About the Authors
JANET IWASA is an Assistant Professor in the Biochemistry Department at the University of Utah. She received her bachelor’s degree from Williams College and a Ph.D. in Cell Biology from the University of California, San Francisco, where she first became interested in the visualization of biological processes. As a postdoctoral fellow, she was awarded a fellowship from the National Science Foundation to create a multimedia exhibit with Nobel Laureate Jack Szostak (Harvard University) and the Museum of Science, Boston. She later joined Harvard Medical School as a faculty member in the Department of Cell Biology, where she utilized visualization tools to aid in scientific communication, exploration, and outreach. Janet’s award-winning illustrations and animations have appeared in scientific journals including Nature, Science, and Cell, as well as in the New York Times. She currently runs a group, called the Animation Lab, that focuses on the innovative use of visualization tools for molecular biology research, education and outreach.
WALLACE MARSHALL is Professor of Biochemistry and Biophysics at the University of California San Francisco and an ASCB Fellow. A native Long-Islander, he received his bachelor’s degrees in Electrical Engineering and Biochemistry from the State University of
New York at Stony Brook, and his Ph.D. in Biochemistry from UC San Francisco, where he studied organization of chromosomes within the nucleus with John Sedat. He then moved to Yale University for postdoctoral studies with Joel Rosenbaum, where he became interested in questions of organelle size control and cell organization, using cilia, flagella, and centrioles as model systems. In 2003, he joined the faculty at UCSF where he continues to study questions of cellular organization in a variety of model organisms including green algae, yeast, ciliates, and mammalian cells. In addition to his cell biology research, he teaches Human Metabolism for the UCSF School of Pharmacy, Cell Biology for the UCSF Graduate Division, and a twoweek lab course on cell behavior. In 2014, he served as Program Committee Chair organizing the annual meeting of the American Society for Cell Biology. From 2014 to 2018 he served as co-director of the Physiology summer course at the Marine Biological Laboratory in Woods Hole, Massachusetts. He currently directs the Center for Cellular Construction, a National Science Foundation Science and Technology Center devoted to engineering cells. Both authors are Council members of the American Society for Cell Biology.
Contents
Cover Title Page
Copyright About the Authors
Preface to the Ninth Edition
Acknowledgments
We also wish to thank all reviewers of this and previous editions
Nobel Prizes Awarded for Research in Cell and Molecular Biology Since 1958
Topics of Human Interest
CHAPTER 1: Introduction to the Study of Cell and Molecular Biology
1.1 The Discovery of Cells
1.2 Basic Properties of Cells
1.3 Two Fundamentally Different Classes of Cells
1.4 Viruses and Viroids
1.5 Green Cells: Volvox, an Experiment in Multicellularity
1.6 Engineering Linkage: Tissue Engineering
Analytic Questions
CHAPTER 2: The Chemical Basis of Life
2.1 Covalent Bonds
2.2 Engineering Linkage: Radionuclides for Imaging and Treatment
2.3 Noncovalent Bonds
2.4 Acids, Bases, and Buffers
2.5 The Nature of Biological Molecules
2.6 Green Cells: Chemical Fertilizers
2.7 Four Types of Biological Molecules
2.8 The Formation of Complex Macromolecular Structures
Analytic Questions
CHAPTER 3: Bioenergetics, Enzymes, and Metabolism
3.1 Bioenergetics
3.2 Enzymes as Biological Catalysts
3.3 Metabolism
3.4 Green Cells: Regulation of Metabolism by the Light/Dark Cycle
3.5 Engineering Linkage: Using Metabolism to Image Tumors
Analytic Questions
CHAPTER 4: The Structure and Function of the Plasma Membrane
4.1 Introduction to the Plasma Membrane
4.2 The Chemical Composition of Membranes
4.3 Membrane Proteins
4.4 Membrane Lipids and Membrane Fluidity
4.5 The Dynamic Nature of the Plasma Membrane
4.6 The Movement of Substances across Cell Membranes
4.7 Membrane Potentials and Nerve Impulses
4.8 Green Cells: Electrical Signaling in Plants
4.9 Engineering Linkage: Neurotechnology
Analytic Questions
CHAPTER 5: Aerobic Respiration and the Mitochondrion
5.1 Mitochondrial Structure and Function
5.2 Aerobic Metabolism in the Mitochondrion
5.3 The Role of Mitochondria in the Formation of ATP
5.4 Engineering Linkage: Measuring Blood Oxygen
5.5 Establishment of a Proton-Motive Force
5.6 The Machinery for ATP Formation
5.7 Peroxisomes
5.8 Green Cells: Glyoxysomes
Analytic Questions
CHAPTER 6: Photosynthesis and the Chloroplast
6.1 The Origin of Photosynthesis
6.2 Chloroplast Structure
6.3 An Overview of Photosynthetic Metabolism
6.4 The Absorption of Light
6.5 Green Cells: Chromoplasts
6.6 Photosynthetic Units and Reaction Centers
6.7 Photophosphorylation
6.8 Carbon Dioxide Fixation and the Synthesis of Carbohydrate
6.9 Engineering Linkage: Photodynamic Therapy
Analytic Questions
CHAPTER 7: Interactions Between Cells and Their Environment
7.1 Extracellular Interactions
7.2 Engineering Linkage: Organoids
7.3 Interactions of Cells with Extracellular Materials
7.4 Interactions of Cells with Other Cells
7.5 Tight Junctions: Sealing the Extracellular Space
7.6 Intercellular Communication
7.7 Cell Walls
7.8 Green Cells: Cell Walls and Plant Terrestrialization
Analytic Questions
CHAPTER 8: Cytoplasmic Membrane Systems: Structure, Function, and Membrane Traff...
8.1 An Overview of the Endomembrane System
8.2 A Few Approaches to the Study of Endomembranes
8.3 The Endoplasmic Reticulum
8.4 The Golgi Complex
8.5 Types of Vesicle Transport
8.6 Engineering Linkage: Extracellular Vesicles for Drug Delivery
8.7 Lysosomes
8.8 Green Cells: Plant Cell Vacuoles
8.9 The Endocytic Pathway: Moving Membrane and Materials into the Cell Interior
8.10 Posttranslational Uptake of Proteins by Peroxisomes, Mitochondria, and Chlo...
Analytic Questions
CHAPTER 9: The Cytoskeleton and Cell Motility
9.1 Overview of the Major Functions of the Cytoskeleton
9.2 Structure and Function of Microtubules
9.3 Green Cells: Why the Woodbine Twineth
9.4 Motor Proteins: Kinesins and Dyneins
9.5 Microtubule-Organizing Centers (MTOCs)
9.6 Structure and Function of Cilia and Flagella
9.7 Intermediate Filaments
9.8 Actin and Myosin
9.9 Muscle Organization and Contraction
9.10 Engineering Linkage: Muscle Biomechanics
9.11 Actin-Binding Proteins
9.12 Cellular Motility
9.13 The Bacterial Cytoskeleton
Analytic Questions
CHAPTER 10: The Nature of the Gene and the Genome
10.1 The Concept of a Gene as a Unit of Inheritance
10.2 The Discovery of Chromosomes
10.3 Chromosomes as the Carriers of Genetic Information
10.4 The Chemical Nature of the Gene
10.5 The Complexity of the Genome
10.6 The Stability of the Genome
10.7 Sequencing Genomes: The Footprints of Biological Evolution
10.8 Engineering Linkage: Engineering Genomes
10.9 The Genetic Basis of “Being Human”
10.10 Green Cells: Gene Transfer by Agrobacterium tumefaciens
Analytic Questions
CHAPTER 11: The Central Dogma: DNA to RNA to Protein
11.1 The Relationships among Genes, Proteins, and RNAs
11.2 An Overview of Transcription in Both Prokaryotic and Eukaryotic Cells
11.3 Synthesis and Processing of Eukaryotic Ribosomal and Transfer RNAs
11.4 Synthesis and Structure of Eukaryotic Messenger RNAs
11.5 Small Regulatory RNAs and RNA Silencing Pathways
11.6 Green Cells: Long-Range siRNA Movement
11.7 CRISPR and Other Noncoding RNAs
11.8 Encoding Genetic Information
11.9 Decoding the Codons: The Role of Transfer RNAs
11.10 Translating Genetic Information
11.11 Engineering Linkage: DNA Origami
Analytic Questions
CHAPTER 12: Control of Gene Expression
12.1 Control of Gene Expression in Bacteria
12.2 Engineering Linkage: Building Digital Logic with Genes
12.3 Structure of the Nuclear Envelope
12.4 Chromosomes and Chromatin
12.5 The Nucleus as an Organized Organelle
12.6 An Overview of Gene Regulation in Eukaryotes
12.7 Transcriptional Control
12.8 Green Cells: The ABC Model and MADS Domain Transcription Factors
12.9 RNA Processing Control
12.10 Translational Control
12.11 Posttranslational Control: Determining Protein Stability
Analytic Questions
CHAPTER 13: DNA Replication and Repair
13.1 DNA Replication
13.2 DNA Replication in Bacterial Cells
13.3 The Structure and Functions of DNA Polymerases
13.4 Replication in Viruses
13.5 Engineering Linkage: Storing Data in DNA
13.6 DNA Replication in Eukaryotic Cells
13.7 DNA Repair
13.8 Green Cells: Gamma Gardens
13.9 Between Replication and Repair
Analytic Questions
CHAPTER 14: Cell Division
14.1 The Cell Cycle
14.2 M Phase: Mitosis and Cytokinesis
14.3 Engineering Linkage: The Role of Membrane Tension in Cell Division
14.4 Green Cells: Unique Aspects of Plant Cell Division
14.5 Meiosis
Analytic Questions
CHAPTER 15: Cell Signaling and Signal Transduction: Communication between Cells
15.1 The Basic Elements of Cell Signaling Systems
15.2 A Survey of Extracellular Messages and Their Receptors
15.3 G Protein-Coupled Receptors and Their Second Messengers
15.4 Engineering Linkage: Biosensors in Medicine and Biology
15.5 Protein-Tyrosine Phosphorylation as a Mechanism for Signal Transduction
15.6 Green Cells: Auxin Signaling
15.7 The Role of Calcium as an Intracellular Messenger
15.8 Convergence, Divergence, and Cross-Talk among Different Signaling Pathways
15.9 The Role of NO as an Intracellular Messenger
15.10 Apoptosis (Programmed Cell Death)
Analytic Questions
CHAPTER 16: Cancer
16.1 Basic Properties of a Cancer Cell
16.2 The Causes of Cancer
16.3 Cancer: A Genetic Disorder
16.4 Engineering Linkage: Therapeutic Radiation
16.5 Green Cells: Plant-Based Chemotherapies
16.6 Strategies for Combating Cancer
CHAPTER 17: The Immune Response
17.1 An Overview of the Immune Response
17.2 Green Cells: The Plant Immune System
17.3 The Clonal Selection Theory as It Applies to B Cells
17.4 T Lymphocytes: Activation and Mechanism of Action
17.5 Selected Topics on the Cellular and Molecular Basis of Immunity
17.6 Engineering Linkage: Adoptive T-Cell Therapy
17.7 Signal Transduction Pathways in Lymphocyte Activation
CHAPTER 18: Techniques in Cell and Molecular Biology
18.1 The Light Microscope
18.2 Transmission Electron Microscopy
18.3 Scanning Electron and Atomic Force Microscopy
18.4 The Use of Radioisotopes
18.5 Cell Culture
18.6 The Fractionation of a Cell’s Contents by Differential Centrifugation
18.7 Isolation, Purification, and Fractionation of Proteins
18.8 Determining the Structure of Proteins and Multisubunit Complexes
18.9 Fractionation of Nucleic Acids
18.10 Nucleic Acid Hybridization
18.11 Chemical Synthesis of DNA
18.12 Recombinant DNA Technology
18.13 Enzymatic Amplification of DNA by PCR
18.14 DNA Sequencing
18.15 DNA Libraries
18.16 DNA Transfer into Eukaryotic Cells and Mammalian Embryos
18.17 Gene Editing and Silencing
18.18 The Use of Antibodies
Additional Readings
Glossary
Index
End User License Agreement
List of Tables
CHAPTER 1
TABLE 1.1 A Comparison of Prokaryotic and Eukaryotic Cells
TABLE 1.2 Number and Biomass of Prokaryotes in the World
CHAPTER 2
TABLE 2.1 Strengths of Acids and Bases
TABLE 2.2 Functional Groups
CHAPTER 3
TABLE 3.1 Thermodynamics of the Water–Ice Transformation
TABLE 3.2 Relation between and at
TABLE 3.3 Catalytic Activity of a Variety of Enzymes
TABLE 1 Antibiotics in Clinical Use and Modes of Resistance
CHAPTER 4
TABLE 4.1 Lipid Compositions of Some Biological Membranes
TABLE 4.2 Melting Points of the Common 18-Carbon Fatty Acids
TABLE 4.3 Ion Concentrations Inside and Outside a Typical Mammalian Cell
CHAPTER 5
TABLE 5.1 Standard Redox Potentials of Selected Half-Reactions
CHAPTER 7
TABLE 7.1 Classification of Integrin Receptors Based on Recognition of RGD Seque...
CHAPTER 8
TABLE 1 Sphingolipid Storage Diseases
TABLE 8.1 A Sampling of Lysosomal Enzymes
CHAPTER 9
TABLE 9.1 Properties of Microtubules, Intermediate Filaments, and Actin Filaments
TABLE 9.2 Properties and Distribution of the Major Mammalian Intermediate Filame...
CHAPTER 10
TABLE 10.1 Seven Traits of Mendel’s Pea Plants
CHAPTER 11
TABLE 11.1 Eukaryotic Nuclear RNA Polymerases
CHAPTER 13
TABLE 13.1 Some of the Proteins Required for Replication
CHAPTER 15
TABLE 15.1 Examples of Physiologic Processes Mediated by GPCRs and Heterotrimeric G Proteins
TABLE 1 Effects of Combinations of Hormones, at Supramaximal Concentrations, on Adenylyl...
TABLE 2 Purification of the Regulatory Component of Adenylyl Cyclase from Rabbit Liver
TABLE 1 Human Diseases Linked to the G Protein Pathway
TABLE 15.2 Summary of Cellular Responses Elicited by Adding IP3 to Either Permeabilized or...
TABLE 15.3 Examples of Hormone-Induced Responses Mediated by cAMP
TABLE 15.4 Examples of Mammalian Proteins Activatedby Ca2+
CHAPTER 16
TABLE 1 Characterization of the Polymerase Product
TABLE 16.1 Tumor-Suppressor Genes
TABLE 16.2 Examples of Small-Molecule Targeted Therapies That Either Have Been FDA-Approved...
CHAPTER 17
TABLE 17.1 Selected Cytokines
TABLE 17.2 Classes of Human Immunoglobulins
TABLE 1 Cytotoxic Activity of Spleen Cells from Various Strains of Mice Injected 7 Days ...
TABLE 2 Inhibition of Antigen Presentation with NH4Cl and Chloroquine
CHAPTER 18
TABLE 18.1 Properties of a Variety of Radioisotopes Used in Biological Research
List of Illustrations
CHAPTER 1
FIGURE 1.1 The discovery of cells. (a) One of Robert Hooke’s more ornate compound (double-l...
FIGURE 1.2 HeLa cells, such as the ones pictured here, were the first human cells to be kep...
FIGURE 1.3 Levels of cellular and molecular organization. The brightly colored photograph o...
FIGURE 1.4 Cell reproduction. This mammalian oocyte has recently undergone a highly unequal...
FIGURE 1.5 Acquiring energy. A living cell of the filamentous alga Spirogyra. The ribbon-li...
FIGURE 1.6 Self-regulation. The left panel depicts the normal development of a sea urchin i...
FIGURE 1.7 Cellular activities are often analogous to this Rube Goldberg machine in which o...
FIGURE 1.8 The structure of cells. Schematic diagrams of a “generalized” bacterial (a), pla...
FIGURE 1.9 Earth’s biogeologic clock. A portrait of the past five billion years of Earth’s ...
FIGURE 1.10 The structure of a eukaryotic cell. This epithelial cell lines the male reproduc...
FIGURE 1.11 The cytoplasm of a eukaryotic cell is a crowded compartment. This colorized elec...
FIGURE 1.12 Cell division in eukaryotes requires the assembly of an elaborate chromosome-sep...
FIGURE 1.13 Bacterial conjugation. Electron micrograph showing a conjugating pair of bacteri...
FIGURE 1.14 The difference between prokaryotic and eukaryotic flagella. (a) The bacterium Sa...
FIGURE 1.15 Cyanobacteria. (a) Electron micrograph of a cyanobacterium showing the cytoplasm...
FIGURE 1.16 Vorticella, a complex ciliated protist. A number of these unicellular organisms ...
FIGURE 1.17 Pathways of cell differentiation. A few of the types of differentiated cells pre...
FIGURE 1 An adult muscle stem cell. (a) A portion of a muscle fiber, with its many nuclei...
FIGURE 2 Embryonic stem cells; their isolation and potential use. (a) Micrograph of a mam...
FIGURE 3 Steps taken to generate induced pluripotent stem (iPS) cells for use in correcti...
FIGURE 1.18 Six model organisms. (a) Escherichia coli is a rod-shaped bacterium that lives i...
FIGURE 1.19 Relative sizes of cells and cell components. These structures differ in size by ...
FIGURE 1.20 The synthetic biologist’s toolkit of the future? Such a toolkit would presumably.
FIGURE 1.21 Tobacco mosaic virus (TMV). (a) Model of a portion of a TMV particle. The protei...
FIGURE 1.22 Virus diversity. The structures of (a) an adenovirus, (b) a human immunodeficien...
FIGURE 1.23 A virus infection. (a) Micrograph showing a late stage in the infection of a bac...
FIGURE 1 A model depicting stages in the evolution of eukaryotes. Starting from a prokary...
FIGURE 2 Prokaryotes with complex internal membrane systems. Electron micrograph of Gemma...
FIGURE 3 Domains of life. (a) A phylogenetic tree based on rRNA sequence comparisons show...
FIGURE 1.24 Evolutionary experiments in multicelluarity. (a) Volvox carteri, a multicellular...
FIGURE 1.25 Replacement blood vessel that incorporates living cells, created using tissue en...
CHAPTER 2
FIGURE 2.1 A representation of the arrangement of electrons in a number of
common atoms. El...
FIGURE 2.2 Gamma knife machine. Left panel shows hemispherical array of cobalt ingots, each...
FIGURE 2.3 The dissolution of a salt crystal. When placed in water, the Na+ and Cl ions of...
FIGURE 2.4 Noncovalent ionic bonds play an important role in holding the protein molecule o...
FIGURE 2.5 Hydrogen bonds form between a bonded electronegative atom, such as nitrogen or o...
FIGURE 2.6 In a hydrophobic interaction, the nonpolar (hydrophobic) molecules are forced in...
FIGURE 2.7 Van der Waals forces. (a) As two atoms approach each other, they experience a we...
FIGURE 2.8 Hydrogen bond formation between neighboring water molecules. Each H atom of the ...
FIGURE 2.9 The importance of water in protein structure. The water molecules (each with a s...
FIGURE 2.10 The structure of cholesterol illustrates how carbon atoms (represented by the bl...
FIGURE 2.11 Monomers and polymers; polymerization and hydrolysis. (a) Polysaccharides, prote...
FIGURE 2.12 An overview of the types of biological molecules that make up various cellular s...
FIGURE 2.13 Fertilizers, both organic and chemical, provide elements such as nitrogen and ph...
FIGURE 2.14 The structures of sugars. (a) Straight-chain formula of fructose, a ketohexose [...
FIGURE 2.15 Stereoisomerism of glyceraldehyde. (a) The four groups bonded to a carbon atom (...
FIGURE 2.16 Aldotetroses. Because they have two asymmetric carbon atoms, aldotetroses can ex...
FIGURE 2.17 Formation of an α- and β-pyranose. When a molecule of glucose undergoes self-rea...
FIGURE 2.18 Disaccharides. Sucrose and lactose are two of the most common disaccharides. Suc...
FIGURE 2.19 Three polysaccharides with identical sugar monomers but dramatically different p...
FIGURE 2.20 Chitin is the primary component of the outer skeleton of this grasshopper....
FIGURE 2.21 Fats and fatty acids. (a) The basic structure of a triacylglycerol (also called ...
FIGURE 2.22 Soaps consist of fatty acids. In this schematic drawing of a soap micelle, the n...
FIGURE 2.23 The structure of steroids. All steroids share the basic four-ring skeleton. The ...
FIGURE 2.24 The phospholipid phosphatidylcholine. The molecule consists of a glycerol backbo...
FIGURE 2.25 Four examples of the thousands of biological structures composed predominantly o...
FIGURE 2.26 Amino acid structure. Ball-and-stick model (a) and chemical formula (b) of a gen...
FIGURE 2.27 Amino acid stereoisomerism. Because the α-carbon of all amino acids except glyci...
FIGURE 2.28 The chemical structure of amino acids. These 20 amino acids represent those most...
FIGURE 2.29 The ionization of charged, polar amino acids. (a) The side chain of glutamic aci...
FIGURE 2.30 Disposition of hydrophilic and hydrophobic amino acid residues in the soluble pr...
FIGURE 2.31 Scanning electron micrograph of a red blood cell from a person with sickle cell ...
FIGURE 2.32 The alpha helix. (a) The helical path around a central axis taken by the polypep...
FIGURE 2.33 The β sheet. (a) Each polypeptide of a β sheet assumes an extended but pleated c...
FIGURE 2.34 A ribbon model of ribonuclease. The regions of α helix are depicted as spirals a...
FIGURE 2.35 An X-ray diffraction pattern of myoglobin. The pattern of spots is produced as a...
FIGURE 2.36 NMR spectroscopy reveals tertiary structure without crystalization. (a) An NMR s...
FIGURE 2.37 The three-dimensional structure of myoglobin. Most of the amino acids in the ter...
FIGURE 2.38 Types of noncovalent bonds maintaining the conformation of proteins....
FIGURE 2.39 Different sequence, similar structure. Two proteins, actin and MreB, with primar...
FIGURE 2.40 Proteins are built of structural units, or domains. The mammalian enzyme phospho...
FIGURE 2.41 Dynamic movements within the enzyme acetylcholinesterase. A portion of the enzym...
FIGURE 2.42 Proteins with quaternary structure. (a) Ribbon diagram of transforming growth fa...
FIGURE 2.43 Pyruvate dehydrogenase: a multiprotein complex. (a) Threedimensional reconstruc...
FIGURE 2.44 Protein–protein interactions. (a) A model illustrating the complementary molecul...
FIGURE 2.45 Denaturation and refolding of ribonuclease. A native ribonuclease molecule (with...
FIGURE 2.46 Two alternate pathways by which a newly synthesized or denatured protein could a...
FIGURE 2.47 Along the folding pathway. The image on the left shows the native tertiary struc...
FIGURE 1 A contrast in structure. (a) Tertiary structure of the normal (PrPC) protein as...
FIGURE 2 Alzheimer’s disease. (a) The defining characteristics of brain tissue from a per...
FIGURE 3 Formation of the Aβ peptide. The Aβ peptide is carved from the amyloid precursor...
FIGURE 4 A neuroimaging technique that reveals the presence of amyloid in the brain. Thes...
FIGURE 2.48 The role of molecular chaperones in encouraging protein folding. The steps are d...
FIGURE 1 An early model of the GroEL complex built according to data from electron micros...
FIGURE 2 Reconstructions of GroEL based on high-resolution electron micrographs of specim...
FIGURE 3 Conformational change in GroEL. (a) The model on the left shows a surface view o...
FIGURE 4 A schematic illustration of the proposed steps that occur during the GroEL-GroES...
FIGURE 2.49 Identifying proteins by mass spectrometry. A protein is isolated from a cell or ...
FIGURE 2.50 A network of protein–protein interactions. Each red line represents an interacti...
FIGURE 2.51 Protein–protein interactions of hub proteins. (a) The enzyme RNA polymerase II, ...
FIGURE 2.52 The computational design of a protein that is capable of binding specifically to...
FIGURE 2.53 Development of a protein-targeting drug, such as Gleevec. (a) Typical steps in d...
FIGURE 2.54 Distribution of polar, charged amino acid residues in the enzyme malate dehydrog...
FIGURE 2.55 The dramatic effect on conformation that can result from a single amino acid sub...
FIGURE 2.56 Nucleotides and nucleotide strands of RNA. (a) Nucleotides are the monomers from...
FIGURE 2.57 Nitrogenous bases in nucleic acids. Of the four standard bases found in RNA, ade...
FIGURE 2.58 RNAs can assume complex shapes. (a) This ribosomal RNA is an integral component ...
FIGURE 2.59 Reconstruction of a ribosome from the cytoplasm of a wheat germ cell. This recon...
FIGURE 2.60 Example of a phase-separated compartment formed by Whi3, an RNA-binding protein ...
CHAPTER 3
FIGURE 3.1 Examples of energy transduction. (a) Conversion of mechanical energy to electric...
FIGURE 3.2 A change in a system’s internal energy. In this example, the system is defined a...
FIGURE 3.3 Events are accompanied by an increase in the entropy of the universe. (a) A suga...
FIGURE 3.4 When water freezes, its entropy decreases because the water molecules of ice exi...
FIGURE 3.5 ATP hydrolysis. Adenosine triphosphate (ATP) is hydrolyzed as part of many bioch...
FIGURE 3.6 A few roles for ATP hydrolysis. In the cell, energy from ATP hydrolysis may be u...
FIGURE 3.7 Steady state versus equilibrium. (a) As long as this ameba can continue to take ...
FIGURE 3.8 Activation energy and enzymatic reactions. Even though the formation of glucose ...
FIGURE 3.9 The effect of lowering activation energy on the rate of a reaction. The bell-sha...
FIGURE 3.10 Formation of an enzyme–substrate complex. Schematic drawing of the reaction cata...
FIGURE 3.11 The active site of an enzyme. (a) Diagrammatic representation of the active site...
FIGURE 3.12 Three mechanisms by which enzymes accelerate reactions: (a)
maintaining precise ...
FIGURE 3.13 Diagrammatic representation of the catalytic mechanism of chymotrypsin. The reac...
FIGURE 3.14 An example of induced fit. When a glucose molecule binds to the enzyme hexokinas...
FIGURE 3.15 Electron density map of a single hydrogen bond (green-dotted line). This map sho...
FIGURE 3.16 Myoglobin: the movie. In this example of time-resolved X-ray crystallography, th...
FIGURE 3.17 The relationship between the rate (velocity) of an enzyme-catalyzed reaction and...
FIGURE 3.18 A Lineweaver-Burk plot of the reciprocals of velocity and substrate concentratio...
FIGURE 3.19 Dependence of the rate of an enzyme-catalyzed reaction on (a) pH, and (b) temper...
FIGURE 3.20 Competitive inhibition. Because of their molecular similarity, competitive inhib...
FIGURE 3.21 The effects of inhibitors on enzyme kinetics. The effect of both competitive and...
FIGURE 3.22 Three stages of metabolism. The catabolic pathways (green arrows rightward) conv...
FIGURE 3.23 The oxidation state of a carbon atom depends on the other atoms to which it is b...
FIGURE 3.24 The steps of glycolysis....
FIGURE 3.25 Free-energy profile of glycolysis in cardiac muscle tissue. The step numbers cor...
FIGURE 3.26 The transfer of energy during chemical oxidation. The oxidation of glyceraldehyd...
N A D +
FIGURE 3.27 The structure of and its reduction to NADH. When the 2′ OH of the ribose mo...
FIGURE 3.28 Ranking compounds by phosphate transfer potential. Those phosphorylated compound...
FIGURE 3.29 Fermentation. Most cells carry out aerobic respiration, which depends on molecul...
FIGURE 3.30 Feedback inhibition. Starting with two substrates, A and B, a series of three en...
FIGURE 3.31 Glycolysis versus gluconeogenesis. Whereas most of the reactions are the same in...
FIGURE 3.32 In the silk plant (Albizia julibrissin), leaves extend to collect light
FIGURE 3.33 Brain tumor visualized using PET scanning to detect accumulation of 18F-FDG. Red...
CHAPTER 4
FIGURE 4.1 The trilaminar appearance of membranes. (a) Electron micrograph showing the thre...
FIGURE 4.2 A summary of membrane functions in a plant cell. (1) An example of membrane comp...
FIGURE 4.3 The plasma membrane contains a lipid bilayer. (a) Calculating the surface area o...
FIGURE 4.4 Plasma membranes as lipids plus proteins. (a) A representation of the plasma mem...
FIGURE 4.5 The myelin sheath. Electron micrograph of a nerve cell axon surrounded by a myel...
FIGURE 4.6 The chemical structures of membrane lipids. (a) The structures of phosphoglyceri...
FIGURE 4.7 The cholesterol molecules (shown in green) of a lipid bilayer are oriented with ...
FIGURE 4.8 The dynamic properties of the plasma membrane. (a) The leading edge of a moving ...
FIGURE 4.9 Liposomes. A schematic diagram of a stealth liposome containing a hydrophilic po...
FIGURE 4.10 The asymmetric distribution of phospholipids (and cholesterol) in the plasma mem...
FIGURE 4.11 Two types of linkages that join sugars to a polypeptide chain. The N-glycosidic ...
FIGURE 4.12 Blood group antigens. Whether a person has type A, B, AB, or O blood is determin...
FIGURE 4.13 Three classes of membrane proteins. (a) Integral proteins typically contain one ...
FIGURE 4.14 The interactions between membrane proteins and lipids. (a) Aquaporin is a membra...
FIGURE 4.15 Freeze fracture: a technique for investigating cell membrane structure. (a) When...
FIGURE 4.16 Solubilization of membrane proteins with detergents. The nonpolar ends of the de...
FIGURE 4.17 Structure of integral proteins. (a) Tertiary structure of the photosynthetic rea...
FIGURE 4.18 Glycophorin A, an integral protein with a single transmembrane domain. The singl...
FIGURE 4.19 Accommodating various amino acid residues within transmembrane helices. (a) In t...
FIGURE 4.20 Hydropathy plot for glycophorin A, a single membrane-spanning protein. Hydrophob...
FIGURE 4.21 An experiment employing site-directed mutagenesis to learn about dynamic changes...
FIGURE 4.22 Use of EPR spectroscopy to monitor changes in conformation of a bacterial ion ...
FIGURE 4.23 The structure of the lipid bilayer depends on the temperature. The bilayer shown...
FIGURE 4.24 Lipid rafts. (a) Image of the upper surface of an artificial lipid bilayer conta...
FIGURE 4.25 The possible movements of phospholipids in a membrane. The figure shows the type...
FIGURE 4.26 The use of cell fusion to reveal the mobility of membrane proteins. (a) Outline ...
FIGURE 4.27 Measuring the diffusion rates of membrane proteins by fluorescence recovery afte...
FIGURE 4.28 Patterns of movement of integral membrane proteins. Depending on the cell type a...
FIGURE 4.29 Experimental demonstration that diffusion of phospholipids within the plasma mem...
FIGURE 4.30 Differentiated functions of the plasma membrane of an epithelial cell. The apica...
FIGURE 4.31 Differentiation of the mammalian sperm plasma membrane as revealed by fluorescen...
FIGURE 4.32 The plasma membrane of the human erythrocyte. (a) Scanning electron micrograph o...
FIGURE 4.33 Four basic mechanisms by which solute molecules move across membranes. The relat...
FIGURE 4.34 The relationship between partition coefficient and membrane permeability. In thi...
FIGURE 4.35 The effects of differences in the concentration of solutes on opposite sides of ...
FIGURE 4.36 The effects of osmosis on a plant cell. (a) Aquatic plants living in fresh water...
FIGURE 4.37 Passage of water molecules through an aquaporin channel. (a) Snapshot from a mol...
FIGURE 4.38 Measuring ion channel conductance by patch-clamp recording. (a) In this techniqu... K +
FIGURE 4.39 Three-dimensional structure of the bacterial KcsA channel and the selection of ..
FIGURE 4.40 Schematic illustration of the hinge-bending model for the opening of the KcsA ch...
FIGURE 4.41 The structure of one subunit of a eukaryotic, voltage-gated K+ channel. A two-di...
FIGURE 4.42 Three-dimensional structure of a voltage-gated mammalian K+ channel. (a) The cry...
FIGURE 4.43 Conformational states of a voltage-gated ion channel. (a) Threedimensional m...
FIGURE 1 The electric organs of Torpedo consist of stacks of modified neuromuscular junct...
FIGURE 2 Steps used in the isolation of the nAChR. (a) Structure of a synthetic compound,...
FIGURE 3 The top portion of the figure shows an SDS–polyacrylamide gel following electrop...
FIGURE 4 Electron micrograph of negatively stained, receptor-rich membranes from the elec...
FIGURE 5 (a) An electron density map of a slice through nAChR obtained by analyzing elect...
FIGURE 6 Ribbon drawings illustrating the changes that occur within the nAChR conducting ...
FIGURE 4.44 Facilitated diffusion. A schematic model for the facilitated diffusion of glucos...
FIGURE 4.45 The kinetics of facilitated diffusion compared to that of simple physical diffus...
FIGURE 4.46 The Na+/K+-ATPase. (a) Simplified schematic model of the transport cycle as desc...
FIGURE 4.47 Control of acid secretion in the stomach. In the resting state, the H+ K+-ATPase...
FIGURE 1 An explanation for the debilitating effects on lung function from the absence of...
FIGURE 4.48 Bacteriorhodopsin: a light-driven proton pump. The protein contains seven membra...
FIGURE 4.49 Secondary transport: the use of energy stored in an ionic gradient. The ..
FIGURE 4.50 A schematic model of the transport cycle of a secondary transporter. Four differ...
FIGURE 4.51 The structure of a nerve cell. (a) Schematic drawing of a simple K + . K + / Na + /K + .
neuron with a m...
FIGURE 4.52 Measuring a membrane’s resting potential. A potential is measured when a differe...
FIGURE 4.53 Formation of an action potential. (a) Time 1, upper left box: The membrane in th...
FIGURE 4.54 Propagation of an impulse results from the local flow of ions. An action potenti...
FIGURE 4.55 Saltatory conduction. During saltatory conduction, only the membrane in the noda...
FIGURE 4.56 The neuromuscular junction is the site where branches from a motor axon form syn...
FIGURE 4.57 The sequence of events during synaptic transmission with acetylcholine as the ne...
FIGURE 4.58 Action potentials in plant cells. (a) The Venus flytrap catches insects in a tra...
FIGURE 4.59 Cochlear implant showing a flexible silicone elastomer containing platinum elect...
CHAPTER 5
FIGURE 5.1 Mitochondria. (a) A living fibroblast viewed with a phase-contrast microscope. M...
FIGURE 5.2 Mitochondrial fusion and fission. (a) The dynamic nature of these organelles is ...
FIGURE 5.3 The structure of a mitochondrion. (a) Scanning electron micrograph of a macerate...
FIGURE 5.4 Porins. Gram-negative bacteria have a lipid-containing outer membrane outside of...
FIGURE 5.5 An overview of carbohydrate metabolism in eukaryotic cells. The reactions of gly...
FIGURE 5.6 An overview of glycolysis showing some of the key steps. These include the two r...
FIGURE 5.7 The tricarboxylic acid (TCA) cycle. The cycle begins with the condensation of ox...
FIGURE 5.8 Catabolic pathways generate compounds that are fed into the TCA cycle. (a) Oxida...
FIGURE 5.9 The glycerol phosphate shuttle. In the glycerol phosphate shuttle, electrons are...
FIGURE 5.10 A summary of the process of oxidative phosphorylation. In the first step of the ...
FIGURE 1 Skeletal muscles contain a mix of fast-twitch (or type II) fibers (darkly staine...
FIGURE 5.11 Measuring the standard oxidation–reduction (redox) potential. The sample half-ce...
FIGURE 5.12 Structures of the oxidized and reduced forms of three types of electron carriers...
FIGURE 5.13 Iron-sulfur centers. Structures of (a) and (b) ironsulfur cen...
FIGURE 5.14 The arrangement of several carriers in the electron-transport chain. The diagram...
FIGURE 5.15 Experimental use of inhibitors to determine the sequence of carriers in the elec...
FIGURE 5.16 Electron-tunneling pathways for the yeast cytochrome -cytochrome c peroxidase c...
FIGURE 5.17 The electron-transport chain of the inner mitochondrial membrane. The respirator...
FIGURE 5.18 Experimental demonstration that cytochrome oxidase is a proton pump. When purifi...
FIGURE 5.19 Structure and a proposed mechanism of action of complex I of the respiratory cha...
FIGURE 5.20 A summary of cytochrome oxidase activity. A model showing the flow of electrons ...
FIGURE 5.21 Pulse oximeter. Red and infrared light emitting diodes transmit light through th...
FIGURE 5.22 Visualizing the proton-motive force. Fluorescence micrograph of a cultured cell ...
FIGURE 5.23 The machinery for ATP synthesis. Three-dimensional electron micrograph of isolat...
FIGURE 5.24 An experiment to drive ATP formation in membrane vesicles reconstituted with the...
FIGURE 5.25 The structure of ATP synthase. (a) Schematic diagram of the ATP synthase from...
FIGURE 5.26 Visualizing the oligomeric ring of a chloroplast ATP synthase. Atomic force mi...
FIGURE 5.27 The structural basis of catalytic site conformation. (a) A section through the F...
FIGURE 5.28 The binding change mechanism for ATP synthesis. (a) Schematic drawing showing ch...
FIGURE 5.29 Direct observation of rotational catalysis. (a) To carry out the experiment, a m...
FIGURE 5.30 A model in which proton diffusion is coupled to the rotation of the ring of the...
c c
FIGURE 5.31 Summary of the major activities during aerobic respiration in a mitochondrion.
FIGURE 5.32 The structure and function of peroxisomes. (a) Electron micrograph of a section ...
FIGURE 1 Mitochondrial abnormalities in skeletal muscle. (a) Ragged-red fibers. These deg...
FIGURE 2 A premature aging phenotype caused by increased mutations in mtDNA. The photogra...
FIGURE 5.33 Glyoxysome localization within plant seedlings. Light micrograph of a section th...
CHAPTER 6
FIGURE 6.1 Photosynthetic green sulfur bacteria are present as a ring of peripheral cells t...
FIGURE 6.2 The functional organization of a leaf. The section of the leaf shows several lay...
FIGURE 6.3 The internal structure of a chloroplast. (a) Transmission electron micrograph th...
FIGURE 6.4 Thylakoid membranes. Electron micrograph of a section through a portion of a chl...
FIGURE 6.5 An overview of the energetics of photosynthesis and aerobic respiration....
FIGURE 6.6 The structure of chlorophyll The molecule consists of a porphyrin ring (which...
FIGURE 6.7 Absorption spectrum for several photosynthetic pigments of higher plants. The ba...
FIGURE 6.8 Action spectrum for photosynthesis. The action spectrum (red line) indicates the...
FIGURE 6.9 Chromoplasts within the cells of a red pepper....
FIGURE 6.10 The transfer of excitation energy. Energy is transferred randomly through a netw...
FIGURE 6.11 An overview of the flow of electrons during the light-dependent reactions of pho...
FIGURE 6.12 The functional organization of photosystem II. (a) A schematic model of the huge...
FIGURE 6.13 Plastoquinone. The acceptance of two electrons and two protons reduces PQ (plast...
FIGURE 6.14 Measuring the kinetics of release. The plot shows the response by isolated ch...
FIGURE 6.15 Electron transport between PSII and PSI. The flow of a pair of electrons is indi...
FIGURE 6.16 The functional organization of photosystem I. The path taken by electrons is ind...
FIGURE 6.17 Summary of the light-dependent reactions. (a) Three-dimensional structures of th...
FIGURE 6.18 Simplified scheme for cyclic photophosphorylation. Absorption of light by PSI ex...
FIGURE 6.19 Chromatogram showing the results of an experiment in which algal cells were incu...
FIGURE 6.20 Converting into carbohydrate. (a) The reaction catalyzed by ribulose bisphos...
FIGURE 6.21 An overview of the various stages of photosynthesis....
FIGURE 6.22 Redox control of the Calvin cycle. In the light, ferredoxin is reduced, and a fr...
FIGURE 6.23 The reactions of photorespiration. Rubisco can catalyze two different reactions ...
FIGURE 6.24 The cellular basis of photorespiration. Electron micrograph of a portion of a le...
FIGURE 6.25 Structure and function in plants. Electron micrograph of a transverse section...
FIGURE 6.26 Aminolevulinic acid (ALA) is converted to the photosensitizing agent protoporphy...
CHAPTER 7
FIGURE 7.1 An overview of cell organization into tissues and of their interactions with one...
FIGURE 7.2 The glycocalyx. (a) The basal surface of an ectodermal cell of an early chick em...
FIGURE 7.3 The extracellular matrix (ECM) of cartilage cells. (a) Scanning electron microgr...
FIGURE 7.4 The basement membrane (basal lamina). (a) Scanning electron micrograph of human ...
FIGURE 7.5 An overview of the macromolecular organization of the extracellular matrix. The ...
FIGURE 7.6 The structure of collagen I. This figure depicts several levels of organization ...
FIGURE 7.7 The corneal stroma consists largely of layers of collagen fibrils of uniform dia...
FIGURE 7.8 The type IV collagen network of the basement membrane. Electron micrograph of a ...
FIGURE 7.9 The structure of a cartilage-type proteoglycan complex. (a) Schematic representa...
FIGURE 7.10 Structure of fibronectin and its importance during embryonic development. (a) A ...
FIGURE 7.11 The role of cell migration during embryonic development. (a) A summary of some o...
FIGURE 7.12 A model of the basement membrane scaffold. Basement membranes contain two networ...
FIGURE 7.13 Fluorescence microscopy image of a brain organoid. Neurons, labeled green, are e...
FIGURE 7.14 Integrin conformations. (a) Schematic drawing of a complete integrin in the bent...
FIGURE 7.15 The role of integrins in platelet aggregation. (a) Blood clots form when platele...
FIGURE 7.16 Steps in the process of cell spreading. Scanning electron micrographs showing th...
FIGURE 7.17 Focal adhesions are sites where cells adhere to their substratum and transmit si...
FIGURE 7.18 Experimental demonstration of forces exerted by focal adhesions. Scanning-electr...
FIGURE 7.19 Measuring the force necessary to activate integrins. In this experiment, an inte...
FIGURE 7.20 Hemidesmosomes. Hemidesmosomes are differentiated sites at the basal surfaces of...
FIGURE 7.21 Experimental demonstration of cell–cell recognition. When cells from different p...
FIGURE 7.22 Overview of cell adhesion molecules. (a) Selectins. All three types of known sel...
FIGURE 1 Steps in the movement of neutrophils from the bloodstream during inflammation. T...
FIGURE 2 Steps leading to the metastatic spread of an epithelial cancer (a carcinoma). (a...
FIGURE 7.23 Cadherins and morphogenesis. (a) During gastrulation, cells in the upper layer o...
FIGURE 7.24 Cadherins form tip links of stereocilia. Cadherin 23 and protocadherin 15 form t...
FIGURE 7.25 An intercellular junctional complex. (a) Schematic diagram showing the junctiona...
FIGURE 7.26 Schematic model of the molecular architecture of an adherens junction and desomo...
FIGURE 7.27 Specialized cadherins, called desmoglein and desmocollin, bind to one another in...
FIGURE 7.28 Tight junctions. (a) Electron micrograph of a section through the apical region ...
FIGURE 7.29 The molecular composition of tight junction strands. Electron micrograph of a fr...
FIGURE 7.30 Gap junctions. (a) Electron micrograph of a section through a gap junction perpe...
FIGURE 7.31 Results of an experiment demonstrating the passage of lowmolecular-weight solut...
FIGURE 1 Micrograph showing recording microelectrodes being inserted into living cells of...
FIGURE 2 Calcium waves induced by mechanical stimulation in (a) a control culture of rat ...
FIGURE 3 (a) When a small mass of C6 glioma cells is implanted into the brain of a rat, t...
FIGURE 7.32 Plasmodesmata. (a) Electron micrograph of a section through a plasmodesma of a f...
FIGURE 7.33 Tunneling nanotubes. Scanning electron micrograph showing two cultured neuroendo...
FIGURE 7.34 The plant cell wall. (a) Electron micrograph of a plant cell surrounded by its c...
FIGURE 7.35 Synthesis of plant cell wall macromolecules. (a) Freeze-fracture replica of the ...
CHAPTER 8
FIGURE 8.1 Membrane-bound compartments of the cytoplasm. The cytoplasm of this root cap cel...
FIGURE 8.2 An overview of the biosynthetic/secretory and endocytic pathways that unite endo...
FIGURE 8.3 Autoradiography reveals the sites of synthesis and subsequent transport of secre...
FIGURE 8.4 The use of green fluorescent protein (GFP) reveals the movement of proteins with...
FIGURE 8.5 Microscopic images of a mammalian cell expressing multiple fluorophores to visua...
FIGURE 8.6 Isolation of a microsomal fraction by differential centrifugation. (a) When a ce...
FIGURE 8.7 Formation of coated vesicles in a cell-free system. Electron micrograph of a lip...
FIGURE 8.8 The use of genetic mutants in the study of secretion. (a) The first leg of the b...
FIGURE 8.9 Inhibition of gene expression with RNA interference. (a) A control
Drosophila S2...
FIGURE 8.10 The rough endoplasmic reticulum (RER). (a) Schematic diagram showing the stacks ...
FIGURE 8.11 The smooth ER (SER). Electron micrograph of a Leydig cell from the testis showin...
FIGURE 8.12 Polarized structure of a secretory cell. (a) Drawing of a mucussecreting goblet...
FIGURE 8.13 A schematic model of the synthesis of a secretory protein (or a lysosomal enzyme...
FIGURE 8.14 A schematic model for the synthesis of an integral membrane protein that contain...
FIGURE 8.15 Maintenance of membrane asymmetry. As each protein is synthesized in the RER, it...
FIGURE 8.16 Modifying the lipid composition of membranes. (a) Histogram indicating the perce...
FIGURE 8.17 Steps in the synthesis of the core portion of an N-linked oligosaccharide in the...
FIGURE 8.18 Quality control: ensuring that misfolded proteins do not proceed forward. Based ...
FIGURE 8.19 A model of the mammalian unfolded protein response (UPR). The ER contains transm...
FIGURE 8.20 Visualizing membrane traffic with the use of a fluorescent tag. This series of p...
FIGURE 8.21 The Golgi complex. (a) Schematic model of a portion of a Golgi complex from an e...
FIGURE 8.22 Regional differences in membrane composition across the Golgi stack. (a) Reduced...
FIGURE 8.23 Steps in the glycosylation of a typical mammalian N-linked oligosaccharide in th...
FIGURE 8.24 The dynamics of transport through the Golgi complex. (a) In the vesicular transp...
FIGURE 8.25 Coated vesicles. These electron micrographs show the membranes of these vesicles...
FIGURE 8.26 Proposed movement of materials by vesicular transport between membranous compart...
FIGURE 8.27 Proposed roles of the COPII coat proteins in generating membrane curvature, asse...
FIGURE 8.28 Structure of vesicle coats. (a) Molecular model of the outer Sec13–Sec31 cage of...
FIGURE 8.29 Retrieving ER proteins. Resident proteins of the ER contain amino
acid sequences...
FIGURE 8.30 Targeting lysosomal enzymes to lysosomes. (a) Lysosomal enzymes are recognized b...
FIGURE 8.31 The formation of clathrin-coated vesicles at the TGN. Clathrincoated vesicles t...
FIGURE 1 Lysosomal storage disorders. Electron micrograph of a section through a portion ...
FIGURE 8.32 Proposed steps in the targeting of transport vesicles to target membranes. (a) A...
FIGURE 8.33 A model of the interactions between v- and t-SNAREs leading to membrane fusion a...
FIGURE 8.34 Illustration of extracellular vesicles. Extracellular vesicles can vary in size ...
FIGURE 8.35 Lysosomes. Portion of a phagocytic Kupffer cell of the liver showing at least 10...
FIGURE 8.36 Autophagy. Electron micrograph of a mitochondrion and peroxisome enclosed in a d...
FIGURE 8.37 A summary of the autophagic pathway. The steps are described in the text....
FIGURE 8.38 Plant cell vacuoles. (a) Each of the cylindrical leaf cells of the aquatic plant...
FIGURE 8.39 Receptor-mediated endocytosis. This sequence of micrographs shows the steps in t...
FIGURE 8.40 Coated pits. (a) Electron micrograph of a replica formed on the extracellular su...
FIGURE 8.41 Clathrin triskelions. Electron micrograph of a metal-shadowed preparation of cla...
FIGURE 8.42 Molecular organization of a coated vesicle. (a) Schematic drawing of the surface...
FIGURE 8.43 The role of dynamin in the formation of clathrin-coated vesicles. (a) The clathr...
FIGURE 8.44 A structural model depicting the changes in protein conformation that occur upon...
FIGURE 1 (a) A handmade model of an “empty basket” that would form the surface lattice of...
FIGURE 2 HMG CoA reductase activity in normal fibroblasts was measured following addition...
FIGURE 3 Fibroblast cells from either a control subject (closed circles) or a patient wit...
