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ویرایش: 6 نویسندگان: Eduardo Zeiger, Lincoln Taiz, Ian Max Moller, Angus Murphy سری: ISBN (شابک) : 9781605352558, 2014030480 ناشر: Sinauer سال نشر: 2014 تعداد صفحات: 888 زبان: English فرمت فایل : PDF (درصورت درخواست کاربر به PDF، EPUB یا AZW3 تبدیل می شود) حجم فایل: 95 مگابایت
کلمات کلیدی مربوط به کتاب فیزیولوژی و توسعه گیاهان: کتاب درسی
در صورت تبدیل فایل کتاب Plant Physiology and Development به فرمت های PDF، EPUB، AZW3، MOBI و یا DJVU می توانید به پشتیبان اطلاع دهید تا فایل مورد نظر را تبدیل نمایند.
توجه داشته باشید کتاب فیزیولوژی و توسعه گیاهان نسخه زبان اصلی می باشد و کتاب ترجمه شده به فارسی نمی باشد. وبسایت اینترنشنال لایبرری ارائه دهنده کتاب های زبان اصلی می باشد و هیچ گونه کتاب ترجمه شده یا نوشته شده به فارسی را ارائه نمی دهد.
در طول تاریخ بیست و دو ساله خود، نویسندگان فیزیولوژی گیاهی به طور مداوم این کتاب را به روز کرده اند تا آخرین پیشرفت های زیست شناسی گیاهی را در خود جای دهد و پیشرفت های آموزشی را که توسط پذیرندگان درخواست شده است، اجرا کند. این موضوع باعث شده است که فیزیولوژی گیاهی معتبرترین، جامع ترین و پرکاربردترین کتاب درسی زیست شناسی گیاهی در بخش بالایی باشد. در ویرایش ششم، بخش رشد و توسعه (واحد III) برای ارائه چرخه زندگی کامل گیاهان بذر از جوانه زنی تا پیری سازماندهی مجدد و گسترش یافته است. با توجه به این پیشرفت، متن به فیزیولوژی و توسعه گیاهان تغییر نام داد. مانند قبل، واحد III با فصل های به روز شده در دیواره های سلولی و سیگنال ها و انتقال سیگنال آغاز می شود. فصل اخیر گسترش یافته است تا شامل بحثی در مورد مولکول های سیگنال دهنده اصلی، مانند یون های کلسیم و هورمون های گیاهی باشد. فصل جدید و یکپارچه ای با عنوان سیگنال هایی از نور خورشید جایگزین دو فصل ویرایش پنجم درباره پاسخ های نور آبی و فیتوکروم شده است. این فصل شامل فیتوکروم و همچنین گیرنده های نور آبی و UV و مسیرهای سیگنال دهی آنها از جمله فتوتروپین ها، کریپتوکروم ها و UVR8 می باشد. فصلهای بعدی در بخش III به توصیف مراحل رشد از جنینزایی تا پیری و بسیاری از عوامل فیزیولوژیکی و محیطی که آنها را تنظیم میکنند اختصاص دارد. نتیجه به دانش آموزان درک بهتری از ادغام هورمون ها و سایر عوامل سیگنال دهی در تنظیم رشد ارائه می دهد. سازمان جدید واحد III دارای مزیت اضافهای است که افزونگی را به حداقل میرساند و کاهش تعداد فصلهای واحد را از 13 به 11 ممکن میسازد. تجدید نظر ایان مکس مولر متعاقباً تمام فصول کتاب را ویرایش کرد تا از کیفیت و سطح سازگاری یکسانی اطمینان حاصل کند.
Throughout its twenty-two year history, the authors of Plant Physiology have continually updated the book to incorporate the latest advances in plant biology and implement pedagogical improvements requested by adopters. This has made Plant Physiology the most authoritative, comprehensive, and widely used upper-division plant biology textbook. In the Sixth Edition, the Growth and Development section (Unit III) has been reorganized and expanded to present the complete life cycle of seed plants from germination to senescence. In recognition of this enhancement, the text has been renamed Plant Physiology and Development. As before, Unit III begins with updated chapters on Cell Walls and Signals and Signal Transduction. The latter chapter has been expanded to include a discussion of major signaling molecules, such as calcium ions and plant hormones. A new, unified chapter entitled Signals from Sunlight has replaced the two Fifth-Edition chapters on Phytochrome and Blue Light Responses. This chapter includes phytochrome, as well as the blue and UV light receptors and their signaling pathways, including phototropins, cryptochromes, and UVR8. The subsequent chapters in Unit III are devoted to describing the stages of development from embryogenesis to senescence and the many physiological and environmental factors that regulate them. The result provides students with an improved understanding of the integration of hormones and other signaling agents in developmental regulation. The new organization of Unit III has the added benefit that it minimizes redundancy, making it possible to reduce the number of chapters in the Unit from 13 to 11. Angus Murphy of the University of Maryland has headed up a team of authors and editors to implement the revision. Ian Max Moller has subsequently edited all the book chapters to ensure an even high quality and consistency level.
Cover Half Title Title Page Copyright Page Brief Contents Editors Reviewers Preface Media and Supplements Table of Contents Chapter 1 Plant and Cell Architecture Plant Classification and Life Cycles Plant Life Processes: Unifying Principles Plant life cycles alternate between diploid and haploid generations Overview of Plant Structure Plant cells are surrounded by rigid cell walls Plasmodesmata allow the free movement of molecules between cells New cells originate in dividing tissues called meristems Biological membranes are phospholipid bilayers that contain proteins Plant Cell Organelles The Endomembrane System The nucleus contains the majority of the genetic material Gene expression involves both transcription and translation The endoplasmic reticulum is a network of internal membranes Secretion of proteins from cells begins with the rough ER Glycoproteins and polysaccharides destined for secretion are processed in the Golgi apparatus The plasma membrane has specialized regions involved in membrane recycling Vacuoles have diverse functions in plant cells Independently Dividing or Fusing Organelles Derived from the Endomembrane System Oil bodies are lipid-storing organelles Microbodies play specialized metabolic roles in leaves and seeds Independently Dividing, Semiautonomous Organelles Proplastids mature into specialized plastids in different plant tissues Chloroplast and mitochondrial division are independent of nuclear division The plant cytoskeleton consists of microtubules and microfilaments The Plant Cytoskeleton Actin, tubulin, and their polymers are in constant flux in the living cell Cortical microtubules move around the cell by treadmilling Cytoskeletal motor proteins mediate cytoplasmic streaming and directed organelle movement Cell Cycle Regulation Each phase of the cell cycle has a specific set of biochemical and cellular activities The cell cycle is regulated by cyclins and cyclin-dependent kinases Mitosis and cytokinesis involve both microtubules and the endomembrane system Plant Cell Types Dermal tissues cover the surfaces of plants Ground tissues form the bodies of plants Vascular tissues form transport networks between different parts of the plant Summary Web Material Suggested Reading Chapter 2 Genome Structure and Gene Expression Nuclear Genome Organization The nuclear genome is packaged into chromatin Centromeres, telomeres, and nucleolar organizer regions contain repetitive sequences Transposons are mobile sequences within the genome Meiosis halves the number of chromosomes and allows for the recombination of alleles Chromosome organization is not random in the interphase nucleus Polyploids contain multiple copies of the entire genome Phenotypic and physiological responses to polyploidy are unpredictable The role of polyploidy in evolution is still unclear Plant Cytoplasmic Genomes: Mitochondria and Plastids The endosymbiotic theory describes the origin of cytoplasmic genomes Organellar genetics do not obey Mendelian principles Organellar genomes vary in size RNA polymerase II binds to the promoter region of most protein-coding genes Transcriptional Regulation of Nuclear Gene Expression Conserved nucleotide sequences signal transcriptional termination and polyadenylation Epigenetic modifications help determine gene activity Noncoding RNAs regulate mRNA activity via the RNA interference (RNAi) pathway Posttranscriptional Regulation of Nuclear Gene Expression All RNA molecules are subject to decay Posttranslational regulation determines the life span of proteins Tools for Studying Gene Function Mutant analysis can help elucidate gene function Molecular techniques can measure the activity of genes Gene fusions can introduce reporter genes Genetic Modification of Crop Plants Transgenes can confer resistance to herbicides or plant pests Genetically modified organisms are controversial Plant Cytoplasmic Genomes: Mitochondria and Plastids Summary Web Material Suggested Reading Chapter 3 Water and Plant Cells Water in Plant Life The Structure and Properties of Water Water is a polar molecule that forms hydrogen bonds Water molecules are highly cohesive Water is an excellent solvent Water has distinctive thermal properties relative to its size Water has a high tensile strength Diffusion and Osmosis Diffusion is the net movement of molecules by random thermal agitation Diffusion is most effective over short distances Osmosis describes the net movement of water across a selectively permeable barrier Water Potential The chemical potential of water represents the free-energy status of water Three major factors contribute to cell water potential Water potentials can be measured Water Potential of Plant Cells Water enters the cell along a water potential gradient Water can also leave the cell in response to a water potential gradient Cell Wall and Membrane Properties Small changes in plant cell volume cause large changes in turgor pressure Water potential and its components vary with growth conditions and location within the plant The rate at which cells gain or lose water is influenced by plasma membrane hydraulic conductivity Aquaporins facilitate the movement of water across plasma membranes Plant Water Status Solute accumulation helps cells maintain turgor and volume Physiological processes are affected by plant water status Summary Web Material Suggested Reading Chapter 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 pathways Solute accumulation in the xylem can generate “root pressure” The xylem consists of two types of transport cells Water Transport through the Xylem Water moves through the xylem by pressure-driven bulk flow Water movement through the xylem requires a smaller pressure gradient than movement through living cells What pressure difference is needed to lift water 100 meters to a treetop? The cohesion–tension theory explains water transport in the xylem Xylem transport of water in trees faces physical challenges Water Movement from the Leaf to the Atmosphere Plants minimize the consequences of xylem cavitation Leaves have a large hydraulic resistance The driving force for transpiration is the difference in water vapor concentration Water loss is also regulated by the pathway resistances Stomatal control couples leaf transpiration to leaf photosynthesis The cell walls of guard cells have specialized features An increase in guard cell turgor pressure opens the stomata Overview: The Soil–Plant– Atmosphere Continuum The transpiration ratio measures the relationship between water loss and carbon gain Summary Web Material Suggested Reading Chapter 5 Mineral Nutrition 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 the addition of fertilizers Some mineral nutrients can be absorbed by leaves Negatively charged soil particles affect the adsorption of mineral nutrients Soil, Roots, and Microbes Soil pH affects nutrient availability, soil microbes, and root growth Some plants develop extensive root systems Excess mineral ions in the soil limit plant growth Root systems differ in form but are based on common structures Different areas of the root absorb different mineral ions Nutrient availability influences root growth Mycorrhizal symbioses facilitate nutrient uptake by roots Nutrients move between mycorrhizal fungi and root cells Summary Web Material Suggested Reading Chapter 6 Solute Transport Passive and Active Transport Transport of Ions across Membrane Barriers 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 Channels 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 The genes for many transporters have been identified Transporters exist for diverse nitrogen-containing compounds Cation transporters are diverse Anion transporters have been identified Aquaporins have diverse functions Transporters for metal and metalloid ions transport essential micronutrients Plasma membrane H+-ATPases are highly regulated P-type ATPases The tonoplast H+-ATPase drives solute accumulation in vacuoles 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 Web Material Suggested Reading Chapter 7 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 Photosynthesis takes place in complexes containing light-harvesting antennas and photochemical reaction centers Action spectra relate light absorption to photosynthetic activity Light drives the reduction of NADP+ and the formation of ATP The chemical reaction of photosynthesis is driven by light Oxygen-evolving organisms have two photosystems that operate in series Organization of the Photosynthetic Apparatus The chloroplast is the site of photosynthesis Thylakoids contain integral membrane proteins Photosystems I and II are spatially separated in the thylakoid membrane Anoxygenic photosynthetic bacteria have a single reaction center The antenna funnels energy to the reaction center Organization of Light-Absorbing Antenna Systems Antenna systems contain chlorophyll and are membrane-associated Many antenna pigment–protein complexes have a common structural motif Electrons from chlorophyll travel through the carriers organized in the Z scheme Mechanisms of Electron Transport Energy is captured when an excited chlorophyll reduces an electron acceptor molecule The reaction center chlorophylls of the two photosystems absorb at different wavelengths Water is oxidized to oxygen by PSII The PSII reaction center is a multi-subunit pigment–protein complex Pheophytin and two quinones accept electrons from PSII Electron flow through the cytochrome complex also transports protons Plastoquinone and plastocyanin carry electrons between photosystems II and I The PSI reaction center reduces NADP+ Cyclic electron flow generates ATP but no NADPH Proton Transport and ATP Synthesis in the Chloroplast Some herbicides block photosynthetic electron flow Repair and Regulation of the Photosynthetic Machinery Carotenoids serve as photoprotective agents Some xanthophylls also participate in energy dissipation The PSII reaction center is easily damaged PSI is protected from active oxygen species Genetics, Assembly, and Evolution of Photosynthetic Systems Thylakoid stacking permits energy partitioning between the photosystems Chloroplast genes exhibit non-Mendelian patterns of inheritance Most chloroplast proteins are imported from the cytoplasm Complex photosynthetic organisms have evolved from simpler forms The biosynthesis and breakdown of chlorophyll are complex pathways Mechanisms of Electron Transport Summary Web Material Suggested Reading Chapter 8 Photosynthesis: The Carbon Reactions The Calvin–Benson Cycle The Calvin–Benson cycle has three phases: carboxylation, reduction, and regeneration The fixation of CO2 via carboxylation of ribulose 1,5-bisphosphate and the reduction of the product 3-phosphoglycerate yield triose phosphates The regeneration of ribulose 1,5-bisphosphate ensures the continuous assimilation of CO2 An induction period precedes the steady state of photosynthetic CO2 assimilation Many mechanisms regulate the Calvin–Benson cycle Rubisco-activase regulates the catalytic activity of rubisco Light regulates the Calvin–Benson cycle via the ferredoxin–thioredoxin system Light-dependent ion movements modulate enzymes of the Calvin–Benson cycle Light controls the assembly of chloroplast enzymes into supramolecular complexes The C2 Oxidative Photosynthetic Carbon Cycle The oxygenation of ribulose 1,5-bisphosphate sets in motion the C2 oxidative photosynthetic carbon cycle Enzymes of the plant C2 oxidative photosynthetic carbon cycle derive from different ancestors Photorespiration is linked to the photosynthetic electron transport system Cyanobacteria use a proteobacterial pathway for bringing carbon atoms of 2-phosphoglycolate back to the Calvin–Benson cycle The C2 oxidative photosynthetic carbon cycle interacts with many metabolic pathways Production of biomass may be enhanced by engineering photorespiration Inorganic Carbon–Concentrating Mechanisms Inorganic Carbon–Concentrating Mechanisms: The C4 Carbon Cycle Malate and aspartate are the primary carboxylation products of the C4 cycle The C4 cycle assimilates CO2 by the concerted action of two different types of cells The C4 cycle uses different mechanisms for decarboxylation of four-carbon acids transported to bundle sheath cells Bundle sheath cells and mesophyll cells exhibit anatomical and biochemical differences The C4 cycle also concentrates CO2 in single cells Light regulates the activity of key C4 enzymes Photosynthetic assimilation of CO2 in C4 plants demands more transport processes than in C3 plants In hot, dry climates, the C4 cycle reduces photorespiration Inorganic Carbon–Concentrating Mechanisms: Crassulacean Acid Metabolism (CAM) Different mechanisms regulate C4 PEPCase and CAM PEPCase CAM is a versatile mechanism sensitive to environmental stimuli Accumulation and Partitioning of Photosynthates—Starch and Sucrose Formation and Mobilization of Chloroplast Starch Chloroplast stroma accumulates starch as insoluble granules during the day Starch degradation at night requires the phosphorylation of amylopectin The export of maltose prevails in the nocturnal breakdown of transitory starch The synthesis and degradation of the starch granule are regulated by multiple mechanisms Sucrose Biosynthesis and Signaling Triose phosphates from the Calvin–Benson cycle build up the cytosolic pool of three important hexose phosphates in the light Sucrose is continuously synthesized in the cytosol Fructose 2,6-bisphosphate regulates the hexose phosphate pool in the light Summary Web Material Suggested Reading Chapter 9 Photosynthesis: Physiological and Ecological Considerations Photosynthesis Is Influenced by Leaf Properties Leaf anatomy and canopy structure maximize light absorption Leaf angle and leaf movement can control light absorption Leaves acclimate to sun and shade environments Effects of Light on Photosynthesis in the Intact Leaf Light-response curves reveal photosynthetic properties Leaves must dissipate excess light energy Absorption of too much light can lead to photoinhibition Effects of Temperature on Photosynthesis in the Intact Leaf Leaves must dissipate vast quantities of heat There is an optimal temperature for photosynthesis Photosynthesis is sensitive to both high and low temperatures Photosynthetic efficiency is temperature-sensitive Effects of Carbon Dioxide on Photosynthesis in the Intact Leaf Atmospheric CO2 concentration keeps rising CO2 diffusion to the chloroplast is essential to photosynthesis CO2 imposes limitations on photosynthesis How will photosynthesis and respiration change in the future under elevated CO2 conditions? Stable Isotopes Record Photosynthetic Properties How do we measure the stable carbon isotopes of plants? Why are there carbon isotope ratio variations in plants? Summary Web Material Suggested Reading Chapter 10 Stomatal Biology Light-dependent Stomatal Opening Guard cells respond to blue light Blue light activates a proton pump at the guard cell plasma membrane Blue light regulates the osmotic balance of guard cells Blue-light responses have characteristic kinetics and lag times Sucrose is an osmotically active solute in guard cells Mediation of Blue-light Photoreception in Guard Cells by Zeaxanthin Reversal of Blue Light–Stimulated Opening by Green Light A carotenoid–protein complex senses light intensity The Resolving Power of Photophysiology Mediation of Blue-light Photoreception in Guard Cells by Zeaxanthin Summary Web Material Suggested Reading Chapter 11 Translocation in the Phloem Pathways of Translocation 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 Translocation: Source to Sink Phloem sap can be collected and analyzed Materials Translocated in the Phloem Sugars are translocated in a nonreducing form Other solutes are translocated in the phloem Rates of Movement An osmotically generated pressure gradient drives translocation in the pressure-flow model The Pressure-Flow Model, a Passive Mechanism for Phloem Transport Some predictions of pressure flow have been confirmed, while others require further experimentation The energy requirement for transport through the phloem pathway is small in herbaceous plants There is no bidirectional transport in single sieve elements, and solutes and water move at the same velocity Sieve plate pores appear to be open channels Pressure gradients in the sieve elements may be modest; pressures in herbaceous plants and trees appear to be similar Alternative models for translocation by mass flow have been suggested Does translocation in gymnosperms involve a different mechanism? Phloem Loading Phloem loading can occur via the apoplast or symplast Abundant data support the existence of apoplastic loading in some species Sucrose uptake in the apoplastic pathway requires metabolic energy Phloem loading is symplastic in some species Phloem loading in the apoplastic pathway involves a sucrose–H+ symporter The polymer-trapping model explains symplastic loading in plants with intermediary-type companion cells The type of phloem loading is correlated with several significant characteristics Phloem loading is passive in several tree species 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 Various sinks partition transport sugars Allocation includes storage, utilization, and transport Source leaves regulate allocation Sink tissues compete for available translocated photosynthate The source adjusts over the long term to changes in the source-to-sink ratio Sink strength depends on sink size and activity Transport of Signaling Molecules Turgor pressure and chemical signals coordinate source and sink activities Proteins and RNAs function as signal molecules in the phloem to regulate growth and development Plasmodesmata function in phloem signaling Summary Web Material Suggested Reading Chapter 12 Respiration and Lipid Metabolism Overview of Plant Respiration Glycolysis Glycolysis metabolizes carbohydrates from several sources Plants have alternative glycolytic reactions The energy-conserving phase of glycolysis extracts usable energy In the absence of oxygen, fermentation regenerates the NAD+ needed for glycolysis Plant glycolysis is controlled by its products The Oxidative Pentose Phosphate Pathway The oxidative pentose phosphate pathway is redox-regulated The oxidative pentose phosphate pathway produces NADPH and biosynthetic intermediates The Citric Acid Cycle Mitochondria are semiautonomous organelles Pyruvate enters the mitochondrion and is oxidized via the citric acid cycle Mitochondrial Electron Transport and ATP Synthesis The citric acid cycle of plants has unique features The electron transport chain catalyzes a flow of electrons from NADH to O2 The electron transport chain has supplementary branches ATP synthesis in the mitochondrion is coupled to electron transport Aerobic respiration yields about 60 molecules of ATP per molecule of sucrose Transporters exchange substrates and products Several subunits of respiratory complexes are encoded by the mitochondrial genome Plants have several mechanisms that lower the ATP yield Short-term control of mitochondrial respiration occurs at different levels Respiration is tightly coupled to other pathways Plants respire roughly half of the daily photosynthetic yield Respiration in Intact Plants and Tissues Different tissues and organs respire at different rates Respiration operates during photosynthesis Environmental factors alter respiration rates Triacylglycerols are stored in oil bodies Lipid Metabolism Fats and oils store large amounts of energy 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 ER Membrane lipids are precursors of important signaling compounds Lipid composition influences membrane function Storage lipids are converted into carbohydrates in germinating seeds Summary Web Material Suggested Reading Chapter 13 Assimilation of Inorganic 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 Converting ammonium to amino acids requires two enzymes Ammonium Assimilation Ammonium can be assimilated via an alternative pathway Transamination reactions transfer nitrogen Amino Acid Biosynthesis Asparagine and glutamine link carbon and nitrogen metabolism Biological Nitrogen Fixation Free-living and symbiotic bacteria fix nitrogen Nitrogen fixation requires microanaerobic or 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 N2 Amides and ureides are the transported forms of nitrogen Sulfur Assimilation Sulfate assimilation requires the reduction of sulfate to cysteine Sulfate is the form of sulfur transported into plants Methionine is synthesized from cysteine Phosphate Assimilation Sulfate assimilation occurs mostly in leaves Cations form noncovalent bonds with carbon compounds Cation Assimilation Roots modify the rhizosphere to acquire iron Oxygen Assimilation Iron cations form complexes with carbon and phosphate The Energetics of Nutrient Assimilation Summary Web Material Suggested Reading Chapter 14 Cell Walls: Structure, Formation, and Expansion Overview of Plant Cell Wall Functions and Structures Plant cell walls vary in structure and function Components differ for primary and secondary cell walls Cellulose microfibrils have an ordered structure and are synthesized at the plasma membrane Matrix polymers are synthesized in the Golgi apparatus and secreted via vesicles Pectins are hydrophilic gel-forming components of the primary cell wall Hemicelluloses are matrix polysaccharides that bind to cellulose New primary cell walls are assembled during cytokinesis and continue to be assembled during growth Primary Cell Wall Structure and Function The primary cell wall is composed of cellulose microfibrils embedded in a matrix of pectins and hemicelluloses Mechanisms of Cell Expansion Microfibril orientation influences growth directionality of cells with diffuse growth Cortical microtubules influence the orientation of newly deposited microfibrils The Extent and Rate of Cell Growth Stress relaxation of the cell wall drives water uptake and cell expansion Acid-induced growth and wall stress relaxation are mediated by expansins Cell wall models are hypotheses about how molecular components fit together to make a functional wall Secondary Cell Wall Structure and Function Many structural changes accompany the cessation of wall expansion Secondary cell walls are rich in cellulose and hemicellulose and often have a hierarchical organization Lignification transforms the SCW into a hydrophobic structure resistant to deconstruction Summary Web Material Suggested Reading Chapter 15 Signals and Signal Transduction Temporal and Spatial Aspects of Signaling Signal Perception and Amplification Receptors are located throughout the cell and are conserved across kingdoms Signals must be amplified intracellularly to regulate their target molecules The MAP kinase signal amplification cascade is present in all eukaryotes Ca2+ is the most ubiquitous second messenger in plants and other eukaryotes Changes in the cytosolic or cell wall pH can serve as second messengers for hormonal and stress responses Reactive oxygen species act as second messengers mediating both environmental and developmental signals Hormones and Plant Development Lipid signaling molecules act as second messengers that regulate a variety of cellular processes Gibberellins promote stem growth and were discovered in relation to the “foolish seedling disease” of rice Auxin was discovered in early studies of coleoptile bending during phototropism Cytokinins were discovered as cell division– promoting factors in tissue culture experiments Ethylene is a gaseous hormone that promotes fruit ripening and other developmental processes Abscisic acid regulates seed maturation and stomatal closure in response to water stress Brassinosteroids regulate photomorphogenesis, germination, and other developmental processes Strigolactones suppress branching and promote rhizosphere interactions Phytohormone Metabolism and Homeostasis Indole-3-pyruvate is the primary intermediate in auxin biosynthesis Gibberellins are synthesized by oxidation of the diterpene ent-kaurene Cytokinins are adenine derivatives with isoprene side chains Abscisic acid is synthesized from a carotenoid intermediate Ethylene is synthesized from methionine via the intermediate ACC Brassinosteroids are derived from the sterol campesterol Signal Transmission and Cell–Cell Communication Strigolactones are synthesized from beta-carotene The cytokinin and ethylene signal transduction pathways are derived from the bacterial two-component regulatory system Hormonal Signaling Pathways Receptor-like kinases mediate brassinosteroid and certain auxin signaling pathways The core ABA signaling components include phosphatases and kinases Plant hormone signaling pathways generally employ negative regulation Several plant hormone receptors encode components of the ubiquitination machinery and mediate signaling via protein degradation Plants have evolved mechanisms for switching off or attenuating signaling responses The cellular response output to a signal is often tissue-specific Cross-regulation allows signal transduction pathways to be integrated Summary Suggested Reading Chapter 16 Signals from Sunlight Plant Photoreceptors Photoresponses are driven by light quality or spectral properties of the energy absorbed Plants responses to light can be distinguished by the amount of light required Phytochrome is the primary photoreceptor for red and far-red light Phytochromes Phytochrome can interconvert between Pr and Pfr forms The phytochrome chromophore and protein both undergo conformational changes in response to red light Pfr is the physiologically active form of phytochrome Pfr is partitioned between the cytosol and the nucleus Phytochrome Responses Phytochrome responses fall into three main categories based on the amount of light required Phytochrome responses vary in lag time and escape time Phytochrome A mediates responses to continuous far-red light Roles for phytochromes C, D, and E are emerging Phytochrome Signaling Pathways Phytochrome B mediates responses to continuous red or white light Phytochrome regulates membrane potentials and ion fluxes Phytochrome interacting factors (PIFs) act early in signaling Phytochrome regulates gene expression Phytochrome signaling involves protein phosphorylation and dephosphorylation Phytochrome-induced photomorphogenesis involves protein degradation Blue-Light Responses and Photoreceptors Blue-light responses have characteristic kinetics and lag times Cryptochromes The activated FAD chromophore of cryptochrome causes a conformational change in the protein Nuclear cryptochromes inhibit COP1-induced protein degradation cry1 and cry2 have different developmental effects Cryptochrome can also bind to transcriptional regulators directly The Coaction of Cryptochrome, Phytochrome, and Phototropins Stem elongation is inhibited by both red and blue photoreceptors The circadian clock is regulated by multiple aspects of light Phototropins Phytochrome interacts with cryptochrome to regulate flowering Blue light induces changes in FMN absorption maxima associated with conformation changes The LOV2 domain is primarily responsible for kinase activation in response to blue light Blue light induces a conformational change that “uncages” the kinase domain of phototropin and leads to autophosphorylation Phototropism requires changes in auxin mobilization Phototropins regulate chloroplast movements via F-actin filament assembly Stomatal opening is regulated by blue light, which activates the plasma membrane H+-ATPase The main signal transduction events of phototropin-mediated stomatal opening have been identified Responses to Ultraviolet Radiation Summary Web Material Suggested Reading Chapter 17 Embryogenesis Overview of Plant Growth and Development Sporophytic development can be divided into three major stages Embryogenesis differs between eudicots and monocots, but also features common fundamental processes Embryogenesis: The Origins of Polarity Apical–basal polarity is established early in embryogenesis Position-dependent mechanisms guide embryogenesis Intercellular signaling processes play key roles in guiding position-dependent development Embryo development features regulated communication between cells The analysis of mutants identifies genes for signaling processes that are essential for embryo organization Auxin functions as a mobile chemical signal during embryogenesis Plant polarity is maintained by polar auxin streams Auxin transport is regulated by multiple mechanisms The GNOM protein establishes a polar distribution of PIN auxin efflux proteins Radial patterning guides formation of tissue layers MONOPTEROS encodes a transcription factor that is activated by auxin The origin of epidermis: a boundary and interface at the edge of the radial axis Procambial precusors for the vascular stele lie at the center of the radial axis The differentiation of cortical and endodermal cells involves the intercellular movement of a transcription factor Meristematic Tissues: Foundations for Indeterminate Growth The root and shoot apical meristems use similar strategies to enable indeterminate growth The Root Apical Meristem The origin of different root tissues can be traced to specific initial cells The root tip has four developmental zones Responses to auxin are mediated by several distinct families of transcription factors Cell ablation experiments implicate directional signaling processes in determination of cell identity Auxin contributes to the formation and maintenance of the RAM Cytokinin is required for normal root development The Shoot Apical Meristem The shoot apical meristem has distinct zones and layers Shoot tissues are derived from several discrete sets of apical initials Factors involved in auxin movement and responses influence SAM formation Embryonic SAM formation requires the coordinated expression of transcription factors A combination of positive and negative interactions determines apical meristem size class homeodomain genes help maintain the proliferative ability of the SAM through regulation of cytokinin and GA levels Localized zones of auxin accumulation promote leaf initiation The Vascular Cambium The maintenance of undetermined initials in various meristem types depends on similar mechanisms Summary Web Material Suggested Reading Chapter 18 Seed Dormancy, Germination, and Seedling Establishment Seed Structure Seed anatomy varies widely among different plant groups Seed Dormancy Dormancy can be imposed on the embryo by the surrounding tissues Embryo dormancy may be caused by physiological or morphological factors Non-dormant seeds can exhibit vivipary and precocious germination The ABA:GA ratio is the primary determinant of seed dormancy Release from Dormancy Some seeds require either chilling or after-ripening to break dormancy Light is an important signal that breaks dormancy in small seeds Seed Germination Germination can be divided into three phases corresponding to the phases of water uptake Seed dormancy can by broken by various chemical compounds The cereal aleurone layer is a specialized digestive tissue surrounding the starchy endosperm Mobilization of Stored Reserves Gibberellins enhance the transcription of mRNA The gibberellin receptor, GID1, promotes the degradation of negative regulators of the gibberellin response DELLA repressor proteins are rapidly degraded GA-MYB is a positive regulator of amylase transcription ABA inhibits gibberellin-induced enzyme production Seedling Growth and Establishment The outer tissues of eudicot stems are the targets of auxin action The minimum lag time for auxin-induced elongation is 10 minutes Auxin promotes growth in stems and coleoptiles, while inhibiting growth in roots Auxin-induced proton extrusion induces cell wall creep and cell elongation Gravitropism involves the lateral redistribution of auxin Tropisms: Growth in Response to Directional Stimuli Polar auxin transport requires energy and is gravity independent According to the starch–statolith hypothesis, specialized amyloplasts serve as gravity sensors in root caps Auxin movements in the root are regulated by specific transporters The gravitropic stimulus perturbs the symmetric movement of auxin from the root tip Gravity perception in eudicot stems and stemlike organs occurs in the starch sheath Gravity sensing may involve pH and calcium ions (Ca2+) as second messengers Phototropism Phototropism is mediated by the lateral redistribution of auxin Phototropism occurs in a series of posttranslational events Photomorphogenesis Gibberellins and brassinosteroids both suppress photomorphogenesis in the dark Hook opening is regulated by phytochrome and auxin Ethylene induces lateral cell expansion Shade Avoidance Decreasing the R:FR ratio causes elongation in sun plants Phytochrome enables plants to adapt to changes in light quality Reducing shade avoidance responses can improve crop yields Vascular Tissue Differentiation Auxin and cytokinin are required for normal vascular development Zinnia suspension-cultured cells can be induced to undergo xylogenesis Xylogenesis involves chemical signaling between neighboring cells Root Growth and Differentiation Root epidermal development follows three basic patterns Auxin and other hormones regulate root hair development Lateral root formation and emergence depend on endogenous and exogenous signals Regions of lateral root emergence correspond with regions of auxin maxima Lateral roots and shoots have gravitropic setpoint angles Summary Web Material Suggested Reading Chapter 19 Vegetative Growth and Organogenesis Leaf Development The Establishment of Leaf Polarity Hormonal signals play key roles in regulating leaf primordia emergence A signal from the SAM initiates adaxial–abaxial polarity Adaxial leaf development requires HD-ZIP III transcription factors ARP genes promote adaxial identity and repress the KNOX1 gene The expression of HD-ZIP III genes is antagonized by miR166 in abaxial regions of the leaf Interactions between adaxial and abaxial tissues are required for blade outgrowth Antagonism between KANADI and HD-ZIP III is a key determinant of adaxial–abaxial leaf polarity Blade outgrowth is auxin dependent and regulated by the YABBY and WOX genes Leaf proximal–distal polarity also depends on specific gene expression In compound leaves, de-repression of the KNOX1 gene promotes leaflet formation Differentiation of Epidermal Cell Types Guard cell fate is ultimately determined by a specialized epidermal lineage Two groups of bHLH transcription factors govern stomatal cell fate transitions Peptide signals regulate stomatal patterning by interacting with cell surface receptors Genetic screens have led to the identification of positive and negative regulators of trichome initiation GLABRA2 acts downstream of the GL1–GL3–TTG1 complex to promote trichome formation Venation Patterns in Leaves Jasmonic acid regulates Arabidopsis leaf trichome development Auxin canalization initiates development of the leaf trace The primary leaf vein is initiated discontinuously from the preexisting vascular system Basipetal auxin transport from the L1 layer of the leaf primordium initiates development of the leaf trace procambium The existing vasculature guides the growth of the leaf trace Higher-order leaf veins differentiate in a predictable hierarchical order Auxin canalization regulates higher-order vein formation Localized auxin biosynthesis is critical for higher-order venation patterns Shoot Branching and Architecture Axillary meristem initiation involves many of the same genes as leaf initiation and lamina outgrowth Auxin, cytokinins, and strigolactones regulate axillary bud outgrowth Strigolactones act locally to repress axillary bud growth Auxin from the shoot tip maintains apical dominance Cytokinins antagonize the effects of strigolactones The initial signal for axillary bud growth may be an increase in sucrose availability to the bud Integration of environmental and hormonal branching signals is required for plant fitness Axillary bud dormancy in woody plants is affected by season, position, and age factors Plants can modify their root system architecture to optimize water and nutrient uptake Root System Architecture Monocots and eudicots differ in their root system architecture Root system architecture changes in response to phosphorous deficiencies Root system architecture responses to phosphorus deficiency involve both local and systemic regulatory networks Mycorrhizal networks augment root system architecture in all major terrestrial ecosystems Secondary Growth The vascular cambium and cork cambium are the secondary meristems where secondary growth originates Phytohormones have important roles in regulating vascular cambium activity and differentiation of secondary xylem and phloem Secondary growth evolved early in the evolution of land plants Secondary growth from the vascular cambium gives rise to secondary xylem and phloem Genes involved in stem cell maintenance, proliferation, and differentiation regulate secondary growth Environmental factors influence vascular cambium activity and wood properties Summary Web Material Suggested Reading Chapter 20 The Control of Flowering and Floral Development Floral Evocation: Integrating Environmental Cues The Shoot Apex and Phase Changes Plant development has three 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 signals Circadian Rhythms: The Clock Within Circadian rhythms exhibit characteristic features Phytochromes and cryptochromes entrain the clock Phase shifting adjusts circadian rhythms to different day–night cycles Photoperiodism: Monitoring Day Length Plants can be classified according to their photoperiodic responses Night breaks can cancel the effect of the dark period The leaf is the site of perception of the photoperiodic signal Photoperiodic timekeeping during the night depends on a circadian clock Plants monitor day length by measuring the length of the night The coincidence model is based on oscillating light sensitivity The coincidence of CONSTANS expression and light promotes flowering in LDPs Phytochrome is the primary photoreceptor in photoperiodism SDPs use a coincidence mechanism to inhibit flowering in long days 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 can involve epigenetic changes in gene expression A range of vernalization pathways may have evolved Long-distance Signaling Involved in Flowering Grafting studies provided the first evidence for a transmissible floral stimulus Florigen is translocated in the phloem The Identification of Florigen The Arabidopsis protein FLOWERING LOCUS T (FT) is florigen Gibberellins and ethylene can induce flowering The transition to flowering involves multiple factors and pathways Floral Meristems and Floral Organ Development The four different types of floral organs are initiated as separate whorls The shoot apical meristem in Arabidopsis changes with development Two major categories of genes regulate floral development Floral meristem identity genes regulate meristem function Homeotic mutations led to the identification of floral organ identity genes The ABC model partially explains the determination of floral organ identity Arabidopsis Class E genes are required for the activities of the A, B, and C genes According to the Quartet Model, floral organ identity is regulated by tetrameric complexes of the ABCE proteins Class D genes are required for ovule formation Floral asymmetry in flowers is regulated by gene expression Summary Web Material Suggested Reading Chapter 21 Gametophytes, Pollination, Seeds, and Fruits Development of the Male and Female Gametophyte Generations Formation of Male Gametophytes in the Stamen Pollen grain formation occurs in two successive stages The multilayered pollen cell wall is surprisingly complex Female Gametophyte Development in the Ovule The Arabidopsis gynoecium is an important model system for studying ovule development The vast majority of angiosperms exhibit Polygonum-type embryo sac development Functional megaspores undergo a series of free nuclear mitotic divisions followed by cellularization Embryo sac development involves hormonal signaling between sporophytic and gametophytic generations Pollination and Fertilization in Flowering Plants Delivery of sperm cells to the female gametophyte by the pollen tube occurs in six phases Adhesion and hydration of a pollen grain on a compatible flower depend on recognition between pollen and stigma surfaces Ca2+-triggered polarization of the pollen grain precedes tube formation Pollen tubes grow by tip growth Receptor-like kinases are thought to regulate the ROP1 GTPase switch, a master regulator of tip growth Style tissue conditions the pollen tube to respond to attractants produced by the synergids of the embryo sac Pollen tube tip growth in the pistil is directed by both physical and chemical cues Double fertilization occurs in three distinct stages Selfing versus Outcrossing Hermaphroditic and monoecious species have evolved floral features to ensure outcrossing Self-incompatibility (SI) is the primary mechanism that enforces outcrossing in angiosperms Cytoplasmic male sterility (CMS) occurs in the wild and is of great utility in agriculture The Brassicaceae sporophytic SI system requires two S-locus genes Gametophytic self-incompatibility (GSI) is mediated by cytotoxic S-RNases and F-box proteins Apomixis: Asexual Reproduction by Seed Apomixis is not an evolutionary dead end Endosperm Development Cellularization of coenocytic endosperm in Arabidopsis progresses from the micropylar to the chalazal region Endosperm development and embryogenesis can occur autonomously Cellularization of the coenocytic endosperm of cereals progresses centripetally Many of the genes that control endosperm development are maternally expressed genes The FIS proteins are members of a Polycomb repressive complex (PRC2) that represses endosperm development Two genes, DEK1 and CR4, have been implicated in aleurone layer differentiation Cells of the starchy endosperm and aleurone layer follow divergent developmental pathways Seed Coat Development Seed coat development appears to be regulated by the endosperm Seed Maturation and Desiccation Tolerance Seed filling and desiccation tolerance phases overlap in most species LEA proteins and nonreducing sugars have been implicated in seed desiccation tolerance The acquisition of desiccation tolerance involves many metabolic pathways During the acquisition of desiccation tolerance, the cells of the embryo acquire a glassy state Specific LEA proteins have been implicated in desiccation tolerance in Coat-imposed dormancy is correlated with longterm seed-viability Abscisic acid plays a key role in seed maturation Fruit Development and Ripening Arabidopsis and tomato are model systems for the study of fruit development Fleshy fruits undergo ripening Ripening involves changes in the color of fruit The causal link between ethylene and ripening was demonstrated in transgenic and mutant tomatoes Fruit softening involves the coordinated action of many cell wall–degrading enzymes Taste and flavor reflect changes in acids, sugars, and aroma compounds Climacteric and non-climacteric fruit differ in their ethylene responses Angiosperms share a range of common molecular mechanisms controlling fruit development and ripening The ripening process is transcriptionally regulated Fruit ripening is under epigenetic control A mechanistic understanding of the ripening process has commercial applications Summary Web Material Suggested Reading Chapter 22 Plant Senescence and Cell Death Programmed Cell Death and Autolysis PCD during normal development differs from that of the hypersensitive response The autophagy pathway captures and degrades cellular constituents within lytic compartments A subset of the autophagy-related genes controls the formation of the autophagosome The autophagy pathway plays a dual role in plant development The Leaf Senescence Syndrome The developmental age of a leaf may differ from its chronological age Leaf senescence may be sequential, seasonal, or stress-induced Developmental leaf senescence consists of three distinct phases The earliest cellular changes during leaf senescence occur in the chloroplast The autolysis of chloroplast proteins occurs in multiple compartments The STAY-GREEN (SGR) protein is required for both LHCP II protein recycling and chlorophyll catabolism Leaf senescence is preceded by a massive reprogramming of gene expression The NAC and WRKY gene families are the most abundant transcription factors regulating leaf senescence Leaf Senescence: The Regulatory Network ROS serve as internal signaling agents in leaf senescence Sugars accumulate during leaf senescence and may serve as a signal Plant hormones interact in the regulation of leaf senescence Leaf Abscission The timing of leaf abscission is regulated by the interaction of ethylene and auxin Whole Plant Senescence Angiosperm life cycles may be annual, biennial, or perennial Whole plant senescence differs from aging in animals The determinacy of shoot apical meristems is developmentally regulated Nutrient or hormonal redistribution may trigger senescence in monocarpic plants The rate of carbon accumulation in trees increases continuously with tree size Summary Web Material Suggested Reading Chapter 23 Biotic Interactions Beneficial Interactions between Plants and Microorganisms Arbuscular mycorrhizal associations and nitrogen-fixing symbioses involve related signaling pathways Nod factors are recognized by the Nod factor receptor (NFR) in legumes Rhizobacteria can increase nutrient availability, stimulate root branching, and protect against pathogens Harmful Interactions between Plants, Pathogens, and Herbivores Mechanical barriers provide a first line of defense against insect pests and pathogens Plant secondary metabolites can deter insect herbivores Plants store constitutive toxic compounds in specialized structures Plants often store defensive chemicals as nontoxic water-soluble sugar conjugates in the vacuole Inducible Defense Responses to Insect Herbivores Constitutive levels of secondary compounds are higher in young developing leaves than in older tissues Modified fatty acids secreted by grasshoppers act as elicitors of jasmonic acid accumulation and ethylene emission Plants can recognize specific components of insect saliva Phloem feeders activate defense signaling pathways similar to those activated by pathogen infections Calcium signaling and activation of the MAP kinase pathway are early events associated with insect herbivory Jasmonic acid activates defense responses against insect herbivores Jasmonic acid acts through a conserved ubiquitin ligase signaling mechanism Hormonal interactions contribute to plant–insect herbivore interactions JA initiates the production of defense proteins that inhibit herbivore digestion Herbivore damage induces systemic defenses Glutamate receptor-like (GLR) genes are required for long-distance electrical signaling during herbivory Herbivore-induced volatiles can repel herbivores and attract natural enemies Herbivore-induced volatiles can serve as longdistance signals between plants Defense responses to herbivores and pathogens are regulated by circadian rhythms Herbivore-induced volatiles can also act as systemic signals within a plant Plant Defenses against Pathogens Insects have evolved mechanisms to defeat plant defenses Microbial pathogens have evolved various strategies to invade host plants Pathogens produce effector molecules that aid in the colonization of their plant host cells Pathogen infection can give rise to molecular “danger signals” that are perceived by cell surface pattern recognition receptors R genes provide resistance to individual pathogens by recognizing strain-specific effectors Effectors released by phloem-feeding insects also activate NBS–LRR receptors Exposure to elicitors induces a signal transduction cascade The hypersensitive response is a common defense against pathogens Phytoalexins with antimicrobial activity accumulate after pathogen attack A single encounter with a pathogen may increase resistance to future attacks The main components of the salicylic acid signaling pathway for SAR have been identified Interactions of plants with nonpathogenic bacteria can trigger systemic resistance through a process called induced systemic res Plant Defenses against Other Organisms Some plant parasitic nematodes form specific associations through the formation of distinct feeding structures Plants compete with other plants by secreting allelopathic secondary metabolites into the soil Some plants are biotrophic pathogens of other plants Summary Web Material Suggested Reading Chapter 24 Abiotic Stress Defining Plant Stress Physiological adjustment to abiotic stress involves trade-offs between vegetative and reproductive development Adaptation to stress involves genetic modification over many generations Acclimation allows plants to respond to environmental fluctuations Acclimation and Adaptation Environmental Factors and Their Biological Impacts on Plants Water deficit decreases turgor pressure, increases ion toxicity, and inhibits photosynthesis Light stress can occur when shade-adapted or shade-acclimated plants are subjected to full sunlight Salinity stress has both osmotic and cytotoxic effects Temperature stress affects a broad spectrum of physiological processes Heavy metals can both mimic essential mineral nutrients and generate ROS Flooding results in anaerobic stress to the root Mineral nutrient deficiencies are a cause of stress During freezing stress, extracellular ice crystal formation causes cell dehydration Ozone and ultraviolet light generate ROS that cause lesions and induce PCD Combinations of abiotic stresses can induce unique signaling and metabolic pathways Stress-Sensing Mechanisms in Plants Sequential exposure to different abiotic stresses sometimes confers cross-protection Signaling Pathways Activated in Response to Abiotic Stress Early-acting stress sensors provide the initial signal for the stress response The signaling intermediates of many stress-response pathways can interact Acclimation to stress involves transcriptional regulatory networks called Chloroplast genes respond to high-intensity light by sending stress signals to the nucleus A self-propagating wave of ROS mediates systemic acquired acclimation Hormonal interactions regulate normal development and abiotic stress responses Epigenetic mechanisms and small RNAs provide additional protection against stress Developmental and Physiological Mechanisms That Protect Plants against Abiotic Stress Plants adjust osmotically to drying soil by accumulating solutes Submerged organs develop aerenchyma tissue in response to hypoxia Antioxidants and ROS-scavenging pathways protect cells from oxidative stress Molecular chaperones and molecular shields protect proteins and membranes during abiotic stress Plants can alter their membrane lipids in response to temperature and other abiotic stresses Exclusion and internal tolerance mechanisms allow plants to cope with toxic ions Phytochelatins and other chelators contribute to internal tolerance of toxic metal ions Plants use cryoprotectant molecules and antifreeze proteins to prevent ice crystal formation ABA signaling during water stress causes the massive efflux of K+ and anions from guard cells Plants can alter their morphology in response to abiotic stress Metabolic shifts enable plants to cope with a variety of abiotic stresses Developing crops with enhanced tolerance to abiotic stress conditions is a major goal of agricultural research The process of recovery from stress can be dangerous to the plant and requires a coordinated adjustment of plant metabolism and Summary Web Material Suggested Reading Glossary Illustration Credits Photo Credits Subject Index Blank Page