Fundamentals of Plant Physiology
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Fundamentals of Plant Physiology emphasizes essential concepts, offering a concise, accessible, and focused approach to the topic while maintaining a high standard of scientific accuracy and pedagogical richness for which the Taiz franchise is renowned. Fundamentals of Plant Physiology is a concise and accessible textbook that meets the needs of undergraduate students studying plant physiology and structure/function classes.
This fundamentals version is a response to requests from educators for an up-to-date and accessible text, specifically tailored for students who may not have extensive training in organic chemistry, genetics, or molecular biology. The second edition has been updated to reflect the remarkable advancements in plant sciences. These advancements, coupled with rapid progress in molecular breeding genome editing techniques, have enabled the development of crops that can sustain various challenges, including climate change, soil degradation, and the introduction of new pathogens and pests.
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- Oxford University Press Academic US
- 9780197614181
- 9780197614167
- ePub
- 2
- Lincoln Taiz
- English
- 2024-04-23
- 100
- 2
- 2
Kaflar
- Cover Page
- Title page
- Copyright page
- About the Cover
- Table of Contents
- Preface
- Editors
- Digital Resources: to accompany Fundamentals of Plant Physiology 2e
- Enhanced E-Book for the Student (ISBN 9780197614181)
- For the Instructor
- 1 Plant and Cell Architecture
- 1.1 Plant Life Processes: Unifying Principles
- Plant life cycles alternate between diploid and haploid generations
- 1.2 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
- 1.3 Plant Tissue 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
- 1.4 Plant Cell Compartments
- Biological membranes are lipid bilayers that contain proteins
- Lipids
- Proteins
- 1.5 The Plant Cytoskeleton
- The plant cytoskeleton consists of microtubules and microfilaments
- Actin, tubulin, and their polymers are in constant flux in the living cell
- Cytoskeletal motor proteins mediate cytoplasmic streaming and directed organelle movement
- 1.6 The Nucleus
- Gene expression involves transcription, translation, and protein processing
- Posttranslational modification of proteins determines their location, activity, and longevity
- 1.7 The Endomembrane System
- The endoplasmic reticulum is a network of internal membranes
- Cell wall matrix polysaccharides, secretory proteins, and glycoproteins are processed in the Golgi apparatus
- The plasma membrane has specialized regions involved in membrane recycling
- Vacuoles have diverse functions in plant cells
- Oil bodies are lipid-storing organelles
- Peroxisomes play specialized metabolic roles in leaves and seeds
- 1.8 Independently Dividing Semiautonomous Organelles
- Plastidial and mitochondrial division are independent of nuclear division in land plants
- 1.9 Cell Cycle Regulation
- Each phase of the cell cycle has a specific set of biochemical and cellular activities
- Mitosis and cytokinesis involve both microtubules and the endomembrane system
- Summary
- 1.1 Plant Life Processes: Unifying Principles
- 1.2 Overview of Plant Structure
- 1.3 Plant Tissue Types
- 1.4 Plant Cell Compartments
- 1.5 The Plant Cytoskeleton
- 1.6 The Nucleus
- 1.7 The Endomembrane System
- 1.8 Independently Dividing Semiautonomous Organelles
- 1.9 Cell Cycle Regulation
- Suggested Reading
- List of Key Terms
- 2 Water and Plant Cells
- 2.1 Water in Plant Life
- 2.2 The Structure and Properties of Water
- Water is a polar molecule that forms hydrogen bonds
- Water is an excellent solvent
- Water has distinctive thermal properties relative to its size
- Water has a high surface tension
- Water has a high tensile strength
- 2.3 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
- 2.4 Water Potential
- The chemical potential of water represents the free-energy status of water
- Three major factors contribute to water potential
- Solutes
- Pressure
- Gravity
- Water potentials can be measured
- 2.5 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
- Water potential and its components vary with growth conditions and location within the plant
- 2.6 Cell Wall and Membrane Properties
- Small changes in plant cell volume cause large changes in turgor pressure
- The rate at which cells gain or lose water is influenced by plasma membrane hydraulic conductivity
- Aquaporins facilitate the movement of water across membranes
- 2.7 Plant Water Status
- Physiological processes are affected by plant water status
- Solute accumulation helps cells maintain turgor and volume
- Summary
- 2.1 Water in Plant Life
- 2.2 The Structure and Properties of Water
- 2.3 Diffusion and Osmosis
- 2.4 Water Potential
- 2.5 Water Potential of Plant Cells
- 2.6 Cell Wall and Membrane Properties
- 2.7 Plant Water Status
- Suggested Reading
- List of Key Terms
- 3 Water Balance of Plants
- 3.1 Water in the Soil
- Soil water potential is affected by solutes, surface tension, and gravity
- Water moves through the soil by bulk flow
- 3.2 Water Absorption by Roots
- Water moves in the root via the apoplast, symplasm, and transmembrane pathways
- Solute accumulation in the xylem can generate “root pressure”
- 3.3 Water Transport through the Xylem
- The xylem consists of two types of transport cells
- 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 m to a treetop?
- The cohesion–tension theory explains water transport in the xylem
- Xylem transport of water in trees faces physical challenges
- Plants have several mechanisms to overcome losses of xylem conductivity caused by embolism
- 3.4 Water Movement from the Leaf to the Atmosphere
- Leaves have a large hydraulic resistance
- The driving force for transpiration is the difference in water vapor concentration
- Water loss is also affected by the pathway resistances
- Stomatal control couples leaf transpiration to leaf photosynthesis
- The cell walls of guard cells have specialized features
- Changes in guard cell turgor pressure cause stomata to open and close
- Internal and external signals regulate the osmotic balance of guard cells
- The transpiration ratio measures the relationship between water loss and carbon gain
- 3.5 Overview: The Soil–Plant–Atmosphere Continuum
- Summary
- 3.1 Water in the Soil
- 3.2 Water Absorption by Roots
- 3.3 Water Transport through the Xylem
- 3.4 Water Movement from the Leaf to the Atmosphere
- 3.5 Overview: The Soil–Plant–Atmosphere Continuum
- Suggested Reading
- List of Key Terms
- 4 Mineral Nutrition
- 4.1 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
- Group 1: Deficiencies in mineral nutrients that are part of carbon compounds
- Nitrogen
- Sulfur
- Phosphorus
- Group 2: Deficiencies in mineral nutrients that are important for structural integrity
- Silicon
- Boron
- Group 3: Deficiencies in mineral nutrients that remain in ionic form
- Potassium
- Calcium
- Magnesium
- Chlorine
- Zinc
- Sodium
- Group 4: Deficiencies in mineral nutrients that are involved in redox reactions
- Iron
- Manganese
- Copper
- Nickel
- Molybdenum
- Plant tissue analysis reveals mineral deficiencies
- 4.2 Treating Nutritional Deficiencies
- Crop yields can be improved by the addition of fertilizers
- Some mineral nutrients can be absorbed by leaves
- 4.3 Soil, Roots, and Microbes
- Negatively charged soil particles affect the adsorption of mineral nutrients
- Soil pH affects nutrient availability, soil microbes, and root growth
- Excess mineral ions in the soil limit plant growth
- Some plants develop extensive root systems
- Root systems differ in form but are based on common structures
- Different areas of the root absorb mineral ions differently
- Nutrient availability influences root growth and development
- Mycorrhizal symbioses facilitate nutrient uptake by roots
- Nutrients move between mycorrhizal fungi and root cells
- 4.4 Fertilizer Production and Uses
- Summary
- 4.1 Essential Nutrients, Deficiencies, and Plant Disorders
- 4.2 Treating Nutritional Deficiencies
- 4.3 Soil, Roots, and Microbes
- 4.4 Fertilizer Production and Use
- Suggested Reading
- List of Key Terms
- 5 Assimilation of Inorganic Nutrients
- 5.1 Nitrogen in the Environment
- Nitrogen passes through several forms in a biogeochemical cycle
- Unassimilated ammonium or nitrate may be dangerous
- 5.2 Nitrate Assimilation
- Many factors regulate nitrate reductase
- Nitrite reductase converts nitrite to ammonium
- Both roots and shoots assimilate nitrate
- Nitrate can be transported in both xylem and phloem
- Transceptor contributes to nitrate signaling
- 5.3 Ammonium Assimilation
- Converting ammonium to amino acids requires two enzymes
- Ammonium can be assimilated via an alternative pathway
- Transamination reactions transfer nitrogen
- Asparagine and glutamine link carbon and nitrogen metabolism
- 5.4 Amino Acid Biosynthesis
- 5.5 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
- 5.6 Sulfur Assimilation
- Sulfate is the form of sulfur transported into plants
- Sulfate assimilation occurs mostly in leaves
- Methionine is synthesized from cysteine
- 5.7 Phosphate Assimilation
- miRNAs contribute to phosphate and sulfate signaling
- 5.8 Oxygen Assimilation
- 5.9 The Energetics of Nutrient Assimilation
- Summary
- 5.1 Nitrogen in the Environment
- 5.2 Nitrate Assimilation
- 5.3 Ammonium Assimilation
- 5.4 Amino Acid Biosynthesis
- 5.5 Biological Nitrogen Fixation
- 5.6 Sulfur Assimilation
- 5.7 Phosphate Assimilation
- 5.8 Oxygen Assimilation
- 5.9 The Energetics of Nutrient Assimilation
- Suggested Reading
- List of Key Terms
- 6 Solute Transport
- 6.1 Passive and Active Transport
- 6.2 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
- 6.3 Membrane Transport Processes
- Channels enhance diffusion across membranes
- Carriers bind and transport specific substances
- Primary active transport requires energy
- Secondary active transport is driven by ion gradients
- Kinetic analyses can elucidate transport mechanisms
- 6.4 Membrane Transport Proteins
- Transporters exist for diverse nitrogen-containing compounds
- Cation transporters are diverse
- Cation channels
- Cation carriers
- Cation function in cells
- Anion transporters have been identified
- Transporters for metal and metalloid ions transport essential micronutrients
- Aquaporins have diverse functions
- Plasma membrane H+-ATPases are highly regulated P-type ATPases
- The tonoplast H+-ATPase drives solute accumulation in vacuoles
- H+-pyrophosphatases and P-type H+-ATPases also pump protons at the tonoplast
- 6.5 Transport in Stomatal Guard Cells
- Blue light induces stomatal opening
- Abscisic acid and high CO2 induce stomatal closing
- 6.6 Ion Transport in Roots
- Solutes move through both apoplast and symplasm
- Ions cross both symplasm and apoplast
- Xylem parenchyma cells participate in xylem loading
- Summary
- 6.1 Passive and Active Transport
- 6.2 Transport of Ions across Membrane Barriers
- 6.3 Membrane Transport Processes
- 6.4 Membrane Transport Proteins
- 6.5 Transport in Stomatal Guard Cells
- 6.6 Ion Transport in Roots
- Suggested Reading
- List of Key Terms
- 7 Photosynthesis: The Light Reactions
- 7.1 Photosynthesis in Green Plants
- 7.2 General Concepts
- Light consists of photons with characteristic energies
- Absorption of photosynthetically active light changes the electronic states of chlorophylls
- Photosynthetic pigments absorb the light that powers photosynthesis
- 7.3 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
- 7.4 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
- 7.5 Organization of Light-Absorbing Antenna Systems
- Antenna systems contain chlorophyll and are membrane-associated
- The antenna funnels energy to the reaction center
- Many antenna pigment–protein complexes have a common structural motif
- 7.6 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 PSII reaction center is a multi-subunit pigment–protein complex
- Water is oxidized to oxygen by PSII
- Pheophytin and two quinones accept electrons from PSII
- Electron flow through the cytochrome b6f complex also transports protons
- Plastocyanin carries electrons between the cytochrome b6f complex and photosystem I
- The PSI reaction center oxidizes PC and reduces ferredoxin, which transfers electrons to NADP+
- Some herbicides block photosynthetic electron flow
- 7.7 Proton Transport and ATP Synthesis in the Chloroplast
- Cyclic electron flow augments the output of ATP to balance the chloroplast energy budget
- Summary
- 7.1 Photosynthesis in Green Plants
- 7.2 General Concepts
- 7.3 Key Experiments in Understanding Photosynthesis
- 7.4 Organization of the Photosynthetic Apparatus
- 7.5 Organization of Light-Absorbing Antenna Systems
- 7.6 Mechanisms of Electron Transport
- 7.7 Proton Transport and ATP Synthesis in the Chloroplast
- Suggested Reading
- List of Key Terms
- 8 Photosynthesis: The Carbon Reactions
- 8.1 The Calvin–Benson Cycle
- The Calvin–Benson cycle has three phases: carboxylation, reduction, and regeneration
- 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
- 8.2 The Oxygenation Reaction of Rubisco and Photorespiration
- The oxygenation of ribulose 1,5-bisphosphate sets in motion photorespiration
- Photorespiration is linked to the photosynthetic electron transport system
- Enzymes of plant photorespiration derive from different ancestors
- Photorespiration interacts with many metabolic pathways
- 8.3 Inorganic Carbon-Concentrating Mechanisms
- 8.4 Inorganic Carbon-Concentrating Mechanisms: C4 Photosynthetic Carbon Fixation
- Malate and aspartate are the primary carboxylation products of the C4 cycle
- Kranz-type C4 plants assimilate CO2 by the concerted action of two different types of cells
- The C4 subtypes use different mechanisms to decarboxylate 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 requires more transport processes than in C3 plants
- In hot, dry climates, the C4 cycle reduces photorespiration
- 8.5 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
- 8.6 Accumulation and Partitioning of Photosynthates—Starch and Sucrose
- Summary
- 8.1 The Calvin–Benson Cycle
- 8.2 The Oxygenation Reaction of Rubisco and Photorespiration
- 8.3 Inorganic Carbon-Concentrating Mechanisms
- 8.4 Inorganic Carbon-Concentrating Mechanisms: C4 Photosynthetic Carbon Fixation
- 8.5 Inorganic Carbon-Concentrating Mechanisms: Crassulacean Acid Metabolism (CAM)
- 8.6 Accumulation and Partitioning of Photosynthates—Starch and Sucrose
- Suggested Reading
- List of Key Terms
- 9 Photosynthesis: Physiological and Ecological Considerations
- 9.1 Photosynthesis Is Influenced by Leaf Properties
- Leaf anatomy and canopy structure optimize light absorption
- Leaf angle and leaf movement can control light absorption
- Leaves acclimate to sun and shade environments
- 9.2 Effects of Light on Photosynthesis in the Intact Leaf
- Photosynthetic light-response curves reveal differences in leaf properties
- Leaves must dissipate excess light energy as heat
- The xanthophyll cycle
- Chloroplast movements
- Leaf movements
- Absorption of too much light can lead to photoinhibition
- 9.3 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
- 9.4 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 supply imposes limitations on photosynthesis
- C3 versus C4 plants
- CAM plants
- How will photosynthesis and respiration change in the future under elevated CO2 conditions?
- Summary
- 9.1 Photosynthesis Is Influenced by Leaf Properties
- 9.2 Effects of Light on Photosynthesis in the Intact Leaf
- 9.3 Effects of Temperature on Photosynthesis in the Intact Leaf
- 9.4 Effects of Carbon Dioxide on Photosynthesis in the Intact Leaf
- Suggested Reading
- List of Key Terms
- 10 Translocation in the Phloem
- 10.1 Patterns of Translocation: Source to Sink
- 10.2 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
- Companion cells aid the highly specialized sieve elements
- 10.3 Phloem Loading
- Phloem loading can occur via the apoplast or symplasm
- Apoplastic loading is characteristic of many herbaceous species
- Sucrose loading in the apoplastic pathway requires metabolic energy
- Phloem loading in the apoplastic pathway involves a sucrose–H+ symporter
- Transfer cells are companion cells that are specialized for membrane transport
- Phloem loading is symplasmic in some species
- The oligomer-trapping model explains symplasmic loading in plants with intermediary-type companion cells
- Phloem loading is passive in several tree species
- The type of phloem loading is correlated with several significant characteristics
- 10.4 Long-Distance Transport: A Pressure-Driven Mechanism
- Mass transfer is much faster than diffusion
- The pressure-flow model is a passive mechanism for phloem transport
- The pressure is osmotically generated
- Some predictions of pressure flow have been confirmed, while others require further experimentation
- Functional sieve plate pores appear to be open channels
- Are the pressure gradients in the sieve elements sufficient to drive phloem transport in trees?
- 10.5 Materials Translocated in the Phloem
- Sugars are translocated in a nonreducing form
- Other small organic and inorganic solutes are translocated in the phloem
- Phloem-mobile macromolecules often originate in companion cells
- Damaged sieve elements are sealed off
- 10.6 Phloem Unloading and Sink-to-Source Transition
- Phloem unloading and short-distance transport can occur via symplasmic or apoplastic pathways
- Symplasmic unloading supplies growing vegetative sinks
- Import into seeds, fruits, and storage organs often involves an apoplastic step
- Apoplastic import is active and requires metabolic energy
- The transition of a leaf from sink to source is gradual
- 10.7 Photosynthate Distribution: Allocation and Partitioning
- Allocation includes storage, utilization, and transport
- Source leaves regulate allocation
- Various sinks partition transport sugars
- 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
- 10.8 Transport of Signaling Molecules
- Turgor pressure and chemical signals coordinate source and sink activities
- Mobile RNAs function as signal molecules to regulate growth and development
- Mobile proteins also function as signal molecules to regulate growth and development
- Plasmodesmata function in phloem signaling
- Summary
- 10.1 Patterns of Translocation: Source to Sink
- 10.2 Pathways of Translocation
- 10.3 Phloem Loading
- 10.4 Long-Distance Transport: A Pressure-Driven Mechanism
- 10.5 Materials Translocated in the Phloem
- 10.6 Phloem Unloading and Sink-to-Source Transition
- 10.7 Photosynthate Distribution: Allocation and Partitioning
- 10.8 Transport of Signaling Molecules
- Suggested Reading
- List of Key Terms
- 11 Respiration and Lipid Metabolism
- 11.1 Overview of Plant Respiration
- 11.2 Glycolysis
- Glycolysis metabolizes carbohydrates from several sources
- The energy-conserving phase of glycolysis produces pyruvate, ATP, and NADH
- Plants have alternative glycolytic reactions
- Fermentation regenerates the NAD+ needed for glycolytic ATP production in the absence of oxygen
- 11.3 The Oxidative Pentose Phosphate Pathway
- The oxidative pentose phosphate pathway produces NADPH and biosynthetic intermediates
- The oxidative pentose phosphate pathway is controlled by cellular redox status
- 11.4 The Tricarboxylic Acid Cycle
- Mitochondria are semiautonomous organelles
- Pyruvate enters the mitochondrion and is oxidized via the TCA cycle
- The TCA cycle of plants has unique features
- 11.5 Oxidative Phosphorylation
- The electron transport chain catalyzes a flow of electrons from NADH to O2
- Complex I (NADH dehydrogenase)
- Complex II (succinate dehydrogenase)
- Complex III (cytochrome bc1 complex)
- Complex IV (cytochrome c oxidase)
- The electron transport chain has supplementary branches
- ATP synthesis in the mitochondrion is coupled to electron transport
- Transporters exchange substrates and products
- Aerobic respiration yields about 60 molecules of ATP per molecule of sucrose
- Plants have several mechanisms that lower the ATP yield
- The alternative oxidase
- The uncoupling protein
- Rotenone-insensitive NAD(P)H dehydrogenases
- Respiration is an integral part of a redox and biosynthesis network
- Respiration is controlled at multiple levels
- 11.6 Respiration in Intact Plants and Tissues
- Plants respire roughly half of the daily photosynthetic yield
- Respiratory processes operate during photosynthesis
- Different tissues and organs respire at different rates
- Environmental factors alter respiration rates
- Oxygen
- Temperature
- Carbon dioxide
- 11.7 Lipid Metabolism
- Fats and oils store large amounts of energy
- Triacylglycerols are stored in oil bodies
- Polar glycerolipids are the main structural lipids in membranes
- Glycerolipids are synthesized in the plastids and the ER
- Lipid composition influences membrane function
- Membrane lipids are precursors of important signaling compounds
- Storage lipids are converted into carbohydrates in germinating seeds
- Overview: lipids to sucrose
- Lipase-mediated hydrolysis
- Β–Oxidation of fatty acids
- The glyoxylate cycle
- The mitochondrial role
- Summary
- 11.1 Overview of Plant Respiration
- 11.2 Glycolysis
- 11.3 The Oxidative Pentose Phosphate Pathway
- 11.4 The Tricarboxylic Acid Cycle
- 11.5 Oxidative Phosphorylation
- 11.6 Respiration in Intact Plants and Tissues
- 11.7 Lipid Metabolism
- Suggested Reading
- List of Key Terms
- 12 Signals and Signal Transduction
- 12.1 Temporal and Spatial Aspects of Signaling
- 12.2 Signal Perception and Amplification
- Protein kinases are major signal transduction components
- Kinases also function in signal amplification
- Phosphatases are the “off switch” of protein phosphorylation
- Other protein modifications can reconfigure cellular processes
- Small-molecule second messengers rapidly amplify signals
- 12.3 Hormones and Plant Development
- Auxin was discovered in early studies of phototropism
- Gibberellins promote stem growth and were discovered in studies of the “foolish seedling disease” of rice
- 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, floral sex identity, and other developmental processes
- Strigolactones suppress branching and promote rhizosphere interactions
- 12.4 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
- Ethylene is synthesized from methionine via ACC
- Abscisic acid is synthesized from a carotenoid intermediate
- Brassinosteroids are derived from the sterol campesterol
- Strigolactones are synthesized from β-carotene
- 12.5 Movement of Hormones within the Plant
- Plant polarity is maintained by polar auxin streams
- Auxin uptake
- Auxin efflux
- Auxin transport is regulated by multiple mechanisms
- 12.6 Hormonal Signaling Pathways
- The cytokinin and ethylene signal transduction pathways derive from bacterial two-component regulatory systems
- Phosphorylation also regulates brassinosteroid, ABA, and auxin perception
- Multiple plant hormone receptors use ubiquitination to remove repressor proteins
- Plants use electrical signaling for communication between tissues
- Cross-regulation allows signal transduction pathways to be integrated
- Summary
- 12.1 Temporal and Spatial Aspects of Signaling
- 12.2 Signal Perception and Amplification
- 12.3 Hormones and Plant Development
- 12.4 Phytohormone Metabolism and Homeostasis
- 12.5 Movement of Hormones within the Plant
- 12.6 Hormonal Signaling Pathways
- Suggested Reading
- List of Key Terms
- 13 Signals from Sunlight
- 13.1 Plant Photoreceptors
- Photoresponses are driven by the spectral properties of the energy absorbed
- Plant responses to light can be distinguished by the amount of light required
- 13.2 Phytochromes
- Phytochrome is the primary photoreceptor for red and far-red light
- Phytochrome can interconvert between Pr and Pfr forms
- 13.3 Phytochrome Responses
- Phytochrome responses vary in lag time and escape time
- Phytochrome responses fall into three main categories based on the amount of light required
- Phytochrome A mediates responses to continuous far-red light
- Phytochrome regulates gene expression
- 13.4 Blue-Light Responses and Photoreceptors
- Blue-light responses have characteristic kinetics and lag times
- 13.5 Cryptochromes
- Blue-light irradiation of the cryptochrome FAD chromophore causes a conformational change
- The nucleus is a primary site of cryptochrome action
- Cryptochrome interacts with phytochrome
- 13.6 Phototropins
- Phototropins activated by directional blue light initiate seedling phototropism
- Phototropins regulate chloroplast movements
- Stomatal opening is regulated by blue light, which activates the plasma membrane H+-ATPase
- 13.7 Responses to Ultraviolet Radiation
- Plant photoresponses observed in the field are composite activities
- Summary
- 13.1 Plant Photoreceptors
- 13.2 Phytochromes
- 13.3 Phytochrome Responses
- 13.4 Blue-Light Responses and Photoreceptors
- 13.5 Cryptochromes
- 13.6 Phototropins
- 13.7 Responses to Ultraviolet Radiation
- Suggested Reading
- List of Key Terms
- 14 Seed Dormancy, Germination, and Seedling Establishment
- 14.1 Seed Anatomy
- Seed anatomy varies widely among different plant groups
- 14.2 Seed Dormancy
- There are two basic types of seed dormancy mechanisms: exogenous and endogenous
- Non-dormant seeds can exhibit vivipary and precocious germination
- The ABA:GA ratio is the primary determinant of embryonic seed dormancy
- 14.3 Release from Dormancy
- Light is an important signal that breaks dormancy in small seeds
- Some seeds require either chilling or after-ripening to break dormancy
- Seed dormancy can be broken by various chemical compounds
- 14.4 Seed Germination
- Germination and postgermination can be divided into three phases corresponding to the phases of water uptake
- 14.5 Mobilization of Stored Reserves
- Cereal seeds are a model for understanding starch mobilization
- Legume seeds are a model for understanding protein mobilization
- Oilseeds are a model for understanding lipid remobilization
- 14.6 Cell Expansion Mechanisms
- Polarized growth and cell expansion involve the cytoskeleton and cell wall modification
- Microfibril orientation influences growth directionality of cells with diffuse growth
- Acid-induced growth and cell wall yielding are mediated by expansins
- 14.7 Seedling Growth and Establishment
- The development of emerging seedlings is strongly influenced by light
- Gibberellins and brassinosteroids both suppress photomorphogenesis in darkness
- Hook opening is regulated by phytochrome, auxin, and ethylene
- Vascular differentiation begins during seedling emergence
- The root tip has specialized cells
- Ethylene and other hormones regulate root hair development
- 14.8 Differential Growth Enables Successful Seedling Establishment
- Ethylene affects microtubule orientation and induces lateral cell expansion
- Auxin promotes growth in stems and coleoptiles, while inhibiting growth in roots
- The minimum lag time for auxin-induced elongation is 10 minutes
- Auxin-induced proton extrusion loosens the cell wall
- 14.9 Tropisms: Growth in Response to Directional Stimuli
- Gravitropism involves the lateral redistribution of auxin
- The gravitropic stimulus perturbs the symmetric movements of auxin
- Gravity perception is triggered by the sedimentation of amyloplasts
- Gravity sensing may involve pH and calcium ions (Ca2+) as second messengers
- Thigmotropism involves signaling by Ca2+, pH, and reactive oxygen species
- Hydrotropism involves ABA signaling and asymmetric cytokinin responses
- Phototropins are the light receptors involved in phototropism
- Phototropism is mediated by the lateral redistribution of auxin
- Shoot phototropism occurs in a series of steps
- Summary
- 14.1 Seed Anatomy
- 14.2 Seed Dormancy
- 14.3 Release from Dormancy
- 14.4 Seed Germination
- 14.5 Mobilization of Stored Reserves
- 14.6 Cell Expansion Mechanisms
- 14.7 Seedling Growth and Establishment
- 14.8 Differential Growth Enables Successful Seedling Establishment
- 14.9 Tropisms: Growth in Response to Directional Stimuli
- Suggested Reading
- List of Key Terms
- 15 Vegetative Growth and Senescence
- 15.1 The Root Apical Meristem (RAM)
- The root tip has four developmental zones
- Auxin and cytokinin contribute to the maintenance and function of the RAM
- 15.2 The Shoot Apical Meristem (SAM)
- The shoot apical meristem has distinct cytohistological zones and layers
- Cell identity in the SAM is determined by position rather thanby cell lineage
- The size of the SAM is determined by cell number
- 15.3 Leaf Structure and Phyllotaxy
- The arrangement of leaves around the shoot is called phyllotaxy
- Auxin-dependent patterning of the shoot apex begins during embryogenesis
- 15.4 Differentiation of Epidermal Cell Types
- A specialized epidermal lineage produces guard cells
- 15.5 Venation Patterns in Leaves
- The primary leaf vein is initiated in the leaf primordium
- Auxin canalization initiates development of the leaf trace
- 15.6 Shoot Branching and Architecture
- Auxin, cytokinins, and strigolactones regulate axillary bud outgrowth
- The initial signal for axillary bud growth may be an increase in sucrose availability to the bud
- 15.7 Shade Avoidance
- Reducing shade avoidance responses can improve crop yields
- 15.8 Root Branching and Architecture
- Lateral root primordia arise from initial cells in the pericycle
- Lateral root formation can be divided into four distinct stages
- Plants can modify their root system architecture to optimize water and nutrient uptake
- Monocots and eudicots differ in their root system architecture
- Root system architecture changes in response to phosphorus deficiencies
- 15.9 Secondary Growth
- The vascular cambium produces secondary xylem and phloem
- The cork cambium produces the periderm
- 15.10 Plant Senescence
- Programmed cell death (PCD) is a normal aspect of development in all eukaryotic organisms
- During leaf senescence, nutrients are remobilized from the source leaf to vegetative or reproductive sinks
- The developmental age of a leaf may differ from its chronological age
- Leaf senescence may be sequential, seasonal, or stress-induced
- The earliest cellular changes during leaf senescence occur in the chloroplast
- Reactive oxygen species serve as internal signaling agents in leaf senescence
- Plant hormones interact in the regulation of leaf senescence
- 15.11 Leaf Abscission
- The timing of leaf abscission is regulated by the interaction of ethylene and auxin
- 15.12 Whole Plant Senescence
- Angiosperm life cycles may be annual, biennial, or perennial
- Nutrient or hormonal redistribution may trigger senescence in monocarpic plants
- Summary
- 15.1 The Root Apical Meristem (RAM)
- 15.2 The Shoot Apical Meristem (SAM)
- 15.3 Leaf Structure and Phyllotaxy
- 15.4 Differentiation of Epidermal Cell Types
- 15.5 Venation Patterns in Leaves
- 15.6 Shoot Branching and Architecture
- 15.7 Shade Avoidance
- 15.8 Root Branching and Architecture
- 15.9 Secondary Growth
- 15.10 Plant Senescence
- 15.11 Leaf Abscission
- 15.12 Whole Plant Senescence
- Suggested Reading
- List of Key Terms
- 16 Flowering and Double Fertilization
- 16.1 Floral Evocation: Integrating Environmental Cues
- 16.2 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 epigenetic regulation
- 16.3 Photoperiodism: Monitoring Day Length
- Plants can be classified according to their photoperiodic responses
- Photoperiodism is one of many plant processes controlled by a circadian rhythm
- Circadian rhythms exhibit characteristic features
- Circadian rhythms adjust to different day–night cycles
- The leaf is the site of perception of the photoperiodic signal
- Plants monitor day length by measuring the length of the night
- Night breaks can cancel the effect of the dark period
- Photoperiodic timekeeping during the night depends on a circadian clock
- The coincidence model links oscillating light sensitivity and photoperiodism
- Phytochrome is the primary photoreceptor in photoperiodism
- 16.4 Vernalization: Promoting Flowering with Cold Treatment
- 16.5 Long-Distance Signaling Involved in Flowering
- Gibberellins and ethylene can induce flowering
- 16.6 Floral Meristems and Floral Organ Development
- The SAM in Arabidopsis changes with development
- The four different types of floral organs are initiated as separate whorls
- Two major categories of genes regulate floral development
- The ABC model partially explains the determination of floral organ identity
- 16.7 Pollen Development in the Anther
- 16.8 Embryo Sac Development in the Ovule
- Functional megaspores undergo a series of free nuclear mitotic divisions followed by cellularization
- 16.9 Pollination and Double Fertilization in Flowering Plants
- Two sperm cells are delivered to the female gametophyte by the pollen tube
- Pollination begins with adhesion and hydration of a pollen grain on a compatible flower
- Pollen tubes grow by tip growth
- Double fertilization results in the formation of the zygote and the primary endosperm cell
- Summary
- 16.1 Floral Evocation: Integrating Environmental Cues
- 16.2 The Shoot Apex and Phase Changes
- 16.3 Photoperiodism: Monitoring Day Length
- 16.4 Vernalization: Promoting Flowering with Cold Treatment
- 16.5 Long-Distance Signaling Involved in Flowering
- 16.6 Floral Meristems and Floral Organ Development
- 16.7 Pollen Development in the Anther
- 16.8 Embryo Sac Development in the Ovule
- 16.9 Pollination and Double Fertilization in Flowering Plants
- Suggested Reading
- List of Key Terms
- 17 Seed and Fruit Development
- 17.1 Seed Structure
- 17.2 Establishment of the Embryonic Axis
- 17.3 Seed Endosperm Development
- Endosperm development and embryogenesis can occur autonomously
- Cells of the starchy endosperm and aleurone layer follow divergent developmental pathways
- 17.4 Seed Coat Development
- 17.5 Seed Maturation and Desiccation Tolerance
- Embryos accumulate proteins and sugars during desiccation
- Coat-imposed dormancy correlates with long-term seed viability
- 17.6 Fruit Development and Ripening
- Auxin and gibberellic acid (GA) regulate fruit set and parthenocarpy
- Mature dehiscent fruits open spontaneously to release seeds
- Fleshy fruits have extensive mesocarp layers
- Fleshy fruits undergo ripening
- Ripening involves changes in the color of fruit
- Fruit softening involves the coordinated action of many cell wall–degrading enzymes
- Taste and flavor reflect changes in acids, sugars, aroma, and other compounds
- Ethylene regulates the ripening of many fruits
- Climacteric and non-climacteric fruit differ in their ethylene responses
- Commercial control of fruit ripening
- Summary
- 17.1 Seed Structure
- 17.2 Establishment of the Embryonic Axis
- 17.3 Seed Endosperm Development
- 17.4 Seed Coat Development
- 17.5 Seed Maturation and Desiccation Tolerance
- 17.6 Fruit Development and Ripening
- Suggested Reading
- List of Key Terms
- 18 Biotic Interactions
- 18.1 Plant Interactions with Beneficial Microorganisms
- Rhizobacteria can increase nutrient availability, stimulate root branching, and protect against pathogens
- 18.2 Herbivore Interactions That Harm Plants
- Mechanical barriers provide a first line of defense against insect pests and pathogens
- Plants’ specialized metabolites can deter insect herbivores
- Plants store constitutive toxic compounds in specialized structures
- Plants often store defense chemicals as nontoxic water-soluble sugar conjugates in the vacuole
- 18.3 Inducible Defense Responses to Insect Herbivores
- Plants can recognize specific components of insect saliva
- Jasmonate activates defense responses against insect herbivores
- Hormonal interactions contribute to plant–insect herbivore interactions
- Jasmonate initiates the production of defense proteins that inhibit herbivore digestion
- Herbivore damage induces systemic defenses
- Herbivore-induced volatiles can repel herbivores and attract natural enemies
- Herbivore-induced volatiles can serve as long-distance signals between plants
- Herbivore-induced volatiles can also act as systemic signals within a plant
- Insects have evolved mechanisms to defeat plant defenses
- 18.4 Plant Defenses against Pathogens
- Microbial pathogens have evolved various strategies to invade host plants
- Pathogens produce effector molecules that aid in the colonization of their plant host cells
- Plants can detect pathogens through perception of pathogen-derived “danger signals”
- Resistosomes recognize strain-specific effectors
- The hypersensitive response is a common defense against pathogens
- A single encounter with a pathogen may increase resistance to future attacks
- Phytoalexins with antimicrobial activity accumulate after pathogen attack
- RNA interference plays a central role in antiviral immune responses in plants
- Some plant parasitic nematodes form specific associations through the formation of distinct feeding structures
- Plants compete with other plants by secreting allelopathic specialized metabolites into the soil
- Some plants are parasites of other plants
- Summary
- 18.1 Plant Interactions with Beneficial Microorganisms
- 18.2 Herbivore Interactions That Harm Plants
- 18.3 Inducible Defense Responses to Insect Herbivores
- 18.4 Plant Defenses against Pathogens
- Suggested Reading
- List of Key Terms
- 19 Abiotic Stress
- 19.1 Defining Plant Stress
- Physiological adjustment to abiotic stress involves trade-offs between vegetative and reproductive development
- 19.2 Acclimation versus Adaptation
- 19.3 Environmental Stressors
- Water deficit decreases turgor pressure, increases ion toxicity, and inhibits photosynthesis
- Temperature stress affects a broad spectrum of physiological processes
- Flooding causes anaerobic stress to roots
- Salinity stress has both osmotic and cytotoxic effects
- During freezing stress, extracellular ice crystal formation causes cell dehydration
- Heavy metals can both mimic essential mineral nutrients and generate ROS
- Combinations of abiotic stresses can induce unique signaling and metabolic pathways
- Sequential exposure to different abiotic stresses sometimes confers cross-protection
- 19.4 Stress-Sensing Mechanisms in Plants
- Plants use a variety of mechanisms to sense abiotic stress
- Chloroplasts and mitochondria respond to abiotic stress by sending stress signals to the nucleus
- Plant-wide waves of Ca2+ and ROS mediate systemic acquired acclimation
- Hormonal interactions regulate a wide range of abiotic stress responses
- Epigenetic mechanisms are also involved in sensing abiotic stress
- 19.5 Developmental and Physiological Mechanisms That Protect Plants against Abiotic Stress
- Plants can activate developmental programs that alter their phenotype
- Leaf area
- Leaf orientation
- Trichomes
- Cuticle
- Root-to-shoot ratio
- Plants adjust osmotically to drying soils 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 structure in response to temperature and other abiotic stresses
- Exclusion and internal tolerance mechanisms allow plants to cope with toxic ions
- Plants use cryoprotectant molecules and antifreeze proteins to prevent ice crystal formation
- Summary
- 19.1 Defining Plant Stress
- 19.2 Acclimation versus Adaptation
- 19.3 Environmental Stressors
- 19.4 Stress-Sensing Mechanisms in Plants
- 19.5 Developmental and Physiological Mechanisms That Protect Plants against Abiotic Stress
- Suggested Reading
- List of Key Terms
- Appendix Plant Breeding and Genome Engineering
- A.1 Genetic Modification of Plants
- A.2 Editing Plant Genomes with Sequence-Specific Nucleases
- A.3 Genetic Engineering of Plants Remains Controversial
- List of Key Terms
- Glossary
- A
- B
- C
- D
- E
- F
- G
- H
- I
- J
- K
- L
- M
- N
- O
- P
- Q
- R
- S
- T
- U
- V
- W
- X
- Z
- Illustration Credits
- CHAPTER 1
- CHAPTER 2
- CHAPTER 3
- CHAPTER 4
- CHAPTER 5
- CHAPTER 6
- CHAPTER 7
- CHAPTER 8
- CHAPTER 9
- CHAPTER 10
- CHAPTER 11
- CHAPTER 12
- CHAPTER 13
- CHAPTER 14
- CHAPTER 15
- CHAPTER 16
- CHAPTER 17
- CHAPTER 18
- CHAPTER 19
- APPENDIX
- Index
- List of Illustrations
- List of Tables
- Images
- 1 Plant and Cell Architecture
- 2 Water and Plant Cells
- 3 Water Balance of Plants
- 4 Mineral Nutrition
- 5 Assimilation of Inorganic Nutrients
- 6 Solute Transport
- 7 Photosynthesis: The Light Reactions
- 8 Photosynthesis: The Carbon Reactions
- 9 Photosynthesis: Physiological and Ecological Considerations
- 10 Translocation in the Phloem
- 11 Respiration and Lipid Metabolism
- 12 Signals and Signal Transduction
- 13 Signals from Sunlight
- 14 Seed Dormancy, Germination, and Seedling Establishment
- 15 Vegetative Growth and Senescence
- 16 Flowering and Double Fertilization
- 17 Seed and Fruit Development
- 18 Biotic Interactions
- 19 Abiotic Stress
- Appendix Plant Breeding and Genome Engineering