Verfügbarkeit: Plant physiology

Plant physiology:

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Bibliographische Detailangaben
Hauptverfasser:	Taiz, Lincoln (VerfasserIn), Zeiger, Eduardo (VerfasserIn)
Format:	Buch
Sprache:	English
Veröffentlicht:	Sunderland, Mass. Sinauer 2006
Ausgabe:	4. ed.
Schlagworte:	Fisiologia vegetal Fysiologie Physiologie végétale Planten Plant physiology Entwicklungsphysiologie Pflanzen Pflanzenphysiologie Lehrbuch
Online-Zugang:	Inhaltsverzeichnis
Beschreibung:	XXVI, 764 S. zahlr. Ill., graph. Darst.
ISBN:	0878938567 9780878938568

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Datensatz im Suchindex

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adam_text	Table Preface v Authors and Contributors vu 1 Plant Cells Plant Life: Unifying Principles Overview of Plant Structure Plant cells are surrounded by rigid cell walls New cells are produced by dividing tissues called meristems Three major tissue systems make up the plant body The Plant Cell Biological membranes are phospholipid bilayers that contain proteins The nucleus contains most of the genetic material of the cell Protein synthesis involves transcription and translation The endoplasmic reticulum is a network of internal membranes Secretion of proteins from cells begins with the rough ER 11 Golgi stacks produce and distribute secretory products Proteins and polysaccharides destined for secretion are processed in the Golgi apparatus Two models for intra-Golgi transport have been proposed Specific coat proteins facilitate vesicle budding Vacuoles Mitochondria and chloroplasts are sites of energy conversion Mitochondria and chloroplasts are semiautonomous organelles Different plastid types are interconvertible Microbodies play specialized metabolic roles in leaves and seeds Oleosomes are lipid-storing organelles The Cytoskeleton Plant cells contain microtubules, intermediate filaments Microtubules and disassemble Microtubules function in mitosis and cytokinesis Motor proteins mediate cytoplasmic streaming and organelle movements Cell Cycle Regulation Each phase of the cell cycle has a specific set of biochemical and cellular activities The cell cycle is regulated by cyclin-dependent kinases Plasmodesmata There are two types of plasmodesmata: primary and secondary Plasmodesmata have a complex internal structure Macromolecular traffic through plasmodesmata is important for developmental signaling Summary 29 Ł XÜ UNITI Transport and 3 Water and Plant Cells Water in Plant Life The Structure and Properties of Water The polarity of water molecules gives rise to hydrogen bonds The polarity of water makes it an excellent solvent The thermal properties of water result from hydrogen bonding The cohesive and adhesive properties of water are due to hydrogen bonding Water has a high tensile strength Water Transport Processes Diffusion is the movement of molecules by random thermal agitation Diffusion is rapid over short distances but extremely slow over long distances Pressure-driven bulk flow drives long-distance water transport Osmosis is driven by a water potential gradient 4 Water Balance of Plants Water in the Soil A negative hydrostatic pressure in soil water lowers soil water potential Water moves through the soil by bulk flow Water Absorption by Roots Water moves in the root via the apoplast, symplast, and transmembrane Solute accumulation in the xylem can generate root pressure Water Transport through the Xylem The xylem consists of two types of tracheary elements Water movement through the xylem requires less pressure than movement through living cells What pressure difference is needed to lift water meters to a treetop? The cohesion-tension theory explains water transport in the xylem Xylem transport of water in trees faces physical challenges The chemical potential of water represents the free- energy status of water Three major factors contribute to cell water potential Water enters the cell along a water potential gradient Water can also leave the cell in response to a water potential gradient Small changes in plant cell volume cause large changes in Water transport rates depend on driving force and hydraulic conductivity Aquaporins facilitate the movement of water across cell membranes The water potential concept helps us evaluate the water status of a plant The components of water potential vary with growth conditions and location within the plant Summary Plants minimize the consequences of xylem cavitation Water Movement from the Leaf to the Atmosphere The driving force for water loss is the difference in water vapor concentration Water loss is also regulated by the pathway resistances Stomatal photosynthesis The cell walls of guard cells have specialized features An increase in guard cell stornata The transpiration ratio measures the relationship between water loss and carbon gain Overview: The Soil-Plant-Atmosphere Continuum Summary Table 5 Mineral Essential Nutrients, Deficiencies, and Plant Disorders Special techniques are used in nutritional studies Nutrient solutions can sustain rapid plant growth Mineral deficiencies disrupt plant metabolism and function Analysis of plant tissues reveals mineral deficiencies Treating Nutritional Deficiencies Crop yields can be improved by addition of fertilizers 83 Some mineral nutrients can be absorbed by leaves Soil, Roots, and Microbes Negatively charged soil particles affect the adsorption 6 Solute Transport Passive and Active Transport Transport of Ions across a Membrane Barrier Different diffusion rates for cations and anions produce diffusion potentials How does membrane potential relate to ion distribution? The Nernst equation distinguishes between active and passive transport Proton transport is a major determinant of the membrane potential Membrane Transport Processes Channel transporters enhance diffusion across membranes Carriers bind and transport specific substances Primary active transport requires energy Secondary active transport uses stored energy Kinetic analyses can elucidate transport mechanisms Membrane Transport Proteins 105 106 of mineral nutrients Soil root growth Excess minerals in the soil limit plant growth Plants develop extensive root systems Root systems differ in form but are based on common structures Different areas of the root absorb different mineral ions Mycorrhizal fungi facilitate nutrient uptake by roots 89 Nutrients move from the mycorrhizal fungi to the root cells Summary The genes for many transporters have been identified Transporters exist for diverse nitrogen-containing compounds Cation transporters are diverse Some Metals are transported by ZIP proteins Aquaporins may have novel functions The plasma membrane H+-ATPase has several functional domains The tonoplast H+-ATPase drives solute accumulation into H+-pyrophosphatases also pump protons at the tonoplast Ion Transport in Roots Solutes move through both apoplast and symplast Ions cross both symplast and apoplast Xylem parenchyma cells participate in xylem loading Summary Xiv UNITU Biochemistry and Metabolism 1 Photosynthesis: The Light Reactions Photosynthesis in Higher Plants General Concepts Light has characteristics of both a particle and a wave When molecules absorb or emit light, they change their electronic state Photosynthetic pigments absorb the light that powers photosynthesis Key Experiments in Understanding Photosynthesis Action spectra relate light absorption to photosynthetic activity Photosynthesis takes place in complexes containing light-harvesting antennas and photochemical reaction centers The chemical reaction of photosynthesis is driven by light Light drives the reduction of NADP and the formation of ATP Oxygen-evolving organisms have two photosystems that operate in series Organization of the Photosynthetic Apparatus The Thylakoids contain integral membrane proteins Photosystems I and II are spatially separated in the thylakoid membrane Anoxygenic photosynthetic bacteria have a single reaction center Organization of Light-Absorbing Antenna Systems The antenna funnels energy to the reaction center Many antenna complexes have a common structural motif Mechanisms of Electron Transport Electrons from chlorophyll travel through the carriers organized in the Z scheme Energy is captured when an excited chlorophyll reduces an electron acceptor molecule The reaction center chlorophylls of the two photosystems absorb at different wavelengths The photosystem II reaction center is a multisubunit pigment-protein complex Water is oxidized to oxygen by photosystem II Pheophytin and two photosystem II Electron flow through the cytochrome b6 transports protons Plastoquinone and plastocyanin carry electrons between photosystems II and I The photosystem I reaction center reduces NADP+ Cyclic electron flow generates ATP but no NADPH Some herbicides block photosynthetic electron flow Proton Transport and ATP Synthesis in the Chloroplast Repair and Regulation of the Photosynthetic Machinery Carotenoids serve as photoprotective agents Some xanthophylls also participate in energy dissipation The photosystem II reaction center is easily damaged Photosystem I is protected from active oxygen species Thylakoid stacking permits energy partitioning between the photosystems Genetics, Assembly, and Evolution of Photosynthetic Systems Chloroplast, cyanobacterial, been sequenced Chloroplast inheritance Many cytoplasm The biosynthesis and breakdown of chlorophyll are complex pathways Complex photosynthetic organisms have evolved from simpler forms Summary Table of Contents XV б Photosynthesis: Carbon Reactions The Calvin Cycle The Calvin cycle has three stages: carboxylation, reduction, and regeneration The carboxylation of ribulose-l,5-bisphosphate is catalyzed by the enzyme rubisco Operation of the Calvin cycle requires the regeneration of ribulose-l,5-bisphosphate The Calvin cycle regenerates its own biochemical components The Calvin cycle uses energy very efficiently Regulation of the Calvin Cycle Light regulates the Calvin cycle The activity of rubisco increases in the light The ferredoxin-thioredoxin system regulates the Calvin cycle Light-dependent ion movements regulate Calvin cycle enzymes The C2 Oxidative Photosynthetic Carbon Cycle Photosynthetic CO2 fixation and photorespiratory oxygénation Photorespiration electron transport system The biological function of investigation COj-Concentrating Mechanisms I. CO2 and HCO3- Pumps II. The C4 Carbon Cycle Malate C4 cycle Two different types of cells participate in the C4 cycle The C4 cycle concentrates CO2 in the bundle sheath cells The C4 cycle also concentrates CO2 in single cells The C4 cycle has higher energy demand than the Calvin cycle Light regulates the activity of key C4 enzymes In hot, dry climates, the C4 cycle reduces photorespiration III. Crassulacean Acid Metabolism (CAM) The during the day Some CAM plants change the pattern of CO2 uptake in response to environmental conditions Starch and Sucrose Chloroplast degraded at night Starch is synthesized in the Starch degradation requires phosphorylation of amylopectin Triose the pool of hexose phosphates in the cytosol Fructose-6-phosphate can be converted to fructose- 1,6-bisphosphate by two different enzymes Fructose-2,6-bisphosphate is an important regulatory compound The hexose phosphate pool is regulated by fructose-2,6- bisphosphate Sucrose is continuously synthesized in the cytosol Summary У Light, Leaves, and Photosynthesis Units in the Measurement of Light Leaf anatomy maximizes light absorption Plants compete for sunlight Leaf angle and leaf movement can control light absorption Plants acclimate and adapt to sun and shade Photosynthetic Responses to Light by the Intact Leaf Light-response curves reveal photosynthetic properties Leaves must dissipate excess light energy Absorption of too much light can lead to photoinhibition Photosynthetic Responses to Temperature Leaves must dissipate vast quantities of heat Photosynthesis is temperature sensitive Photosynthetic Responses to Carbon Dioxide Atmospheric CO2 concentration keeps rising CO2 diffusion to the photosynthesis Patterns of light absorption generate gradients of CO2 fixation CO2 imposes limitations on photosynthesis Crassulacean Acid Metabolism Carbon isotope ratio variations reveal different photosynthetic pathways How do we measure the carbon isotopes of plants? Why are there carbon isotope ratio variations in plants? Summary xvi 10 Translocation Pathways of Sugar is translocated in phloem sieve elements Mature sieve elements are living cells specialized for translocation Large pores in cell walls are the prominent feature of sieve elements Damaged sieve elements are sealed off Companion cells aid the highly specialized sieve elements Patterns of Source-to-sink pathways follow anatomic and developmental patterns Materials Translocated in the Phloem Phloem sap can be collected and analyzed Sugars are translocated in nonreducing form Rates of Movement The Pressure-Flow Model for Phloem Transport A pressure gradient drives pressure-flow model The predictions of mass flow have been confirmed Sieve plate pores are open channels There is no bidirectional transport in single sieve elements The energy requirement for transport through the phloem pathway is small Pressure gradients are sufficient to drive a mass flow of phloem sap Significant questions about the pressure-flow model still exist Phloem Loading Phloem loading can occur via the apoplast or symplast Sucrose uptake in the apoplastic pathway requires metabolic energy Phloem loading in the apoplastic pathway involves a sucrose-H 1 symporter Phloem loading is symplastic in plants with intermediary cells The polymer-trapping model explains symplastic loading The type of phloem loading is correlated with plant family and with climate Phloem Unloading and Sink-to-Source Transition Phloem unloading and short-distance transport can occur via symplastic or apoplastic pathways Transport into sink tissues requires metabolic energy The transition of a leaf from sink to source is gradual Photosynthate Distribution: Allocation and Partitioning Allocation includes storage, utilization, and transport Various sinks partition transport sugars Source leaves regulate allocation Sink tissues compete for available translocated photosynthate Sink strength depends on sink size and activity The source adjusts over the long term to changes in the source-to-sink ratio The Transport of Signaling Molecules Turgor and sink activities Signal molecules in the phloem regulate growth and development Summary 11 Respiration and Overview of Plant Respiration Glycolysis: A Cytosolic and Plastidic Process Glycolysis converts carbohydrates into pyruvate, producing NADH and ATP Plants have alternative glycolytic reactions In the absence of O,, fermentation regenerates the NAD+ needed for glycolysis Fermentation does not liberate all the energy available in each sugar molecule Plant glycolysis is controlled by its products The pentose phosphate pathway produces NADPH and biosynthetic intermediates The Citric Acid Cycle: A Mitochondrial Matrix Process Mitochondria are semiautonomous organelles Pyruvate enters the mitochondrion and is oxidized via the citric acid cycle The citric acid cycle of plants has unique features Mitochondrial Electron Transport and ATP Synthesis The electron transport chain catalyzes a flow of electrons from NADH to O2 Some electron transport enzymes are unique to plant mitochondria Table of ATP synthesis in the mitochondrion is coupled to electron transport Transporters exchange substrates and products Aerobic respiration yields about per molecule of sucrose Several subunits of respiratory complexes are encoded by the mitochondrial genome Plants have several mechanisms that lower the ATP yield Mitochondrial respiration is controlled by key metabolites Respiration is tightly coupled to other pathways Respiration in Intact Plants and Tissues Plants respire roughly half of the daily photosynthetic yield Respiration operates during photosynthesis Different tissues and organs respire at different rates Mitochondrial function is crucial during pollen development Environmental factors alter respiration rates Lipid Fats and oils store large amounts of energy Triacylglycerols are stored in oil bodies Polar glycerolipids are the main structural lipids in membranes Fatty acid biosynthesis consists of cycles of two-carbon addition Glycerolipids are synthesized in the plastids and the 282 Lipid Membrane lipids are precursors of important signaling compounds Storage lipids are converted into carbohydrates in germinating seeds Summary 12 Assimilation of Mineral Nutrients Nitrogen in the Environment Nitrogen passes through several forms in a biogeochemical cycle Unassimilated ammonium or nitrate may be dangerous Nitrate Assimilation Many factors regulate nitrate reductase Nitrite reductase converts nitrite to ammonium Both roots and shoots assimilate nitrate Ammonium Assimilation Converting ammonium to enzymes Ammonium can be assimilated via an alternative pathway Transamination reactions transfer nitrogen Asparagine and metabolism Amino Biological Nitrogen Fixation Free-living and symbiotic bacteria fix nitrogen Nitrogen fixation requires anaerobic conditions Symbiotic nitrogen fixation occurs in specialized structures Establishing symbiosis requires an exchange of signals Nod factors produced by bacteria act as signals for symbiosis Nodule formation involves phytohormones The nitrogenase enzyme complex fixes N-, Amides and ureides are the transported forms of nitrogen Sulfur Assimilation Sulfate Sulfate to Sulfate Methionine is synthesized from Phosphate Assimilation Cation Assimilation Cations form noncovalent bonds with carbon compounds Roots modify the rhizosphere to acquire iron Iron forms complexes with carbon and phosphate Oxygen Assimilation The Energetics of Nutrient Assimilation Summary XVÍ¡¡ 13 Secondary Metabolites and Plant Defense Cutin, Waxes, and Suberin Cutin, waxes, and suberin are made up of hydrophobic compounds Cutin, waxes, and suberin help reduce transpiration and pathogen invasion Secondary Metabolites Secondary metabolites defend plants against herbivores and pathogens Secondary metabolites are divided into three major groups Terpenes Terpenes isoprene There are two pathways for Isopentenyl diphosphate and its form larger Some development Terpenes plants Phenolic Compounds Phenylalanine is an intermediate in the biosynthesis of most plant phenolics Some simple phenolics are activated by ultraviolet light 323 The release of phenolics into the soil may limit the growth of other plants Lignin macromolecule There are four major groups of flavonoids Anthocyanins are colored flavonoids that attract animals Flavonoids may protect against damage by ultraviolet light Isoflavonoids have antimicrobial activity Tannins deter feeding by herbivores Nitrogen-Containing Compounds Alkaloids have dramatic physiological effects on animals Cyanogenic glycosides release the poison hydrogen cyanide Glucosinolates release volatile toxins Nonprotein herbivores Induced Plant Defenses against Insect Herbivores Plants can recognize specific components of insect saliva Jasmonic acid is a plant hormone that activates many defense responses Some plant proteins inhibit herbivore digestion Herbivore damage induces systemic defenses Herbivore-induced functions Plant Defense against Pathogens Some antimicrobial compounds are synthesized before pathogen attack Infection induces additional antipathogen defenses Some plants recognize specific substances released from pathogens Exposure to elicitors induces a signal transduction cascade A single encounter with a pathogen may increase resistance to future attacks Summary UNIT III Growth and Development M Gene Expression and Signal Transduction Table of 15 Cell Walls: Structure, Biogenesis, and Expansion The Structure and Synthesis of Plant Cell Walls Plant cell walls have varied architecture The primary cell wall is composed of cellulose micro- fibrils embedded in a polysaccharide matrix Cellulose microfibrils are synthesized at the plasma membrane Matrix polymers are synthesized in the Golgi and secreted via vesicles Hemicelluloses are matrix polysaccharides that bind to cellulose Pectins are gel-forming components of the matrix Structural proteins become cross-linked in the wall New primary walls are assembled during cytokinesis Secondary walls form in some cells after expansion ceases Patterns of Cell Expansion Microfibril orientation influences growth directionality of cells with diffuse growth Cortical microtubules influence the orientation of newly deposited microfibrils The Rate of Cell Elongation Stress relaxation of the cell wall drives water uptake and cell elongation The rate of cell expansion is governed by two growth equations Acid-induced growth is mediated by expansins Glucanases and other hydrolytic enzymes may modify the matrix Structural changes accompany the cessation of wall expansion Wall Degradation and Plant Defense Enzymes mediate wall hydrolysis and degradation Oxidative bursts accompany pathogen attack Wall fragments can act as signaling molecules Summary 16 Growth and Development Overview of Plant Growth and Development Sporophytic development can be divided into three major stages Development can be analyzed at the molecular level Embryogenesis: The pattern of embryogenesis differs in dicots and monocots The axial polarity of the plant is established by the embryo Position-dependent signaling guides embryogenesis Auxin may function as a morphogen during embryogenesis Genes control apical-basal patterning Embryogenesis genes have diverse biochemical functions MONOPTEROS activity is inhibited by a repressor protein Gene expression patterns correlate with auxin GNOM gene proteins Radial patterning establishes fundamental tissue layers Two genes regulate protoderm differentiation Cytokinin stimulates cell divisions for vascular elements Two genes control the differentiation of cortical and endodermal tissues through intercellular communication Intercellular communication is central to plant development Shoot Apical The shoot apical meristem forms at a position where auxin is low Forming an embryonic SAM requires many genes Shoot apical meristems vary in size and shape The shoot apical meristem contains distinct zones and layers Groups of relatively stable initial cells have been identified SAM function may require intercellular protein movement Protein turnover may spatially restrict gene activity Stem cell population is maintained by a transcriptional feedback loop Root Apical Meristem High auxin levels stimulate the formation of the root apical meristem The root tip has four developmental zones Specific root initials produce different root tissues Root apical meristems contain several types of initials Vegetative XX Table of Peridirmi Local auxin concentrations in the SAM control leaf initiation Three developmental axes describe the leaf s planar form Spatially regulated gene expression controls leaf pattern MicroRNAs regulate the sidedness of the leaf Branch roots and shoots have different origins Senescence and Programmed Cell Death Plants exhibit various types of senescence Senescence involves ordered cellular and biochemical changes Programmed cell death is a specialized type of senescence Summary 17 Phytochrome and Light Control of Plant Development The Photochemical and Biochemical Properties of Phytochrome Phytochrome can interconvert between Pr and Pfr forms Pfr is the physiologically active form of phytochrome Characteristics of Phytochrome-lnduced Responses Phytochrome responses vary in lag time and escape time Phytochrome responses can be distinguished by the amount of light required Very low-fluence responses are nonphotoreversible Low-fluence responses are High-irradiance responses are proportional to the irradiance and the duration Structure and Function of Phytochrome Proteins Phytochrome has several important functional domains Phytochrome is a light-regulated protein kinase Pfr is partitioned between the cytosol and nucleus Phytochromes are encoded by a multigene family Genetic Analysis of Phytochrome Function Phytochrome A mediates responses to continuous far-red light Phytochrome or white light Roles for phytochromes C, D, and Phy gene family interactions are complex PHY gene functions have diversified during evolution Phytochrome Signaling Pathways Phytochrome regulates membrane potentials and ion fluxes Phytochrome regulates gene expression Phytochrome interacting factors (PIFs) act early in phy signaling Phytochrome associates with protein kinases and phosphatases Phytochrome-induced gene expression involves protein degradation Circadian Rhythms The circadian oscillator involves a transcriptional negative feedback loop Ecological Functions Phytochrome regulates the sleep movements of leaves Phytochrome enables plant adaptation to light quality changes Decreasing the R:FR ratio causes elongation in sun plants Small seeds typically require a high R:FR ratio for germination Phytochrome interactions are important early in germination Reducing shade avoidance responses can improve crop yields Phytochrome responses show ecotypic variation Phytochrome action can be modulated Summary 1 and Morphogenesis The Photophysiology of Blue-Light Responses Blue light stimulates asymmetric growth and bending How do plants sense the direction of the light signal? Blue light rapidly inhibits stem elongation Blue light regulates gene expression Blue light stimulates Blue light activates a proton pump at the guard cell plasma membrane Table of Blue-light responses have characteristic kinetics and lag times Blue light regulates osmotic relations of guard cells Sucrose is an osmotically active solute in guard cells Blue-Light Photoreceptors Cryptochromes elongation Phototropins mediate blue light-dependent phototropism and The carotenoid zeaxanthin mediates blue-light photoreception in guard cells Green light reverses blue light-stimulated opening The xanthophyll cycle confers plasticity to the responses to light Summary 19 Auxin: The Growth Hormone The Emergence of the Auxin Concept Identification, Biosynthesis, and Metabolism of Auxin The principal auxin in higher plants is indole-3-acetic acid IAA is synthesized in meristems and young dividing tissues Multiple pathways exist for the biosynthesis of IAA IAA can also be synthesized from indole-3-glycerol phosphate Seeds and storage organs contain large amounts of covalently bound auxin IAA is degraded by multiple pathways IAA partitions between the cytosol and the chloroplasts Auxin Transport Polar transport requires energy and is gravity independent A chemiosmotic model has been proposed to explain polar transport P-glycoproteins are also auxin transport proteins Inhibitors of auxin transport block auxin influx and efflux Auxin is also transported nonpolarly in the phloem Auxin transport is regulated by multiple mechanisms Polar auxin transport is required for development Actions of Auxin: Cell Elongation Auxins promote growth in stems and coleoptiles, while inhibiting growth in roots The outer tissues of dicot stems are the targets of auxin action The minimum lag time for auxin-induced growth is ten minutes Auxin rapidly increases the extensibility of the cell wall Auxin-induced proton extrusion increases cell extension Auxin-induced proton extrusion may involve both activation and synthesis Actions of Auxin: Phototropism and Gravitropism Phototropism is mediated by the lateral redistribution of auxin Gravitropism involves lateral redistribution of auxin Dense plastids serve as gravity sensors Gravity sensing may involve messengers Auxin is redistributed laterally in the root cap Developmental Effects of Auxin Auxin regulates apical dominance Auxin transport regulates floral bud development and phyllotaxy Auxin promotes the formation of lateral and adventitious roots Auxin induces vascular differentiation Auxin delays the onset of leaf abscission Auxin promotes fruit development Synthetic auxins have a variety of commercial uses Auxin Signal Transduction Pathways A ubiquitin E3 Auxin-induced genes are negatively regulated by AUX/IAA proteins Auxin binding to SCFTIR1 stimulates AUX/IAA destruction Auxin-induced genes fall into two classes: early and late Rapid auxin responses may involve a different receptor protein Summary XXii Table of Contents Gibberellins: Their Discovery and Chemical Structure Gibberellins were discovered by studying a disease of rice Gibberellic acid was first purified from Gibberella culture filtrates All gibberellins are based on an enf-gibberellane skeleton Effects of Gibberellins on Growth and Development Gibberellins can stimulate stem growth Gibberellins regulate the transition from juvenile to adult phases Gibberellins influence floral initiation and sex determination Gibberellins promote pollen development and tube growth Gibberellins promote fruit set and parthenocarpy Gibberellins promote seed development and germination Commercial uses of gibberellins and GA biosynthesis inhibitors Biosynthesis and Catabolism of Gibberellins Gibberellins are synthesized via the terpenoid pathway Some enzymes in the regulated Gibberellin regulates its own metabolism GA biosynthesis occurs at multiple cellular sites Environmental conditions can influence GA biosynthesis GA, and GA4 have intrinsic bioactivity for stem growth Plant height can be genetically engineered Dwarf mutants often have other defects in addition to dwarfism Gibberellin Signaling: Significance of Response Mutants Mutations of negative regulators of GA may produce slender or dwarf phenotypes Negative regulators with agricultural importance Gibberellins signal the degradation of transcriptional repressors F-box proteins target degradation A possible GA receptor has been identified in rice Gibberellin Responses: The Cereal Aleurone Layer G A Aleurone cells may have two types of GA receptors GA signaling requires several second messengers Gibberellins enhance the transcription of a-amylase mRNA GAMYB is a positive regulator of a-amylase transcription DELLA Gibberellin Responses: Flowering in Long-Day Plants There are multiple independent pathways to flowering The long day and gibberellin pathways interact GAMYB regulates flowering and male fertility MicroRNAs regulate MYBs after transcription Gibberellin Responses: Stem Growth The shoot apical meristem interior lacks bioactive GA Gibberellins stimulate cell elongation and cell division GAs regulate the transcription of cell cycle kinases Auxin promotes GA biosynthesis and signaling Summary 21 Cytokinins: Regulators of Cell Division Cell Division and Plant Development Differentiated plant cells can resume division Diffusible factors may control cell division Plant tissues and organs can be cultured The Discovery, Identification, and Properties of Cytokinins Kinetin was discovered as a breakdown product of DNA 545 Zeatin was the first natural cytokinin discovered Some synthetic compounds can mimic or antagonize cytokinin action Cytokinins occur in both free and bound forms The hormonally active cytokinin is the free base Some plant pathogenic bacteria, fungi, insects, and nematodes secrete free cytokinins Table of Biosynthesis, Metabolism, and Transport of Cytokinins Crown gall cells have acquired a gene for cytokinin synthesis IPT catalyzes the first step in cytokinin biosynthesis Cytokinins from the root are transported to the shoot via the xylem A signal from the shoot regulates the transport of zeatin ribosides from the root Cytokinins are rapidly metabolized by plant tissues The Biological Roles of Cytokinins Cytokinins regulate cell division in shoots and roots Cytokinins regulate specific components of the cell cycle The auxinrcytokinin ratio regulates morphogenesis in cultured tissues Cytokinins modify apical dominance and promote lateral bud growth Cytokinins induce bud formation in a moss Cytokinin overproduction has been implicated in genetic tumors Cytokinins delay leaf senescence Cytokinins promote movement of nutrients Cytokinins promote Cytokinins promote cell expansion in leaves and cotyledons Cytokinin-regulated processes are revealed in plants that overproduce cytokinins Cellular and Molecular Modes of Cytokinin Action A cytokinin receptor related to bacterial two- component receptors has been identified Cytokinins increase expression of the type-A response regulator genes via activation of the type-B ARR genes Histidine phosphotransferases are also involved in cytokinin signaling Summary 22 Ethylene: Structure, Biosynthesis, and Measurement of Ethylene The properties of Bacteria, fungi, and plant organs produce Regulated biosynthesis determines the physiological activity of Environmental stresses and auxins promote biosynthesis Ethylene synthase stabilization Ethylene inhibitors Ethylene Developmental and Physiological Effects of Ethylene Ethylene Leaf epinasty results when ACC from the root is transported to the shoot Ethylene The hooks of dark-grown seedlings are maintained by ethylene Ethylene species Ethylene aquatic species Ethylene hairs Ethylene Ethylene Some defense responses are mediated by Ethylene cause abscission Ethylene Ethylene Ethylene component system histidine kinases High-affinity binding of requires a copper cofactor Unbound of the response pathway A serine ethylene EIN2 encodes Ethylene Genetic epistasis reveals the order of the signaling components Summary XXiv Table of Contents 23 Abscisic Acid: A Seed Maturation and Occurrence, Chemical Structure, and Measurement of ABA The chemical structure of ABA determines its physiological activity ABA is assayed by biological, physical, and chemical methods Biosynthesis, Metabolism, and Transport of ABA ABA is synthesized from a carotenoid intermediate ABA concentrations in tissues are highly variable ABA can be inactivated by oxidation or conjugation ABA is translocated in vascular tissue Developmental and Physiological Effects of ABA ABA regulates seed maturation ABA inhibits precocious germination and vivipary ABA promotes seed storage reserve accumulation and desiccation tolerance The seed coat and the embryo can cause dormancy Environmental factors control the release from seed dormancy Seed dormancy is controlled by the ratio of ABA to GA 24 ABA inhibits GA-induced enzyme production ABA closes ABA promotes root growth and inhibits shoot growth at low water potentials ABA promotes leaf senescence independently of ethylene ABA accumulates in dormant buds ABA Signal Transduction Pathways ABA regulates ion channels and the PM-ATPase in guard cells ABA may be perceived by both cell surface and intracellular receptors ABA signaling involves both calcium-dependent and calcium-independent pathways ABA-induced messengers ABA signaling involves protein kinases and phosphatases ABA regulates gene expression Other negative regulators also influence the ABA response Summary Brassinosteroids Brassinosteroid Structure, Occurrence, and Genetic Analysis BR-deficient mutants are impaired in photomorphogenesis Biosynthesis, Metabolism, and Transport of Brassinosteroids Brassinolide is synthesized from campesterol Catabolism and negative feedback contribute to BR homeostasis Brassinosteroids act locally near their sites of synthesis Brassinosteroids: Effects on Growth and Development BRs promote both cell expansion and cell division in shoots BRs both promote and inhibit root growth BRs promote xylem differentiation during vascular development BRs are required for the growth of pollen tubes BRs promote seed germination The Brassinosteroid Signaling Pathway BR-insensitive mutants identified the BR cell surface receptor Phosphorylation activates the BIN2 is a repressor of BR-induced gene expression BESI Prospective Uses of Brassinosteroids in Agriculture Summary Table of Contents 25 The Control of Flowering Floral The shoot apical meristems in Arabidopsis change with development The four different types of floral organs are initiated as separate whorls Three types of genes regulate floral development Meristem function Homeotic mutations led to the identification of floral organ identity genes Three types of homeotic genes control floral organ identity The ABC model explains the determination of floral organ identity Floral Evocation: Internal and External Cues The Shoot Apex and Phase Changes Shoot apical meristems have three developmental phases Juvenile tissues are produced first and are located at the base of the shoot Phase changes can be influenced by nutrients, gibberellins, and other chemical signals Competence and determination are two stages in floral evocation Orcadian Rhythms: The Clock Within Circadian rhythms exhibit characteristic features Phase shifting adjusts circadian rhythms to different day-night cycles Phytochromes and clock Photoperiodism: Monitoring Day Length Plants can be classified according to their photoperiodic responses The leaf is the site of perception of the photoperiodic signal The floral stimulus is transported in the phloem 26 Stress Physiology Water Deficit and Drought Tolerance Drought resistance strategies can vary Decreased leaf area is an early response to water deficit Water deficit stimulates leaf abscission Water deficit enhances root growth Abscisic acid induces deficit Water deficit limits photosynthesis Plants monitor day length by measuring the length of the night Night breaks can cancel the effect of the dark period The circadian clock and photoperiodic timekeeping The coincidence model is based on oscillating light sensitivity The coincidence of promotes flowering in LDPs The coincidence of Heading-date inhibits flowering in SDPs Phytochrome is the primary photoreceptor in photoperiodism A blue-light photoreceptor regulates flowering in some LDPs Vernalization: Promoting Flowering with Cold Vernalization results in competence to flower at the shoot apical meristem Vernalization involves epigenetic changes in gene expression A variety of vernalization mechanisms may have evolved Biochemical Signaling Involved in Flowering Grafting studies have provided evidence for a transmissible floral stimulus Indirect induction implies that the floral stimulus is self-propagating Evidence for antiflorigen has been found in some LDPs Florigen FLOWERING LOCUS photoperiodic floral stimulus Gibberellins and plants The transition to flowering involves multiple factors and pathways Summary Osmotic adjustment of cells helps maintain water balance Water deficit increases resistance to water flow Water deficit increases leaf wax deposition Water deficit alters energy dissipation from leaves CAM plants are adapted to water stress Osmotic stress changes gene expression ABA-dependent and ABA-independent signaling pathways regulate stress tolerance XXVi Table of Contents Heat Stress and Heat Shock High leaf temperature and minimal evaporative cooling lead to heat stress At high temperatures, photosynthesis is inhibited before respiration Plants adapted to cool temperatures acclimate poorly to high temperatures Temperature affects membrane stability Several adaptations protect leaves against excessive heating At higher temperatures, plants produce protective proteins A transcription factor mediates HSPs mediate tolerance to high temperatures Several signaling pathways mediate thermotolerance responses Chilling and Freezing Membrane properties change in response to chilling injury Ice crystal formation and protoplast dehydration kill cells Limitation of ice formation contributes to freezing tolerance Some woody plants can acclimate to very low temperatures Some bacteria living on leaf surfaces increase frost damage Acclimation to freezing involves ABA and protein synthesis Numerous genes are induced during cold acclimation A transcription factor regulates cold-induced gene expression Salinity Stress Salt accumulation in irrigated soils impairs plant function Plants show great diversity for salt tolerance Salt stress causes multiple injury effects Plants use multiple strategies to reduce salt stress Ion exclusion and compartmentation reduce salinity stress Plant adaptations to toxic trace elements Oxygen Deficiency Anaerobic microorganisms are active in water- saturated soils Roots are damaged in anoxic environments Damaged O2-deficient roots injure shoots Submerged organs can acquire O2 through specialized structures Most plant tissues cannot tolerate anaerobic conditions Synthesis of anaerobic stress proteins leads to acclimation to O2 deficit Summary Glossary Author Index Subject Index
adam_txt	Table Preface v Authors and Contributors vu 1 Plant Cells Plant Life: Unifying Principles Overview of Plant Structure Plant cells are surrounded by rigid cell walls New cells are produced by dividing tissues called meristems Three major tissue systems make up the plant body The Plant Cell Biological membranes are phospholipid bilayers that contain proteins The nucleus contains most of the genetic material of the cell Protein synthesis involves transcription and translation The endoplasmic reticulum is a network of internal membranes Secretion of proteins from cells begins with the rough ER 11 Golgi stacks produce and distribute secretory products Proteins and polysaccharides destined for secretion are processed in the Golgi apparatus Two models for intra-Golgi transport have been proposed Specific coat proteins facilitate vesicle budding Vacuoles Mitochondria and chloroplasts are sites of energy conversion Mitochondria and chloroplasts are semiautonomous organelles Different plastid types are interconvertible Microbodies play specialized metabolic roles in leaves and seeds Oleosomes are lipid-storing organelles The Cytoskeleton Plant cells contain microtubules, intermediate filaments Microtubules and disassemble Microtubules function in mitosis and cytokinesis Motor proteins mediate cytoplasmic streaming and organelle movements Cell Cycle Regulation Each phase of the cell cycle has a specific set of biochemical and cellular activities The cell cycle is regulated by cyclin-dependent kinases Plasmodesmata There are two types of plasmodesmata: primary and secondary Plasmodesmata have a complex internal structure Macromolecular traffic through plasmodesmata is important for developmental signaling Summary 29 Ł XÜ UNITI Transport and 3 Water and Plant Cells Water in Plant Life The Structure and Properties of Water The polarity of water molecules gives rise to hydrogen bonds The polarity of water makes it an excellent solvent The thermal properties of water result from hydrogen bonding The cohesive and adhesive properties of water are due to hydrogen bonding Water has a high tensile strength Water Transport Processes Diffusion is the movement of molecules by random thermal agitation Diffusion is rapid over short distances but extremely slow over long distances Pressure-driven bulk flow drives long-distance water transport Osmosis is driven by a water potential gradient 4 Water Balance of Plants Water in the Soil A negative hydrostatic pressure in soil water lowers soil water potential Water moves through the soil by bulk flow Water Absorption by Roots Water moves in the root via the apoplast, symplast, and transmembrane Solute accumulation in the xylem can generate "root pressure" Water Transport through the Xylem The xylem consists of two types of tracheary elements Water movement through the xylem requires less pressure than movement through living cells What pressure difference is needed to lift water meters to a treetop? The cohesion-tension theory explains water transport in the xylem Xylem transport of water in trees faces physical challenges The chemical potential of water represents the free- energy status of water Three major factors contribute to cell water potential Water enters the cell along a water potential gradient Water can also leave the cell in response to a water potential gradient Small changes in plant cell volume cause large changes in Water transport rates depend on driving force and hydraulic conductivity Aquaporins facilitate the movement of water across cell membranes The water potential concept helps us evaluate the water status of a plant The components of water potential vary with growth conditions and location within the plant Summary Plants minimize the consequences of xylem cavitation Water Movement from the Leaf to the Atmosphere The driving force for water loss is the difference in water vapor concentration Water loss is also regulated by the pathway resistances Stomatal photosynthesis The cell walls of guard cells have specialized features An increase in guard cell stornata The transpiration ratio measures the relationship between water loss and carbon gain Overview: The Soil-Plant-Atmosphere Continuum Summary Table 5 Mineral Essential Nutrients, Deficiencies, and Plant Disorders Special techniques are used in nutritional studies Nutrient solutions can sustain rapid plant growth Mineral deficiencies disrupt plant metabolism and function Analysis of plant tissues reveals mineral deficiencies Treating Nutritional Deficiencies Crop yields can be improved by addition of fertilizers 83 Some mineral nutrients can be absorbed by leaves Soil, Roots, and Microbes Negatively charged soil particles affect the adsorption 6 Solute Transport Passive and Active Transport Transport of Ions across a Membrane Barrier Different diffusion rates for cations and anions produce diffusion potentials How does membrane potential relate to ion distribution? The Nernst equation distinguishes between active and passive transport Proton transport is a major determinant of the membrane potential Membrane Transport Processes Channel transporters enhance diffusion across membranes Carriers bind and transport specific substances Primary active transport requires energy Secondary active transport uses stored energy Kinetic analyses can elucidate transport mechanisms Membrane Transport Proteins 105 106 of mineral nutrients Soil root growth Excess minerals in the soil limit plant growth Plants develop extensive root systems Root systems differ in form but are based on common structures Different areas of the root absorb different mineral ions Mycorrhizal fungi facilitate nutrient uptake by roots 89 Nutrients move from the mycorrhizal fungi to the root cells Summary The genes for many transporters have been identified Transporters exist for diverse nitrogen-containing compounds Cation transporters are diverse Some Metals are transported by ZIP proteins Aquaporins may have novel functions The plasma membrane H+-ATPase has several functional domains The tonoplast H+-ATPase drives solute accumulation into H+-pyrophosphatases also pump protons at the tonoplast Ion Transport in Roots Solutes move through both apoplast and symplast Ions cross both symplast and apoplast Xylem parenchyma cells participate in xylem loading Summary Xiv UNITU Biochemistry and Metabolism 1 Photosynthesis: The Light Reactions Photosynthesis in Higher Plants General Concepts Light has characteristics of both a particle and a wave When molecules absorb or emit light, they change their electronic state Photosynthetic pigments absorb the light that powers photosynthesis Key Experiments in Understanding Photosynthesis Action spectra relate light absorption to photosynthetic activity Photosynthesis takes place in complexes containing light-harvesting antennas and photochemical reaction centers The chemical reaction of photosynthesis is driven by light Light drives the reduction of NADP and the formation of ATP Oxygen-evolving organisms have two photosystems that operate in series Organization of the Photosynthetic Apparatus The Thylakoids contain integral membrane proteins Photosystems I and II are spatially separated in the thylakoid membrane Anoxygenic photosynthetic bacteria have a single reaction center Organization of Light-Absorbing Antenna Systems The antenna funnels energy to the reaction center Many antenna complexes have a common structural motif Mechanisms of Electron Transport Electrons from chlorophyll travel through the carriers organized in the "Z scheme" Energy is captured when an excited chlorophyll reduces an electron acceptor molecule The reaction center chlorophylls of the two photosystems absorb at different wavelengths The photosystem II reaction center is a multisubunit pigment-protein complex Water is oxidized to oxygen by photosystem II Pheophytin and two photosystem II Electron flow through the cytochrome b6 transports protons Plastoquinone and plastocyanin carry electrons between photosystems II and I The photosystem I reaction center reduces NADP+ Cyclic electron flow generates ATP but no NADPH Some herbicides block photosynthetic electron flow Proton Transport and ATP Synthesis in the Chloroplast Repair and Regulation of the Photosynthetic Machinery Carotenoids serve as photoprotective agents Some xanthophylls also participate in energy dissipation The photosystem II reaction center is easily damaged Photosystem I is protected from active oxygen species Thylakoid stacking permits energy partitioning between the photosystems Genetics, Assembly, and Evolution of Photosynthetic Systems Chloroplast, cyanobacterial, been sequenced Chloroplast inheritance Many cytoplasm The biosynthesis and breakdown of chlorophyll are complex pathways Complex photosynthetic organisms have evolved from simpler forms Summary Table of Contents XV б Photosynthesis: Carbon Reactions The Calvin Cycle The Calvin cycle has three stages: carboxylation, reduction, and regeneration The carboxylation of ribulose-l,5-bisphosphate is catalyzed by the enzyme rubisco Operation of the Calvin cycle requires the regeneration of ribulose-l,5-bisphosphate The Calvin cycle regenerates its own biochemical components The Calvin cycle uses energy very efficiently Regulation of the Calvin Cycle Light regulates the Calvin cycle The activity of rubisco increases in the light The ferredoxin-thioredoxin system regulates the Calvin cycle Light-dependent ion movements regulate Calvin cycle enzymes The C2 Oxidative Photosynthetic Carbon Cycle Photosynthetic CO2 fixation and photorespiratory oxygénation Photorespiration electron transport system The biological function of investigation COj-Concentrating Mechanisms I. CO2 and HCO3- Pumps II. The C4 Carbon Cycle Malate C4 cycle Two different types of cells participate in the C4 cycle The C4 cycle concentrates CO2 in the bundle sheath cells The C4 cycle also concentrates CO2 in single cells The C4 cycle has higher energy demand than the Calvin cycle Light regulates the activity of key C4 enzymes In hot, dry climates, the C4 cycle reduces photorespiration III. Crassulacean Acid Metabolism (CAM) The during the day Some CAM plants change the pattern of CO2 uptake in response to environmental conditions Starch and Sucrose Chloroplast degraded at night Starch is synthesized in the Starch degradation requires phosphorylation of amylopectin Triose the pool of hexose phosphates in the cytosol Fructose-6-phosphate can be converted to fructose- 1,6-bisphosphate by two different enzymes Fructose-2,6-bisphosphate is an important regulatory compound The hexose phosphate pool is regulated by fructose-2,6- bisphosphate Sucrose is continuously synthesized in the cytosol Summary У Light, Leaves, and Photosynthesis Units in the Measurement of Light Leaf anatomy maximizes light absorption Plants compete for sunlight Leaf angle and leaf movement can control light absorption Plants acclimate and adapt to sun and shade Photosynthetic Responses to Light by the Intact Leaf Light-response curves reveal photosynthetic properties Leaves must dissipate excess light energy Absorption of too much light can lead to photoinhibition Photosynthetic Responses to Temperature Leaves must dissipate vast quantities of heat Photosynthesis is temperature sensitive Photosynthetic Responses to Carbon Dioxide Atmospheric CO2 concentration keeps rising CO2 diffusion to the photosynthesis Patterns of light absorption generate gradients of CO2 fixation CO2 imposes limitations on photosynthesis Crassulacean Acid Metabolism Carbon isotope ratio variations reveal different photosynthetic pathways How do we measure the carbon isotopes of plants? Why are there carbon isotope ratio variations in plants? Summary xvi 10 Translocation Pathways of Sugar is translocated in phloem sieve elements Mature sieve elements are living cells specialized for translocation Large pores in cell walls are the prominent feature of sieve elements Damaged sieve elements are sealed off Companion cells aid the highly specialized sieve elements Patterns of Source-to-sink pathways follow anatomic and developmental patterns Materials Translocated in the Phloem Phloem sap can be collected and analyzed Sugars are translocated in nonreducing form Rates of Movement The Pressure-Flow Model for Phloem Transport A pressure gradient drives pressure-flow model The predictions of mass flow have been confirmed Sieve plate pores are open channels There is no bidirectional transport in single sieve elements The energy requirement for transport through the phloem pathway is small Pressure gradients are sufficient to drive a mass flow of phloem sap Significant questions about the pressure-flow model still exist Phloem Loading Phloem loading can occur via the apoplast or symplast Sucrose uptake in the apoplastic pathway requires metabolic energy Phloem loading in the apoplastic pathway involves a sucrose-H"1" symporter Phloem loading is symplastic in plants with intermediary cells The polymer-trapping model explains symplastic loading The type of phloem loading is correlated with plant family and with climate Phloem Unloading and Sink-to-Source Transition Phloem unloading and short-distance transport can occur via symplastic or apoplastic pathways Transport into sink tissues requires metabolic energy The transition of a leaf from sink to source is gradual Photosynthate Distribution: Allocation and Partitioning Allocation includes storage, utilization, and transport Various sinks partition transport sugars Source leaves regulate allocation Sink tissues compete for available translocated photosynthate Sink strength depends on sink size and activity The source adjusts over the long term to changes in the source-to-sink ratio The Transport of Signaling Molecules Turgor and sink activities Signal molecules in the phloem regulate growth and development Summary 11 Respiration and Overview of Plant Respiration Glycolysis: A Cytosolic and Plastidic Process Glycolysis converts carbohydrates into pyruvate, producing NADH and ATP Plants have alternative glycolytic reactions In the absence of O,, fermentation regenerates the NAD+ needed for glycolysis Fermentation does not liberate all the energy available in each sugar molecule Plant glycolysis is controlled by its products The pentose phosphate pathway produces NADPH and biosynthetic intermediates The Citric Acid Cycle: A Mitochondrial Matrix Process Mitochondria are semiautonomous organelles Pyruvate enters the mitochondrion and is oxidized via the citric acid cycle The citric acid cycle of plants has unique features Mitochondrial Electron Transport and ATP Synthesis The electron transport chain catalyzes a flow of electrons from NADH to O2 Some electron transport enzymes are unique to plant mitochondria Table of ATP synthesis in the mitochondrion is coupled to electron transport Transporters exchange substrates and products Aerobic respiration yields about per molecule of sucrose Several subunits of respiratory complexes are encoded by the mitochondrial genome Plants have several mechanisms that lower the ATP yield Mitochondrial respiration is controlled by key metabolites Respiration is tightly coupled to other pathways Respiration in Intact Plants and Tissues Plants respire roughly half of the daily photosynthetic yield Respiration operates during photosynthesis Different tissues and organs respire at different rates Mitochondrial function is crucial during pollen development Environmental factors alter respiration rates Lipid Fats and oils store large amounts of energy Triacylglycerols are stored in oil bodies Polar glycerolipids are the main structural lipids in membranes Fatty acid biosynthesis consists of cycles of two-carbon addition Glycerolipids are synthesized in the plastids and the 282 Lipid Membrane lipids are precursors of important signaling compounds Storage lipids are converted into carbohydrates in germinating seeds Summary 12 Assimilation of Mineral Nutrients Nitrogen in the Environment Nitrogen passes through several forms in a biogeochemical cycle Unassimilated ammonium or nitrate may be dangerous Nitrate Assimilation Many factors regulate nitrate reductase Nitrite reductase converts nitrite to ammonium Both roots and shoots assimilate nitrate Ammonium Assimilation Converting ammonium to enzymes Ammonium can be assimilated via an alternative pathway Transamination reactions transfer nitrogen Asparagine and metabolism Amino Biological Nitrogen Fixation Free-living and symbiotic bacteria fix nitrogen Nitrogen fixation requires anaerobic conditions Symbiotic nitrogen fixation occurs in specialized structures Establishing symbiosis requires an exchange of signals Nod factors produced by bacteria act as signals for symbiosis Nodule formation involves phytohormones The nitrogenase enzyme complex fixes N-, Amides and ureides are the transported forms of nitrogen Sulfur Assimilation Sulfate Sulfate to Sulfate Methionine is synthesized from Phosphate Assimilation Cation Assimilation Cations form noncovalent bonds with carbon compounds Roots modify the rhizosphere to acquire iron Iron forms complexes with carbon and phosphate Oxygen Assimilation The Energetics of Nutrient Assimilation Summary XVÍ¡¡ 13 Secondary Metabolites and Plant Defense Cutin, Waxes, and Suberin Cutin, waxes, and suberin are made up of hydrophobic compounds Cutin, waxes, and suberin help reduce transpiration and pathogen invasion Secondary Metabolites Secondary metabolites defend plants against herbivores and pathogens Secondary metabolites are divided into three major groups Terpenes Terpenes isoprene There are two pathways for Isopentenyl diphosphate and its form larger Some development Terpenes plants Phenolic Compounds Phenylalanine is an intermediate in the biosynthesis of most plant phenolics Some simple phenolics are activated by ultraviolet light 323 The release of phenolics into the soil may limit the growth of other plants Lignin macromolecule There are four major groups of flavonoids Anthocyanins are colored flavonoids that attract animals Flavonoids may protect against damage by ultraviolet light Isoflavonoids have antimicrobial activity Tannins deter feeding by herbivores Nitrogen-Containing Compounds Alkaloids have dramatic physiological effects on animals Cyanogenic glycosides release the poison hydrogen cyanide Glucosinolates release volatile toxins Nonprotein herbivores Induced Plant Defenses against Insect Herbivores Plants can recognize specific components of insect saliva Jasmonic acid is a plant hormone that activates many defense responses Some plant proteins inhibit herbivore digestion Herbivore damage induces systemic defenses Herbivore-induced functions Plant Defense against Pathogens Some antimicrobial compounds are synthesized before pathogen attack Infection induces additional antipathogen defenses Some plants recognize specific substances released from pathogens Exposure to elicitors induces a signal transduction cascade A single encounter with a pathogen may increase resistance to future attacks Summary UNIT III Growth and Development M Gene Expression and Signal Transduction Table of 15 Cell Walls: Structure, Biogenesis, and Expansion The Structure and Synthesis of Plant Cell Walls Plant cell walls have varied architecture The primary cell wall is composed of cellulose micro- fibrils embedded in a polysaccharide matrix Cellulose microfibrils are synthesized at the plasma membrane Matrix polymers are synthesized in the Golgi and secreted via vesicles Hemicelluloses are matrix polysaccharides that bind to cellulose Pectins are gel-forming components of the matrix Structural proteins become cross-linked in the wall New primary walls are assembled during cytokinesis Secondary walls form in some cells after expansion ceases Patterns of Cell Expansion Microfibril orientation influences growth directionality of cells with diffuse growth Cortical microtubules influence the orientation of newly deposited microfibrils The Rate of Cell Elongation Stress relaxation of the cell wall drives water uptake and cell elongation The rate of cell expansion is governed by two growth equations Acid-induced growth is mediated by expansins Glucanases and other hydrolytic enzymes may modify the matrix Structural changes accompany the cessation of wall expansion Wall Degradation and Plant Defense Enzymes mediate wall hydrolysis and degradation Oxidative bursts accompany pathogen attack Wall fragments can act as signaling molecules Summary 16 Growth and Development Overview of Plant Growth and Development Sporophytic development can be divided into three major stages Development can be analyzed at the molecular level Embryogenesis: The pattern of embryogenesis differs in dicots and monocots The axial polarity of the plant is established by the embryo Position-dependent signaling guides embryogenesis Auxin may function as a morphogen during embryogenesis Genes control apical-basal patterning Embryogenesis genes have diverse biochemical functions MONOPTEROS activity is inhibited by a repressor protein Gene expression patterns correlate with auxin GNOM gene proteins Radial patterning establishes fundamental tissue layers Two genes regulate protoderm differentiation Cytokinin stimulates cell divisions for vascular elements Two genes control the differentiation of cortical and endodermal tissues through intercellular communication Intercellular communication is central to plant development Shoot Apical The shoot apical meristem forms at a position where auxin is low Forming an embryonic SAM requires many genes Shoot apical meristems vary in size and shape The shoot apical meristem contains distinct zones and layers Groups of relatively stable initial cells have been identified SAM function may require intercellular protein movement Protein turnover may spatially restrict gene activity Stem cell population is maintained by a transcriptional feedback loop Root Apical Meristem High auxin levels stimulate the formation of the root apical meristem The root tip has four developmental zones Specific root initials produce different root tissues Root apical meristems contain several types of initials Vegetative XX Table of Peridirmi Local auxin concentrations in the SAM control leaf initiation Three developmental axes describe the leaf's planar form Spatially regulated gene expression controls leaf pattern MicroRNAs regulate the sidedness of the leaf Branch roots and shoots have different origins Senescence and Programmed Cell Death Plants exhibit various types of senescence Senescence involves ordered cellular and biochemical changes Programmed cell death is a specialized type of senescence Summary 17 Phytochrome and Light Control of Plant Development The Photochemical and Biochemical Properties of Phytochrome Phytochrome can interconvert between Pr and Pfr forms Pfr is the physiologically active form of phytochrome Characteristics of Phytochrome-lnduced Responses Phytochrome responses vary in lag time and escape time Phytochrome responses can be distinguished by the amount of light required Very low-fluence responses are nonphotoreversible Low-fluence responses are High-irradiance responses are proportional to the irradiance and the duration Structure and Function of Phytochrome Proteins Phytochrome has several important functional domains Phytochrome is a light-regulated protein kinase Pfr is partitioned between the cytosol and nucleus Phytochromes are encoded by a multigene family Genetic Analysis of Phytochrome Function Phytochrome A mediates responses to continuous far-red light Phytochrome or white light Roles for phytochromes C, D, and Phy gene family interactions are complex PHY gene functions have diversified during evolution Phytochrome Signaling Pathways Phytochrome regulates membrane potentials and ion fluxes Phytochrome regulates gene expression Phytochrome interacting factors (PIFs) act early in phy signaling Phytochrome associates with protein kinases and phosphatases Phytochrome-induced gene expression involves protein degradation Circadian Rhythms The circadian oscillator involves a transcriptional negative feedback loop Ecological Functions Phytochrome regulates the sleep movements of leaves Phytochrome enables plant adaptation to light quality changes Decreasing the R:FR ratio causes elongation in sun plants Small seeds typically require a high R:FR ratio for germination Phytochrome interactions are important early in germination Reducing shade avoidance responses can improve crop yields Phytochrome responses show ecotypic variation Phytochrome action can be modulated Summary 1 and Morphogenesis The Photophysiology of Blue-Light Responses Blue light stimulates asymmetric growth and bending How do plants sense the direction of the light signal? Blue light rapidly inhibits stem elongation Blue light regulates gene expression Blue light stimulates Blue light activates a proton pump at the guard cell plasma membrane Table of Blue-light responses have characteristic kinetics and lag times Blue light regulates osmotic relations of guard cells Sucrose is an osmotically active solute in guard cells Blue-Light Photoreceptors Cryptochromes elongation Phototropins mediate blue light-dependent phototropism and The carotenoid zeaxanthin mediates blue-light photoreception in guard cells Green light reverses blue light-stimulated opening The xanthophyll cycle confers plasticity to the responses to light Summary 19 Auxin: The Growth Hormone The Emergence of the Auxin Concept Identification, Biosynthesis, and Metabolism of Auxin The principal auxin in higher plants is indole-3-acetic acid IAA is synthesized in meristems and young dividing tissues Multiple pathways exist for the biosynthesis of IAA IAA can also be synthesized from indole-3-glycerol phosphate Seeds and storage organs contain large amounts of covalently bound auxin IAA is degraded by multiple pathways IAA partitions between the cytosol and the chloroplasts Auxin Transport Polar transport requires energy and is gravity independent A chemiosmotic model has been proposed to explain polar transport P-glycoproteins are also auxin transport proteins Inhibitors of auxin transport block auxin influx and efflux Auxin is also transported nonpolarly in the phloem Auxin transport is regulated by multiple mechanisms Polar auxin transport is required for development Actions of Auxin: Cell Elongation Auxins promote growth in stems and coleoptiles, while inhibiting growth in roots The outer tissues of dicot stems are the targets of auxin action The minimum lag time for auxin-induced growth is ten minutes Auxin rapidly increases the extensibility of the cell wall Auxin-induced proton extrusion increases cell extension Auxin-induced proton extrusion may involve both activation and synthesis Actions of Auxin: Phototropism and Gravitropism Phototropism is mediated by the lateral redistribution of auxin Gravitropism involves lateral redistribution of auxin Dense plastids serve as gravity sensors Gravity sensing may involve messengers Auxin is redistributed laterally in the root cap Developmental Effects of Auxin Auxin regulates apical dominance Auxin transport regulates floral bud development and phyllotaxy Auxin promotes the formation of lateral and adventitious roots Auxin induces vascular differentiation Auxin delays the onset of leaf abscission Auxin promotes fruit development Synthetic auxins have a variety of commercial uses Auxin Signal Transduction Pathways A ubiquitin E3 Auxin-induced genes are negatively regulated by AUX/IAA proteins Auxin binding to SCFTIR1 stimulates AUX/IAA destruction Auxin-induced genes fall into two classes: early and late Rapid auxin responses may involve a different receptor protein Summary XXii Table of Contents Gibberellins: Their Discovery and Chemical Structure Gibberellins were discovered by studying a disease of rice Gibberellic acid was first purified from Gibberella culture filtrates All gibberellins are based on an enf-gibberellane skeleton Effects of Gibberellins on Growth and Development Gibberellins can stimulate stem growth Gibberellins regulate the transition from juvenile to adult phases Gibberellins influence floral initiation and sex determination Gibberellins promote pollen development and tube growth Gibberellins promote fruit set and parthenocarpy Gibberellins promote seed development and germination Commercial uses of gibberellins and GA biosynthesis inhibitors Biosynthesis and Catabolism of Gibberellins Gibberellins are synthesized via the terpenoid pathway Some enzymes in the regulated Gibberellin regulates its own metabolism GA biosynthesis occurs at multiple cellular sites Environmental conditions can influence GA biosynthesis GA, and GA4 have intrinsic bioactivity for stem growth Plant height can be genetically engineered Dwarf mutants often have other defects in addition to dwarfism Gibberellin Signaling: Significance of Response Mutants Mutations of negative regulators of GA may produce slender or dwarf phenotypes Negative regulators with agricultural importance Gibberellins signal the degradation of transcriptional repressors F-box proteins target degradation A possible GA receptor has been identified in rice Gibberellin Responses: The Cereal Aleurone Layer G A Aleurone cells may have two types of GA receptors GA signaling requires several second messengers Gibberellins enhance the transcription of a-amylase mRNA GAMYB is a positive regulator of a-amylase transcription DELLA Gibberellin Responses: Flowering in Long-Day Plants There are multiple independent pathways to flowering The long day and gibberellin pathways interact GAMYB regulates flowering and male fertility MicroRNAs regulate MYBs after transcription Gibberellin Responses: Stem Growth The shoot apical meristem interior lacks bioactive GA Gibberellins stimulate cell elongation and cell division GAs regulate the transcription of cell cycle kinases Auxin promotes GA biosynthesis and signaling Summary 21 Cytokinins: Regulators of Cell Division Cell Division and Plant Development Differentiated plant cells can resume division Diffusible factors may control cell division Plant tissues and organs can be cultured The Discovery, Identification, and Properties of Cytokinins Kinetin was discovered as a breakdown product of DNA 545 Zeatin was the first natural cytokinin discovered Some synthetic compounds can mimic or antagonize cytokinin action Cytokinins occur in both free and bound forms The hormonally active cytokinin is the free base Some plant pathogenic bacteria, fungi, insects, and nematodes secrete free cytokinins Table of Biosynthesis, Metabolism, and Transport of Cytokinins Crown gall cells have acquired a gene for cytokinin synthesis IPT catalyzes the first step in cytokinin biosynthesis Cytokinins from the root are transported to the shoot via the xylem A signal from the shoot regulates the transport of zeatin ribosides from the root Cytokinins are rapidly metabolized by plant tissues The Biological Roles of Cytokinins Cytokinins regulate cell division in shoots and roots Cytokinins regulate specific components of the cell cycle The auxinrcytokinin ratio regulates morphogenesis in cultured tissues Cytokinins modify apical dominance and promote lateral bud growth Cytokinins induce bud formation in a moss Cytokinin overproduction has been implicated in genetic tumors Cytokinins delay leaf senescence Cytokinins promote movement of nutrients Cytokinins promote Cytokinins promote cell expansion in leaves and cotyledons Cytokinin-regulated processes are revealed in plants that overproduce cytokinins Cellular and Molecular Modes of Cytokinin Action A cytokinin receptor related to bacterial two- component receptors has been identified Cytokinins increase expression of the type-A response regulator genes via activation of the type-B ARR genes Histidine phosphotransferases are also involved in cytokinin signaling Summary 22 Ethylene: Structure, Biosynthesis, and Measurement of Ethylene The properties of Bacteria, fungi, and plant organs produce Regulated biosynthesis determines the physiological activity of Environmental stresses and auxins promote biosynthesis Ethylene synthase stabilization Ethylene inhibitors Ethylene Developmental and Physiological Effects of Ethylene Ethylene Leaf epinasty results when ACC from the root is transported to the shoot Ethylene The hooks of dark-grown seedlings are maintained by ethylene Ethylene species Ethylene aquatic species Ethylene hairs Ethylene Ethylene Some defense responses are mediated by Ethylene cause abscission Ethylene Ethylene Ethylene component system histidine kinases High-affinity binding of requires a copper cofactor Unbound of the response pathway A serine ethylene EIN2 encodes Ethylene Genetic epistasis reveals the order of the signaling components Summary XXiv Table of Contents 23 Abscisic Acid: A Seed Maturation and Occurrence, Chemical Structure, and Measurement of ABA The chemical structure of ABA determines its physiological activity ABA is assayed by biological, physical, and chemical methods Biosynthesis, Metabolism, and Transport of ABA ABA is synthesized from a carotenoid intermediate ABA concentrations in tissues are highly variable ABA can be inactivated by oxidation or conjugation ABA is translocated in vascular tissue Developmental and Physiological Effects of ABA ABA regulates seed maturation ABA inhibits precocious germination and vivipary ABA promotes seed storage reserve accumulation and desiccation tolerance The seed coat and the embryo can cause dormancy Environmental factors control the release from seed dormancy Seed dormancy is controlled by the ratio of ABA to GA 24 ABA inhibits GA-induced enzyme production ABA closes ABA promotes root growth and inhibits shoot growth at low water potentials ABA promotes leaf senescence independently of ethylene ABA accumulates in dormant buds ABA Signal Transduction Pathways ABA regulates ion channels and the PM-ATPase in guard cells ABA may be perceived by both cell surface and intracellular receptors ABA signaling involves both calcium-dependent and calcium-independent pathways ABA-induced messengers ABA signaling involves protein kinases and phosphatases ABA regulates gene expression Other negative regulators also influence the ABA response Summary Brassinosteroids Brassinosteroid Structure, Occurrence, and Genetic Analysis BR-deficient mutants are impaired in photomorphogenesis Biosynthesis, Metabolism, and Transport of Brassinosteroids Brassinolide is synthesized from campesterol Catabolism and negative feedback contribute to BR homeostasis Brassinosteroids act locally near their sites of synthesis Brassinosteroids: Effects on Growth and Development BRs promote both cell expansion and cell division in shoots BRs both promote and inhibit root growth BRs promote xylem differentiation during vascular development BRs are required for the growth of pollen tubes BRs promote seed germination The Brassinosteroid Signaling Pathway BR-insensitive mutants identified the BR cell surface receptor Phosphorylation activates the BIN2 is a repressor of BR-induced gene expression BESI Prospective Uses of Brassinosteroids in Agriculture Summary Table of Contents 25 The Control of Flowering Floral The shoot apical meristems in Arabidopsis change with development The four different types of floral organs are initiated as separate whorls Three types of genes regulate floral development Meristem function Homeotic mutations led to the identification of floral organ identity genes Three types of homeotic genes control floral organ identity The ABC model explains the determination of floral organ identity Floral Evocation: Internal and External Cues The Shoot Apex and Phase Changes Shoot apical meristems have three developmental phases Juvenile tissues are produced first and are located at the base of the shoot Phase changes can be influenced by nutrients, gibberellins, and other chemical signals Competence and determination are two stages in floral evocation Orcadian Rhythms: The Clock Within Circadian rhythms exhibit characteristic features Phase shifting adjusts circadian rhythms to different day-night cycles Phytochromes and clock Photoperiodism: Monitoring Day Length Plants can be classified according to their photoperiodic responses The leaf is the site of perception of the photoperiodic signal The floral stimulus is transported in the phloem 26 Stress Physiology Water Deficit and Drought Tolerance Drought resistance strategies can vary Decreased leaf area is an early response to water deficit Water deficit stimulates leaf abscission Water deficit enhances root growth Abscisic acid induces deficit Water deficit limits photosynthesis Plants monitor day length by measuring the length of the night Night breaks can cancel the effect of the dark period The circadian clock and photoperiodic timekeeping The coincidence model is based on oscillating light sensitivity The coincidence of promotes flowering in LDPs The coincidence of Heading-date inhibits flowering in SDPs Phytochrome is the primary photoreceptor in photoperiodism A blue-light photoreceptor regulates flowering in some LDPs Vernalization: Promoting Flowering with Cold Vernalization results in competence to flower at the shoot apical meristem Vernalization involves epigenetic changes in gene expression A variety of vernalization mechanisms may have evolved Biochemical Signaling Involved in Flowering Grafting studies have provided evidence for a transmissible floral stimulus Indirect induction implies that the floral stimulus is self-propagating Evidence for antiflorigen has been found in some LDPs Florigen FLOWERING LOCUS photoperiodic floral stimulus Gibberellins and plants The transition to flowering involves multiple factors and pathways Summary Osmotic adjustment of cells helps maintain water balance Water deficit increases resistance to water flow Water deficit increases leaf wax deposition Water deficit alters energy dissipation from leaves CAM plants are adapted to water stress Osmotic stress changes gene expression ABA-dependent and ABA-independent signaling pathways regulate stress tolerance XXVi Table of Contents Heat Stress and Heat Shock High leaf temperature and minimal evaporative cooling lead to heat stress At high temperatures, photosynthesis is inhibited before respiration Plants adapted to cool temperatures acclimate poorly to high temperatures Temperature affects membrane stability Several adaptations protect leaves against excessive heating At higher temperatures, plants produce protective proteins A transcription factor mediates HSPs mediate tolerance to high temperatures Several signaling pathways mediate thermotolerance responses Chilling and Freezing Membrane properties change in response to chilling injury Ice crystal formation and protoplast dehydration kill cells Limitation of ice formation contributes to freezing tolerance Some woody plants can acclimate to very low temperatures Some bacteria living on leaf surfaces increase frost damage Acclimation to freezing involves ABA and protein synthesis Numerous genes are induced during cold acclimation A transcription factor regulates cold-induced gene expression Salinity Stress Salt accumulation in irrigated soils impairs plant function Plants show great diversity for salt tolerance Salt stress causes multiple injury effects Plants use multiple strategies to reduce salt stress Ion exclusion and compartmentation reduce salinity stress Plant adaptations to toxic trace elements Oxygen Deficiency Anaerobic microorganisms are active in water- saturated soils Roots are damaged in anoxic environments Damaged O2-deficient roots injure shoots Submerged organs can acquire O2 through specialized structures Most plant tissues cannot tolerate anaerobic conditions Synthesis of anaerobic stress proteins leads to acclimation to O2 deficit Summary Glossary Author Index Subject Index
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spelling	Taiz, Lincoln Verfasser aut Plant physiology Lincoln Taiz ; Eduardo Zeiger 4. ed. Sunderland, Mass. Sinauer 2006 XXVI, 764 S. zahlr. Ill., graph. Darst. txt rdacontent n rdamedia nc rdacarrier Fisiologia vegetal larpcal Fysiologie gtt Physiologie végétale Physiologie végétale rasuqam Planten gtt Plant physiology Entwicklungsphysiologie (DE-588)4152449-4 gnd rswk-swf Pflanzen (DE-588)4045539-7 gnd rswk-swf Pflanzenphysiologie (DE-588)4045580-4 gnd rswk-swf 1\p (DE-588)4123623-3 Lehrbuch gnd-content Pflanzenphysiologie (DE-588)4045580-4 s DE-604 Pflanzen (DE-588)4045539-7 s Entwicklungsphysiologie (DE-588)4152449-4 s 2\p DE-604 Zeiger, Eduardo Verfasser aut Digitalisierung UB Regensburg application/pdf http://bvbr.bib-bvb.de:8991/F?func=service&doc_library=BVB01&local_base=BVB01&doc_number=014833756&sequence=000002&line_number=0001&func_code=DB_RECORDS&service_type=MEDIA Inhaltsverzeichnis 1\p cgwrk 20201028 DE-101 https://d-nb.info/provenance/plan#cgwrk 2\p cgwrk 20201028 DE-101 https://d-nb.info/provenance/plan#cgwrk
spellingShingle	Taiz, Lincoln Zeiger, Eduardo Plant physiology Fisiologia vegetal larpcal Fysiologie gtt Physiologie végétale Physiologie végétale rasuqam Planten gtt Plant physiology Entwicklungsphysiologie (DE-588)4152449-4 gnd Pflanzen (DE-588)4045539-7 gnd Pflanzenphysiologie (DE-588)4045580-4 gnd
subject_GND	(DE-588)4152449-4 (DE-588)4045539-7 (DE-588)4045580-4 (DE-588)4123623-3
title	Plant physiology
title_auth	Plant physiology
title_exact_search	Plant physiology
title_exact_search_txtP	Plant physiology
title_full	Plant physiology Lincoln Taiz ; Eduardo Zeiger
title_fullStr	Plant physiology Lincoln Taiz ; Eduardo Zeiger
title_full_unstemmed	Plant physiology Lincoln Taiz ; Eduardo Zeiger
title_short	Plant physiology
title_sort	plant physiology
topic	Fisiologia vegetal larpcal Fysiologie gtt Physiologie végétale Physiologie végétale rasuqam Planten gtt Plant physiology Entwicklungsphysiologie (DE-588)4152449-4 gnd Pflanzen (DE-588)4045539-7 gnd Pflanzenphysiologie (DE-588)4045580-4 gnd
topic_facet	Fisiologia vegetal Fysiologie Physiologie végétale Planten Plant physiology Entwicklungsphysiologie Pflanzen Pflanzenphysiologie Lehrbuch
url	http://bvbr.bib-bvb.de:8991/F?func=service&doc_library=BVB01&local_base=BVB01&doc_number=014833756&sequence=000002&line_number=0001&func_code=DB_RECORDS&service_type=MEDIA
work_keys_str_mv	AT taizlincoln plantphysiology AT zeigereduardo plantphysiology

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