Verfügbarkeit: Plant biochemistry

Plant biochemistry:

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Bibliographische Detailangaben
Hauptverfasser:	Bowsher, Caroline (VerfasserIn), Steer, Martin (VerfasserIn), Tobin, Alyson (VerfasserIn)
Format:	Buch
Sprache:	English
Veröffentlicht:	London [u.a.] Garland Science 2008
Schlagworte:	Pflanzen Biochemie
Online-Zugang:	Inhaltsverzeichnis
Beschreibung:	XVI, 446 S. Ill., graph. Darst.
ISBN:	9780815341215 0815341210

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Datensatz im Suchindex

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adam_text	Contents Preface Abbreviations vii xv 1 Introduction to Plant Biochemistry 1 2 Approaches to Understanding Metabolic Pathways 5 What we need to understand a metabolic pathway 5 Chromatography 7 Electrophoresis 11 The use of isotopes 14 Current research techniques use a range of molecular biology approaches 16 Unique aspects of plant metabolism and their impact on metabolic flux 26 Metabolic control analysis theory 27 Coarse and fine metabolic control 29 Compartmentation: keeping competitive reactions apart 33 Understanding plant metabolism in the individual cell 33 The isolation of organelles 34 Summary 36 Further Reading 36 3 Plant Cell Structure 39 Cell structure is defined by membranes 40 The plasma membrane 46 Vacuoles and the tonoplast membrane 48 The endomembrane system 49 Cell walls serve to limit osmotic swelling of the enclosed protoplast 54 The nucleus contains the cell s chromatin within a highly specialized structure, the nuclear envelope 56 Mitochondria are ubiquitous organelles, which are the site of cellular respiration 57 Peroxisomes house vital biochemical pathways for many plant cell processes 58 Plastids 59 Summary 63 Further Reading 64 4 Light Reactions of Photosynthesis 65 Bacteria evolved the basic photochemical pathways found in plants today 65 Chlorophyll captures light energy and converts it to a flow of electrons 71 Carotenoids extend the spectral range of light that can be utilized in photosynthesis 74 Photosystem II splits water to form protons and oxygen, and reduces plastoquinone 75 The Q cycle uses plastoquinol to reduce plastocyanin and transport protons into the lumen 78 Photosystem I takes electrons from plastocyanin and reduces ferredoxin, which is used to make NADPH and other reduced compounds 80 ATP synthase utilizes the proton motive force to generate ATP 83 Cyclic photophosphorylation generates ATP independently of water oxidation and NADPH formation 85 Regulation of electron flow pathways in response to fluctuating light levels 86 Scavenging and removal of superoxides, peroxides, and other radicals by dismutases and antioxidants 87 Mechanisms for safely returning the levels of trapped high energy states to the ground state 88 Nonphotochemical quenching and the xanthophyll cycle 89 Summary 90 Further Reading 91 Plant Biochemistry 5 Photosynthetîc Carbon Assimilation 93 Photosynthetic carbon assimilation produces most of the biomass on Earth 93 Carbon dioxide enters the leaf through stornata but water is also lost in the process 94 Carbon dioxide is converted to carbohydrates using energy derived from sunlight 94 The Calvin cycle is used by all photosynthetic eukaryotes to convert carbon dioxide to carbohydrate 96 Discovery of the Calvin cycle 96 There are three phases to the Calvin cycle 97 Calvin cycle intermediates may be used to make other photosynthetic products 108 The Calvin cycle is autocatalytic and produces more substrate than it consumes 108 Calvin cycle activity and regulation 109 Rubisco is a highly regulated enzyme 111 Rubisco oxygenase: the starting point for the photorespiratory pathway 113 The photorespiratory pathway: enzymes in the chloroplast, peroxisome, and mitochondria 113 The isolation and analysis of mutants and the photorespiratory pathway 117 Photorespiration may provide essential amino acids and protect against environmental stress 117 Photorespiration and the loss of photosynthetically fixed carbon 118 Photorespiration uses ATP and reductant 119 Decreasing global carbon dioxide concentrations caused a rapid evolution of C4 photosynthesis 119 C4 photosynthesis concentrates carbon dioxide at the active site of Rubisco 120 Spadai separation of the two carboxylases in C4 leaves 120 Stages of C4 photosynthesis and variations to the basic pathway 122 C3-C4 intermediate species may represent an evolutionary stage between C3 and C4 plants 126 The C4 pathway can exist in single cells of some species 128 Some of the C4 pathway enzymes are light-regulated 129 Crassulacean acid metabolism as a feature of desert plants 130 Temporal separation of the carboxylases in CAM 130 Crassulacean acid metabolism as a flexible pathway 130 Phosphoeno/pyruvate carboxylase in crassulacean acid metabolism plants is regulated by protein phosphorylation 133 Crassulacean acid metabolism is thought to have evolved independently on several occasions 133 C3, C4, and CAM photosynthetic pathways: advantages and disadvantages 134 C3, C4, and CAM plants differ in their facility to discriminate between different isotopes of carbon 138 Summary 140 Further Reading 141 6 Respiration 143 Overview of respiration 143 The main components of plant respiration 144 Plants need energy and precursors for subsequent biosynthesis 144 Glycolysis is the major pathway that fuels respiration 145 Hexose sugars enter into glycolysis and are converted into fructose 1,6-bisphosphate 148 Fructose 1,6-bisphosphate is converted to pyravate 148 Alternative reactions provide flexibility to plant glycolysis 149 Plant glycolysis is regulated by a bottom-up process 151 Glycolysis supplies energy and reducing power for biosynthetic reactions 151 The availability of oxygen determines the fate ofpyruvate 151 The oxidative pentose phosphate pathway is an alternative catabolic route for glucose metabolism 153 The irreversible oxidative decarboxylation of glucose 6-phosphate generates NADPH 153 The second stage of the oxidative pentose phosphate pathway returns any excess pentose phosphates to glycolysis 153 All or part of the OPPP is duplicated in the plastids and cytosol 155 The tricarboxylic acid cycle is located in the mitochondria 155 Pyruvate oxidation marks the link between glycolysis and the tricarboxylic acid cycle 155 The product ofpyruvate oxidation, acetyl CoA, enters the tricarboxylic acid cycle via the citrate synthase reaction 164 Substrates for the tricarboxylic acid cycle are derived mainly from carbohydrates 167 The tricarboxylic acid cycle serves a biosynthetic function in plants 167 Contents xi Anaplerotic reactions are needed to enable intermediates to be withdrawn from the tricarboxylic acid cycle 169 The tricarboxylic acid cycle is regulated at several steps 170 Recent research into a thioredoxin/NADPH redox system for regulating tricarboxylic acid cycle enzymes and other mitochondria! proteins 172 The mitochondria! electron transport chain oxidizes reducing equivalents produced in respiratory substrate oxidation and produces ATP 172 Main protein complexes of the electron transport chain 173 Plant mitochondria possess additional respiratory proteins that provide a branched electron transport chain 175 Plant mitochondria contain four additional NAD(P)H dehydrogenases 176 The physiological function of the alternative NAD(P)H dehydrogenases remains the subject of some speculation 176 Plant mitochondria contain an alternative oxidase that transfers electrons from QH2 to oxygen and provides a bypass of the cytochrome oxidase branch 177 The alternative oxidase is a dimer of two identical polypeptides with a nonheme iron center 178 Alternative oxidase isoforms in plants are encoded by discrete gene families 178 Alternative oxidase activity is regulated by 2-oxo acids and by reduction and oxidation 180 The alternative oxidase adds flexibility to the operation of the mitochondrial electron transport chain 181 The alternative oxidase may prevent the formation of damaging reactive oxygen species within the mitochondria 182 Alternative oxidase appears to play a role in the response of plants to environmental stresses 182 Alternative oxidase and NADH oxidation 183 Plant mitochondria and uncoupling proteins 183 ATP synthesis in plant mitochondria is coupled to the proton electrochemical gradient that forms during electron transport 183 ATP synthase uses the proton motive force to generate ATP 184 Mitochondrial respiration interacts with photosynthesis and photorespiration in the light 187 Emerging research area into supercomplexes and metabolons 191 Summary 191 Further Reading 192 7 Synthesis and Mobilization of Storage and Structural Carbohydrates 195 Role of carbohydrate metabolism in higher plants 195 Sucrose is the major form of carbohydrate transported from source to sink tissue Sucrose phosphate synthase is an important control point in the sucrose biosynthetic pathway in plants Sensing, signaling, and regulation of carbon metabolism by fructose 2,6-bisphosphate Fructose 2,6-bisphosphate enables the cell to regulate the operation of multiple pathways of plant carbohydrate metabolism Fructose 2,6-bisphosphate as a regulatory link between the chloroplast and the cytosol Sucrose breakdown occurs via sucrose synthase and invertase Starch is the principal storage carbohydrate in plants 197 198 200 200 204 205 209 Starch synthesis occurs in plastids of both source and sink tissues 209 Starch formation occurs in water-insoluble starch granules in the plastids 213 The composition and structure of starch affects the properties and functions of starches 215 Starch degradation is different in different plant organs 216 The nature and regulation of starch degradation is poorly understood 216 Transitory starch is remobilized initially by a starch modifying process that takes place at the granule surface during the dark period 218 The regulation of starch degradation is unclear 219 Fractans are probably the most abundant storage carbohydrates in plants after starch and sucrose 220 A model has been proposed for the biosynthesis of the different fractan molecules found in plants 220 Fructan-accumulating plants are abundant in temperate climate zones with seasonal drought or frost 222 Trehalose biosynthesis is not just limited to resurrection plants 222 Trehalose biosynthesis in higher plants and its role in the regulation of carbon metabolism 223 Plant cell wall polysaccharides 224 Synthesis of cell wall sugars and polysaccharides 225 Cellulose 225 Matrix components consist of branched polysaccharides 228 xii Plant Biochemistry Expansins and extensins, proteins that play both enzymatic and structural roles in cell expansion 234 Lignin 234 Summary 235 Further Reading 235 8 Nitrogen and Sulfur Metabolism 237 Nitrogen and sulfur must be assimilated in the plant 237 Apart from oxygen, carbon, and hydrogen, nitrogen is the most abundant element in plants 238 Nitrogen fixation: some plants obtain nitrogen from the atmosphere via a symbiotic association with bacteria 239 Symbiotic nitrogen fixation involves a complex interaction between host plant and microorganism 242 Nodule-forming bacteria (Rhizobiaceae) are composed of the three genera Rhizobium, Bradyrhizobium, and Azorhizobium 242 The nodule environment is generated by interaction between legume plant host and rhizobia 244 Nitrogen fixation is energy expensive, consuming up to 20% of total photosynthates generated 245 Mycorrhizae are associations between soil fungi and plant roots that can enhance the nitrogen nutrition of the plant 246 Most higher plants obtain nitrogen from the soil in the form of nitrate 248 In higher plants there are multiple nitrate carriers with distinct properties and regulation 249 Nitrate reducíase catalyzes the reduction of nitrate to nitrite in the cytosol of root and shoot cells 250 The production of nitrite is rigidly controlled by the expression, catalytic activity, and degradation of NR 251 Nitrite reducíase, localized in the plastids, catalyzes the reduction of nitrite to ammonium 253 Plant cells have the capacity for the transport of ammonium ions 255 Ammonium is assimilated into amino acids 258 Ammonium originates from both primary and secondary sources 258 Ammonium is assimilated by glutaminę synthetase and glutamate synthase, which combine together in the glutaminę synthetase/ glutamate synthase cycle 259 GS is an octameric protein with two isoforms, localized in the cytosol and plastid 259 The GS genes and proteins show discrete cellular localization and different responses to light and nutrients 260 Glutaminę synthetase activity is regulated by metabolites and effectors, and may be modified by phosphorylation and 14-3-3 binding 261 Further evidence of the functions of glutaminę synthetase isoenzymes has come from studies of mutants and transgenic plants 261 Higher plants contain two forms of GOGAT, one is ferredoxin-dependent and the other is NADH-dependent 263 Both Fd- and NADH-GOGAT are located in the plastid and exist as monomeric proteins in most species 263 The tissue and cellular localization of Fd- and NADH-GOGAT provides a clue to their function in higher plants 264 Further evidence of the function of Fd- and NADH-GOGAT has come from the analysis of mutants and transgenic plants 264 Sulfur is an essential macronutrient but it represents only 0.1 % of plant dry matter 265 Sulfate is relatively abundant in the environment and serves as a primary sulfur source for plants 266 The assimilation of sulfate 267 Adenosine ö -phosphosulfate reducíase is composed of two distinct domains 268 Sulfite reducíase is similar in structure to nitrire reducíase 269 Sulfation is an alternative minor assimilation pathway incorporating sulfate into organic compounds 269 Amino acids biosynthesis is essential for plant growth and development 270 Carbon flow is essential to maintain amino acid production 270 There are species differences in the form of nitrogen transported through the xylem 272 Aminotransferase reactions are central to amino acid metabolism by disiribuiing nitrogen from glutamaíe ío oíher amino acids 273 Asparagine, aspartate, and alaninę biosynthesis 275 Glycine and serine biosynthesis 276 The aspartate family of amino acids: lysine, threonine, isoleucine, and methionine 276 The branched chain amino acids valine and leucine 279 Sulfur-containing amino acids cysteine and methionine 280 Contents xiii Glutaminę, arginine, and proline biosynthesis The biosynthesis of the aromatic amino acids: phenylalanine, tyrosine, and tryptophan Histidine biosynthesis Large amounts of nitrogen can be present in nonprotein amino acids Plant storage proteins: why do plants store proteins and what sort of proteins do they store? Vicilins and legumins are the main storage proteins in many dicotyledonous plants Prolamins are major storage proteins in cereals and grasses 2S albumins are important but minor components of seed proteins Where are seed proteins synthesized and how do they reach their storage compartment? Protein stores are degraded and mobilized during seed germination Vegetative organs store proteins, which are very different from seed proteins Despite their diversity, storage proteins share common characteristics Summary Further Reading 9 Lipid Biosynthesis Overview of lipids Fatty acid biosynthesis occurs through the sequential addition of two carbon units The condensation of nine two-carbon units is necessary for the assembly of an 18C fatty acid For the assembly of an 18C fatty acid from acetyl CoA using type II fatty acid synthase, 48 reactions are necessary and at least 12 different proteins involved Acyl-ACP utilization in the plastid Regulation of fatty acid formation Source of NADPH and ATP to support fatty acid biosynthesis Glycerolipids are formed from the incorporation of fatty acids to the glycerol backbone Phosphatidic acid, produced in the plastids or endoplasmic reticulum, is a central intermediate in glycerolipid biosynthesis Lipids function in signaling and defense The products of the oxidation of lipids and the resulting metabolites are collectively known as oxylipins 282 A waxy cuticle coats all land plants 322 Role of suberin as a hydrophobic layer 324 284 285 Storage lipids are primarily a storage form of carbon and chemical energy 325 Release of fatty acids from acyl lipids 328 285 The breakdown of fatty acids occurs via oxidation at the β carbon and subsequent removal of two carbon units 329 286 Summary 333 Further Reading 334 288 290 292 292 296 297 299 299 300 303 303 307 307 312 314 314 315 315 316 318 320 10 Alkaloids 335 Plants produce a vast array of chemicals that deter or attract other organisms 335 Alkaloids, a chemically diverse group that al contain nitrogen along with a number of carbon rings 336 Functions of alkaloids in plants and animals 336 The challenges and complexity of alkaloid biosynthetic pathways 336 Amino acids as precursors in the biosynthesis of alkaloids 338 Terpenoid indole alkaloids are made from tryptamine and the terpenoid secologanin 338 Isoquinoline alkaloids are produced from tyrosine and include many valuable drugs such as morphine and codeine 344 Tropane alkaloids and nicotine are found mainly in the Solanaceae 349 Pyrollizidine alkaloids are found in four main families 354 Purine alkaloids as popular stimulants in beverages, and as poisons and feeding deterrents against herbivores 355 The diversity of alkaloids has arisen through evolution driven by herbivore pressure 356 Summary 360 Further Reading 361 11 Phenolics 363 Plant phenolic compounds are a diverse group with a common aromatic ring structure and a range of biological functions 363 The simple phenolics 364 The more complex phenolics include the flavonoids, which have a characteristic three-membered А, В, С ring structure 367 Lignin is a complex polymer formed mainly from monolignol units 369 xiv Plant Biochemistry The tannins are phenolic polymers that form complexes with proteins 370 Most plant phenolics are synthesized from phenylpropanoids 370 The shikimic acid pathway provides the aromatic amino acid, phenylalanine, from which the phenylpropanoids are all derived 371 The core phenylpropanoid pathway provides the basic phenylpropanoid units that are used to make most of the phenolic compounds in plants 375 Flavonoids are produced from chalcones, formed from the condensation of p-coumaroyl CoA and malonyl CoA 379 Simple phenolics from the basic phenylpropanoid pathway are used in the biosynthesis of the hydrolyzable tannins 391 lignin is formed from monolignol subunits in a complex series of reactions that are still being unraveled 392 Summary 397 Further Reading 398 12 Terpenoids Terpenoids are a diverse group of essential oils that are formed from the fusion of five-carbon isoprene units Terpenoids serve a wide range of biological functions 399 399 402 411 The biosynthesis of terpenoids Subcellular compartmentation is important in the regulation of terpenoid biosynthesis 426 Summary 428 Further Reading 428 Index The colour plate section appears between pages 352 and 353. 431
adam_txt	Contents Preface Abbreviations vii xv 1 Introduction to Plant Biochemistry 1 2 Approaches to Understanding Metabolic Pathways 5 What we need to understand a metabolic pathway 5 Chromatography 7 Electrophoresis 11 The use of isotopes 14 Current research techniques use a range of molecular biology approaches 16 Unique aspects of plant metabolism and their impact on metabolic flux 26 Metabolic control analysis theory 27 Coarse and fine metabolic control 29 Compartmentation: keeping competitive reactions apart 33 Understanding plant metabolism in the individual cell 33 The isolation of organelles 34 Summary 36 Further Reading 36 3 Plant Cell Structure 39 Cell structure is defined by membranes 40 The plasma membrane 46 Vacuoles and the tonoplast membrane 48 The endomembrane system 49 Cell walls serve to limit osmotic swelling of the enclosed protoplast 54 The nucleus contains the cell's chromatin within a highly specialized structure, the nuclear envelope 56 Mitochondria are ubiquitous organelles, which are the site of cellular respiration 57 Peroxisomes house vital biochemical pathways for many plant cell processes 58 Plastids 59 Summary 63 Further Reading 64 4 Light Reactions of Photosynthesis 65 Bacteria evolved the basic photochemical pathways found in plants today 65 Chlorophyll captures light energy and converts it to a flow of electrons 71 Carotenoids extend the spectral range of light that can be utilized in photosynthesis 74 Photosystem II splits water to form protons and oxygen, and reduces plastoquinone 75 The Q cycle uses plastoquinol to reduce plastocyanin and transport protons into the lumen 78 Photosystem I takes electrons from plastocyanin and reduces ferredoxin, which is used to make NADPH and other reduced compounds 80 ATP synthase utilizes the proton motive force to generate ATP 83 Cyclic photophosphorylation generates ATP independently of water oxidation and NADPH formation 85 Regulation of electron flow pathways in response to fluctuating light levels 86 Scavenging and removal of superoxides, peroxides, and other radicals by dismutases and antioxidants 87 Mechanisms for safely returning the levels of trapped high energy states to the ground state 88 Nonphotochemical quenching and the xanthophyll cycle 89 Summary 90 Further Reading 91 Plant Biochemistry 5 Photosynthetîc Carbon Assimilation 93 Photosynthetic carbon assimilation produces most of the biomass on Earth 93 Carbon dioxide enters the leaf through stornata but water is also lost in the process 94 Carbon dioxide is converted to carbohydrates using energy derived from sunlight 94 The Calvin cycle is used by all photosynthetic eukaryotes to convert carbon dioxide to carbohydrate 96 Discovery of the Calvin cycle 96 There are three phases to the Calvin cycle 97 Calvin cycle intermediates may be used to make other photosynthetic products 108 The Calvin cycle is autocatalytic and produces more substrate than it consumes 108 Calvin cycle activity and regulation 109 Rubisco is a highly regulated enzyme 111 Rubisco oxygenase: the starting point for the photorespiratory pathway 113 The photorespiratory pathway: enzymes in the chloroplast, peroxisome, and mitochondria 113 The isolation and analysis of mutants and the photorespiratory pathway 117 Photorespiration may provide essential amino acids and protect against environmental stress 117 Photorespiration and the loss of photosynthetically fixed carbon 118 Photorespiration uses ATP and reductant 119 Decreasing global carbon dioxide concentrations caused a rapid evolution of C4 photosynthesis 119 C4 photosynthesis concentrates carbon dioxide at the active site of Rubisco 120 Spadai separation of the two carboxylases in C4 leaves 120 Stages of C4 photosynthesis and variations to the basic pathway 122 C3-C4 intermediate species may represent an evolutionary stage between C3 and C4 plants 126 The C4 pathway can exist in single cells of some species 128 Some of the C4 pathway enzymes are light-regulated 129 Crassulacean acid metabolism as a feature of desert plants 130 Temporal separation of the carboxylases in CAM 130 Crassulacean acid metabolism as a flexible pathway 130 Phosphoeno/pyruvate carboxylase in crassulacean acid metabolism plants is regulated by protein phosphorylation 133 Crassulacean acid metabolism is thought to have evolved independently on several occasions 133 C3, C4, and CAM photosynthetic pathways: advantages and disadvantages 134 C3, C4, and CAM plants differ in their facility to discriminate between different isotopes of carbon 138 Summary 140 Further Reading 141 6 Respiration 143 Overview of respiration 143 The main components of plant respiration 144 Plants need energy and precursors for subsequent biosynthesis 144 Glycolysis is the major pathway that fuels respiration 145 Hexose sugars enter into glycolysis and are converted into fructose 1,6-bisphosphate 148 Fructose 1,6-bisphosphate is converted to pyravate 148 Alternative reactions provide flexibility to plant glycolysis 149 Plant glycolysis is regulated by a bottom-up process 151 Glycolysis supplies energy and reducing power for biosynthetic reactions 151 The availability of oxygen determines the fate ofpyruvate 151 The oxidative pentose phosphate pathway is an alternative catabolic route for glucose metabolism 153 The irreversible oxidative decarboxylation of glucose 6-phosphate generates NADPH 153 The second stage of the oxidative pentose phosphate pathway returns any excess pentose phosphates to glycolysis 153 All or part of the OPPP is duplicated in the plastids and cytosol 155 The tricarboxylic acid cycle is located in the mitochondria 155 Pyruvate oxidation marks the link between glycolysis and the tricarboxylic acid cycle 155 The product ofpyruvate oxidation, acetyl CoA, enters the tricarboxylic acid cycle via the citrate synthase reaction 164 Substrates for the tricarboxylic acid cycle are derived mainly from carbohydrates 167 The tricarboxylic acid cycle serves a biosynthetic function in plants 167 Contents xi Anaplerotic reactions are needed to enable intermediates to be withdrawn from the tricarboxylic acid cycle 169 The tricarboxylic acid cycle is regulated at several steps 170 Recent research into a thioredoxin/NADPH redox system for regulating tricarboxylic acid cycle enzymes and other mitochondria! proteins 172 The mitochondria! electron transport chain oxidizes reducing equivalents produced in respiratory substrate oxidation and produces ATP 172 Main protein complexes of the electron transport chain 173 Plant mitochondria possess additional respiratory proteins that provide a branched electron transport chain 175 Plant mitochondria contain four additional NAD(P)H dehydrogenases 176 The physiological function of the alternative NAD(P)H dehydrogenases remains the subject of some speculation 176 Plant mitochondria contain an alternative oxidase that transfers electrons from QH2 to oxygen and provides a bypass of the cytochrome oxidase branch 177 The alternative oxidase is a dimer of two identical polypeptides with a nonheme iron center 178 Alternative oxidase isoforms in plants are encoded by discrete gene families 178 Alternative oxidase activity is regulated by 2-oxo acids and by reduction and oxidation 180 The alternative oxidase adds flexibility to the operation of the mitochondrial electron transport chain 181 The alternative oxidase may prevent the formation of damaging reactive oxygen species within the mitochondria 182 Alternative oxidase appears to play a role in the response of plants to environmental stresses 182 Alternative oxidase and NADH oxidation 183 Plant mitochondria and uncoupling proteins 183 ATP synthesis in plant mitochondria is coupled to the proton electrochemical gradient that forms during electron transport 183 ATP synthase uses the proton motive force to generate ATP 184 Mitochondrial respiration interacts with photosynthesis and photorespiration in the light 187 Emerging research area into supercomplexes and metabolons 191 Summary 191 Further Reading 192 7 Synthesis and Mobilization of Storage and Structural Carbohydrates 195 Role of carbohydrate metabolism in higher plants 195 Sucrose is the major form of carbohydrate transported from source to sink tissue Sucrose phosphate synthase is an important control point in the sucrose biosynthetic pathway in plants Sensing, signaling, and regulation of carbon metabolism by fructose 2,6-bisphosphate Fructose 2,6-bisphosphate enables the cell to regulate the operation of multiple pathways of plant carbohydrate metabolism Fructose 2,6-bisphosphate as a regulatory link between the chloroplast and the cytosol Sucrose breakdown occurs via sucrose synthase and invertase Starch is the principal storage carbohydrate in plants 197 198 200 200 204 205 209 Starch synthesis occurs in plastids of both source and sink tissues 209 Starch formation occurs in water-insoluble starch granules in the plastids 213 The composition and structure of starch affects the properties and functions of starches 215 Starch degradation is different in different plant organs 216 The nature and regulation of starch degradation is poorly understood 216 Transitory starch is remobilized initially by a starch modifying process that takes place at the granule surface during the dark period 218 The regulation of starch degradation is unclear 219 Fractans are probably the most abundant storage carbohydrates in plants after starch and sucrose 220 A model has been proposed for the biosynthesis of the different fractan molecules found in plants 220 Fructan-accumulating plants are abundant in temperate climate zones with seasonal drought or frost 222 Trehalose biosynthesis is not just limited to resurrection plants 222 Trehalose biosynthesis in higher plants and its role in the regulation of carbon metabolism 223 Plant cell wall polysaccharides 224 Synthesis of cell wall sugars and polysaccharides 225 Cellulose 225 Matrix components consist of branched polysaccharides 228 xii Plant Biochemistry Expansins and extensins, proteins that play both enzymatic and structural roles in cell expansion 234 Lignin 234 Summary 235 Further Reading 235 8 Nitrogen and Sulfur Metabolism 237 Nitrogen and sulfur must be assimilated in the plant 237 Apart from oxygen, carbon, and hydrogen, nitrogen is the most abundant element in plants 238 Nitrogen fixation: some plants obtain nitrogen from the atmosphere via a symbiotic association with bacteria 239 Symbiotic nitrogen fixation involves a complex interaction between host plant and microorganism 242 Nodule-forming bacteria (Rhizobiaceae) are composed of the three genera Rhizobium, Bradyrhizobium, and Azorhizobium 242 The nodule environment is generated by interaction between legume plant host and rhizobia 244 Nitrogen fixation is energy expensive, consuming up to 20% of total photosynthates generated 245 Mycorrhizae are associations between soil fungi and plant roots that can enhance the nitrogen nutrition of the plant 246 Most higher plants obtain nitrogen from the soil in the form of nitrate 248 In higher plants there are multiple nitrate carriers with distinct properties and regulation 249 Nitrate reducíase catalyzes the reduction of nitrate to nitrite in the cytosol of root and shoot cells 250 The production of nitrite is rigidly controlled by the expression, catalytic activity, and degradation of NR 251 Nitrite reducíase, localized in the plastids, catalyzes the reduction of nitrite to ammonium 253 Plant cells have the capacity for the transport of ammonium ions 255 Ammonium is assimilated into amino acids 258 Ammonium originates from both primary and secondary sources 258 Ammonium is assimilated by glutaminę synthetase and glutamate synthase, which combine together in the glutaminę synthetase/ glutamate synthase cycle 259 GS is an octameric protein with two isoforms, localized in the cytosol and plastid 259 The GS genes and proteins show discrete cellular localization and different responses to light and nutrients 260 Glutaminę synthetase activity is regulated by metabolites and effectors, and may be modified by phosphorylation and 14-3-3 binding 261 Further evidence of the functions of glutaminę synthetase isoenzymes has come from studies of mutants and transgenic plants 261 Higher plants contain two forms of GOGAT, one is ferredoxin-dependent and the other is NADH-dependent 263 Both Fd- and NADH-GOGAT are located in the plastid and exist as monomeric proteins in most species 263 The tissue and cellular localization of Fd- and NADH-GOGAT provides a clue to their function in higher plants 264 Further evidence of the function of Fd- and NADH-GOGAT has come from the analysis of mutants and transgenic plants 264 Sulfur is an essential macronutrient but it represents only 0.1 % of plant dry matter 265 Sulfate is relatively abundant in the environment and serves as a primary sulfur source for plants 266 The assimilation of sulfate 267 Adenosine ö'-phosphosulfate reducíase is composed of two distinct domains 268 Sulfite reducíase is similar in structure to nitrire reducíase 269 Sulfation is an alternative minor assimilation pathway incorporating sulfate into organic compounds 269 Amino acids biosynthesis is essential for plant growth and development 270 Carbon flow is essential to maintain amino acid production 270 There are species differences in the form of nitrogen transported through the xylem 272 Aminotransferase reactions are central to amino acid metabolism by disiribuiing nitrogen from glutamaíe ío oíher amino acids 273 Asparagine, aspartate, and alaninę biosynthesis 275 Glycine and serine biosynthesis 276 The aspartate family of amino acids: lysine, threonine, isoleucine, and methionine 276 The branched chain amino acids valine and leucine 279 Sulfur-containing amino acids cysteine and methionine 280 Contents xiii Glutaminę, arginine, and proline biosynthesis The biosynthesis of the aromatic amino acids: phenylalanine, tyrosine, and tryptophan Histidine biosynthesis Large amounts of nitrogen can be present in nonprotein amino acids Plant storage proteins: why do plants store proteins and what sort of proteins do they store? Vicilins and legumins are the main storage proteins in many dicotyledonous plants Prolamins are major storage proteins in cereals and grasses 2S albumins are important but minor components of seed proteins Where are seed proteins synthesized and how do they reach their storage compartment? Protein stores are degraded and mobilized during seed germination Vegetative organs store proteins, which are very different from seed proteins Despite their diversity, storage proteins share common characteristics Summary Further Reading 9 Lipid Biosynthesis Overview of lipids Fatty acid biosynthesis occurs through the sequential addition of two carbon units The condensation of nine two-carbon units is necessary for the assembly of an 18C fatty acid For the assembly of an 18C fatty acid from acetyl CoA using type II fatty acid synthase, 48 reactions are necessary and at least 12 different proteins involved Acyl-ACP utilization in the plastid Regulation of fatty acid formation Source of NADPH and ATP to support fatty acid biosynthesis Glycerolipids are formed from the incorporation of fatty acids to the glycerol backbone Phosphatidic acid, produced in the plastids or endoplasmic reticulum, is a central intermediate in glycerolipid biosynthesis Lipids function in signaling and defense The products of the oxidation of lipids and the resulting metabolites are collectively known as oxylipins 282 A waxy cuticle coats all land plants 322 Role of suberin as a hydrophobic layer 324 284 285 Storage lipids are primarily a storage form of carbon and chemical energy 325 Release of fatty acids from acyl lipids 328 285 The breakdown of fatty acids occurs via oxidation at the β carbon and subsequent removal of two carbon units 329 286 Summary 333 Further Reading 334 288 290 292 292 296 297 299 299 300 303 303 307 307 312 314 314 315 315 316 318 320 10 Alkaloids 335 Plants produce a vast array of chemicals that deter or attract other organisms 335 Alkaloids, a chemically diverse group that al contain nitrogen along with a number of carbon rings 336 Functions of alkaloids in plants and animals 336 The challenges and complexity of alkaloid biosynthetic pathways 336 Amino acids as precursors in the biosynthesis of alkaloids 338 Terpenoid indole alkaloids are made from tryptamine and the terpenoid secologanin 338 Isoquinoline alkaloids are produced from tyrosine and include many valuable drugs such as morphine and codeine 344 Tropane alkaloids and nicotine are found mainly in the Solanaceae 349 Pyrollizidine alkaloids are found in four main families 354 Purine alkaloids as popular stimulants in beverages, and as poisons and feeding deterrents against herbivores 355 The diversity of alkaloids has arisen through evolution driven by herbivore pressure 356 Summary 360 Further Reading 361 11 Phenolics 363 Plant phenolic compounds are a diverse group with a common aromatic ring structure and a range of biological functions 363 The simple phenolics 364 The more complex phenolics include the flavonoids, which have a characteristic three-membered А, В, С ring structure 367 Lignin is a complex polymer formed mainly from monolignol units 369 xiv Plant Biochemistry The tannins are phenolic polymers that form complexes with proteins 370 Most plant phenolics are synthesized from phenylpropanoids 370 The shikimic acid pathway provides the aromatic amino acid, phenylalanine, from which the phenylpropanoids are all derived 371 The core phenylpropanoid pathway provides the basic phenylpropanoid units that are used to make most of the phenolic compounds in plants 375 Flavonoids are produced from chalcones, formed from the condensation of p-coumaroyl CoA and malonyl CoA 379 Simple phenolics from the basic phenylpropanoid pathway are used in the biosynthesis of the hydrolyzable tannins 391 lignin is formed from monolignol subunits in a complex series of reactions that are still being unraveled 392 Summary 397 Further Reading 398 12 Terpenoids Terpenoids are a diverse group of essential oils that are formed from the fusion of five-carbon isoprene units Terpenoids serve a wide range of biological functions 399 399 402 411 The biosynthesis of terpenoids Subcellular compartmentation is important in the regulation of terpenoid biosynthesis 426 Summary 428 Further Reading 428 Index The colour plate section appears between pages 352 and 353. 431
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spelling	Bowsher, Caroline Verfasser aut Plant biochemistry Caroline Bowsher, Martin Steer, Alyson Tobin London [u.a.] Garland Science 2008 XVI, 446 S. Ill., graph. Darst. txt rdacontent n rdamedia nc rdacarrier Pflanzen (DE-588)4045539-7 gnd rswk-swf Biochemie (DE-588)4006777-4 gnd rswk-swf Pflanzen (DE-588)4045539-7 s Biochemie (DE-588)4006777-4 s DE-604 Steer, Martin Verfasser aut Tobin, Alyson Verfasser aut Digitalisierung UB Regensburg application/pdf http://bvbr.bib-bvb.de:8991/F?func=service&doc_library=BVB01&local_base=BVB01&doc_number=016242628&sequence=000002&line_number=0001&func_code=DB_RECORDS&service_type=MEDIA Inhaltsverzeichnis
spellingShingle	Bowsher, Caroline Steer, Martin Tobin, Alyson Plant biochemistry Pflanzen (DE-588)4045539-7 gnd Biochemie (DE-588)4006777-4 gnd
subject_GND	(DE-588)4045539-7 (DE-588)4006777-4
title	Plant biochemistry
title_auth	Plant biochemistry
title_exact_search	Plant biochemistry
title_exact_search_txtP	Plant biochemistry
title_full	Plant biochemistry Caroline Bowsher, Martin Steer, Alyson Tobin
title_fullStr	Plant biochemistry Caroline Bowsher, Martin Steer, Alyson Tobin
title_full_unstemmed	Plant biochemistry Caroline Bowsher, Martin Steer, Alyson Tobin
title_short	Plant biochemistry
title_sort	plant biochemistry
topic	Pflanzen (DE-588)4045539-7 gnd Biochemie (DE-588)4006777-4 gnd
topic_facet	Pflanzen Biochemie
url	http://bvbr.bib-bvb.de:8991/F?func=service&doc_library=BVB01&local_base=BVB01&doc_number=016242628&sequence=000002&line_number=0001&func_code=DB_RECORDS&service_type=MEDIA
work_keys_str_mv	AT bowshercaroline plantbiochemistry AT steermartin plantbiochemistry AT tobinalyson plantbiochemistry

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