome 83 3.6 Genome-wide histone modifications in
Pi-deficient plants 86 3.7 Conclusions and outlook 89 3.8 Acknowledgements
90 References 90 4 The Role of Post-Translational Enzyme Modifications in
the Metabolic Adaptations of Phosphorus-Deprived Plants 99 William C.
Plaxton and Michael W. Shane 4.1 Introduction 100 4.2 In the beginning
there was protein phosphorylation 101 4.3 Monoubiquitination has emerged as
a crucial PTM that interacts with phosphorylation to control the function
of diverse proteins 104 4.4 Post-translational modification of plant
phosphoenolpyruvate carboxylase by phosphorylation versus
monoubiquitination 107 4.4.1 Activation of PEP carboxylase by in-vivo
phosphorylation appears to be a universal aspect of the plant P-starvation
response 107 4.4.2 PEP carboxylase monoubiquitination: an old dog learns
new tricks 109 4.4.3 Reciprocal control of PEP carboxylase by in-vivo
monoubiquitination and phosphorylation in developing proteoid roots of
P-deficient harsh hakea 111 4.5 Glycosylation is a sweet PTM of
glycoproteins 114 4.5.1 A pair of AtPAP26 glycoforms is upregulated and
secreted by P-deprived Arabidopsis 115 4.5.2 The AtPAP26-S2 glycoform
copurifies with, and appears to interact with, a curculin-like lectin 116
4.6 Concluding remarks 117 Acknowledgements 118 References 119 5 Phosphate
Transporters 125 Yves Poirier and Ji-Yul Jung 5.1 Introduction 125 5.2 The
PHT1 transporters 126 5.2.1 PHT1 structure, activity, and expression
patterns 126 5.3 Control of PHT1 activity 130 5.3.1 Control of PHT1
transcript levels 130 5.3.2 Post-transcriptional control of PHT1 133 5.4
PHO1 and phosphate export 136 5.4.1 PHO1 structure, activity, and
expression patterns 136 5.4.2 Transcriptional control of PHO1 expression
139 5.4.3 Post-transcriptional control of PHO1 139 5.5 Phosphate
transporters of organelles 140 5.5.1 Mitochondrial phosphate transporters
140 5.5.2 Plastidial phosphate transporters 141 5.5.3 The role of PHT2 in
plastid phosphate transport 143 5.5.4 The role of PHT4 in plastid phosphate
transport 143 5.6 Phosphate transporters of other organelles 145 5.6.1
Golgi phosphate transporters 145 5.6.2 Peroxisomal phosphate transporters
146 5.6.3 Vacuolar (tonoplast) phosphate transporters 146 5.7 Concluding
remarks 146 Acknowledgements 147 References 147 6 Molecular Components that
Drive Phosphorus-Remobilisation During Leaf Senescence 159 Aaron P. Smith,
Elena B. Fontenot, Sara Zahraeifard and Sandra Feuer DiTusa 6.1
Introduction 159 6.2 Transcriptomes of senescence and phosphate-deficiency
160 6.3 Major biochemical components that mediate P-remobilisation during
leaf senescence 162 6.3.1 Nucleases 163 6.3.2 Phosphatases 166 6.3.3
Lipid-remodelling enzymes 168 6.3.4 Pi transporters 169 6.4 Regulatory and
signalling components of senescing leaves 170 6.4.1 Transcription factors
170 6.4.2 The SPX superfamily 173 6.4.3 Ubiquitination components and
miRNAs 174 6.5 Role of hormones during leaf senescence 175 6.5.1 Ethylene
and strigolactones 175 6.5.2 Abscisic acid 176 6.5.3 Cytokinins 176 6.6
Concluding remarks 176 Acknowledgements 177 References 177 7 Interactions
Between Nitrogen and Phosphorus Metabolism 187 John A. Raven 7.1
Introduction 188 7.2 Roles of N and P in plants and the extent to which
compounds containing N or P can be substituted by compounds lacking N or P
188 7.3 Variability in the N:P ratio in plants and its metabolic and
ecological significance 195 7.3.1 Fixed N:P ratios: the role of compounds
containing both N and P 195 7.3.2 Protein:RNA ratio, organism N:P ratio,
the Growth Rate Hypothesis 197 7.3.3 Organism N and P concentration as a
function of external supply of N and P 200 7.3.4 Conclusions 201 7.4
Interactions in N and P acquisition and assimilation 201 7.4.1 Structures
involved in acquisition of N and P 202 7.4.2 Secretion of enzymes and
organic anions facilitates root N and P acquisition 204 7.5 Protein
synthesis and protein degradation during P-deprivation: significance for
N-P interaction 207 7.6 General conclusions 207 Acknowledgements 208
References 208 Section III P-deprivation Responses 8 Metabolomics of Plant
Phosphorus-Starvation Response 217 Chris Jones, Jean-Hugues Hatier, Mingshu
Cao, Karl Fraser and Susanne Rasmussen 8.1 Introduction 218 8.2 Metabolomic
approaches 219 8.3 Metabolomic analysis platforms 220 8.4 Data analysis 222
8.5 Metabolomics strategies directed at dissecting responses to P
starvation 223 8.6 Opportunities for metabolomics to contribute to the
development of P-efficient crops 229 8.7 Future prospects 230
Acknowledgements 231 References 231 9 Membrane Remodelling in
Phosphorus-Deficient Plants 237 Meike Siebers, Peter Dörmann and Georg
Hölzl 9.1 Introduction 237 9.2 Membrane lipid remodelling during phosphate
deprivation 238 9.3 Monogalactosyldiacylglycerol (MGDG) 242 9.4
Digalactosyldiacylglycerol (DGDG) 243 9.5 Sulfolipid (SQDG) and
glucuronosyldiacylglycerol (GlcADG) 247 9.6 Phospholipid degradation by
phospholipase D and phosphatidate phosphatase 248 9.7 Phospholipase C (PLC)
249 9.8 Acyl hydrolases 250 9.9 Lipid trafficking under phosphate
starvation 250 9.10 Glucosylceramide, sterol glucoside, and acylated sterol
glucoside 253 9.11 The role of auxin in remodelling of membrane lipid
composition 254 9.12 Improved Pi status by symbiosis with arbuscular
mycorrhizal fungi 255 9.13 Outlook 255 References 256 10 The Role of
Intracellular and Secreted Purple Acid Phosphatases in Plant Phosphorus
Scavenging and Recycling 265 Jiang Tian and Hong Liao 10.1 Introduction 266
10.2 Bioinformatics and structural analysis of plant PAPs 266 10.2.1 PAP
bioinformatics 266 10.2.2 Structural biochemistry of plant PAPs 269 10.3
Biochemical characterisation of plant PAPs 269 10.4 Diverse subcellular
localisation of plant PAPs 271 10.5 Transcriptional and
post-transcriptional regulation of PAP expression by P availability 275
10.5.1 Complex signal transduction pathways integrate nutritional P status
with PAP expression 276 10.5.2 Post-translational PAP modification 277 10.6
Functional analysis of PAPs involved in P mobilization and utilisation 278
10.7 Perspectives 281 Acknowledgements 282 References 282 11 Metabolic
Adaptations of the Non-Mycotrophic Proteaceae to Soils With Low Phosphorus
Availability 289 Hans Lambers, Peta L. Clode, Heidi-Jayne Hawkins, Etienne
Laliberté, Rafael S. Oliveira, Paul Reddell, Michael W. Shane, Mark Stitt
and Peter Weston 11.1 Introduction 290 11.2 Phosphorus nutrition of
Proteaceae, with a focus on south-western Australia 291 11.2.1 Phosphorus
acquisition by non-mycorrhizal roots: cluster roots 291 11.2.2 Proteaceae
species that do not produce cluster roots 298 11.2.3 Phosphorus toxicity
299 11.2.4 High rates of photosynthesis despite low leaf P concentrations
300 11.2.5 Leaf longevity 307 11.2.6 Delayed greening 308 11.2.7 Efficient
and proficient P remobilisation from senescing organs 310 11.2.8 Seed
Preserves 311 11.3 Comparison of species of Proteaceae in south-western
Australia with species elsewhere 312 11.3.1 The Cape Floristic Region in
South Africa 312 11.3.2 Eastern Australia 314 11.3.3 Southern South America
316 11.3.4 Brazil 317 11.4 Perspectives 318 Acknowledgements 323 References
323 12 Algae in a Phosphorus-Limited Landscape 337 Arthur R. Grossman and
Munevver Aksoy 12.1 Introduction 338 12.2 P-deprivation responses of green
algae and vascular plants 339 12.2.1 Phosphatases 342 12.2.2 Nucleases 346
12.2.3 Pi transport 348 12.2.4 Polyphosphates 350 12.2.5 Phospholipids 351
12.3 Control of P deprivation responses 353 12.3.1 PSR1-dependent gene
expression in P-starved algae 356 12.3.2 Low-phosphate bleaching mutants
358 12.4 Future prospects 359 Acknowledgements 360 References 360 Section
IV Significance of Plant-Microbe Interactions for P-Acquisition and
Metabolism 13 Impact of Roots, Microorganisms and Microfauna on the Fate of
Soil Phosphorus in the Rhizosphere 377 Philippe Hinsinger, Laetitia
Herrmann, Didier Lesueur, Agnès Robin, Jean Trap, Kittima Waithaisong and
Claude Plassard 13.1 Introduction 378 13.2 Spatial extension of the
rhizosphere 378 13.2.1 Root architecture and growth 379 13.2.2 Root hairs
and mycorrhizas 380 13.2.3 Root growth-promoting effect of rhizosphere
biota 381 13.3 Mobilisation of inorganic P in the rhizosphere 385 13.3.1
Effect of rhizosphere pH changes 385 13.3.2 Effect of exudation of
carboxylates 387 13.4 Mobilisation of organic P in the rhizosphere 389
13.4.1 Effects of phosphatases 390 13.4.2 Effects of phytases 391 13.5
Microbial P, microbial loop, and P recycling in the rhizosphere 393 13.5.1
Abiotic processes 393 13.5.2 Biotic processes 394 13.6 Conclusions and
future prospects 397 References 398 14 Mycorrhizal Associations and
Phosphorus Acquisition: From Cells to Ecosystems 409 Sally E. Smith, Ian C.
Anderson and F. Andrew Smith 14.1 Introduction 410 14.2 Arbuscular
mycorrhizas 413 14.2.1 Establishment of the symbiosis 413 14.2.2
Specialised AM interfaces in soil and roots are critical for P uptake 413
14.2.3 The AM pathway in plant P nutrition 416 14.2.4 The
'mutualism-parasitism' continuum 417 14.2.5 Some higher-scale issues in AM
symbiosis 418 14.2.6 Significance of AM symbioses in agriculture and
horticulture 419 14.3 Ectomycorrhizas 421 14.3.1 Establishment of the
symbiosis 421 14.3.2 Roles of ectomycorrhizas in plant P nutrition 422
14.3.3 ECM phosphate transporters 423 14.3.4 Solubilisation of inorganic
phosphates by ECM fungi 425 14.3.5 Mobilisation of organic-P sources by ECM
fungi 426 14.3.6 ECM symbioses and forest tree P nutrition: future
challenges 428 14.4 Conclusions 429 References 430 Index 441