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Boundless Biology: 15.4: RNA Processing in Eukaryotes

Boundless Biology
15.4: RNA Processing in Eukaryotes
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table of contents
  1. 1: The Study of Life
    1. 1.1: The Science of Biology
      1. 1.1.0: Introduction to the Study of Biology
      2. 1.1.1: Scientific Reasoning
      3. 1.1.2: The Scientific Method
      4. 1.1.3: Basic and Applied Science
      5. 1.1.4: Publishing Scientific Work
      6. 1.1.5: Branches and Subdisciplines of Biology
    2. 1.2: Themes and Concepts of Biology
      1. 1.2.0: Properties of Life
      2. 1.2.1: Levels of Organization of Living Things
      3. 1.2.2: The Diversity of Life
  2. 2: The Chemical Foundation of Life
    1. 2.1: Atoms, Isotopes, Ions, and Molecules
      1. 2.1.0: Overview of Atomic Structure
      2. 2.1.1: Atomic Number and Mass Number
      3. 2.1.2: Isotopes
      4. 2.1.3: The Periodic Table
      5. 2.1.4: Electron Shells and the Bohr Model
      6. 2.1.5: Electron Orbitals
      7. 2.1.6: Chemical Reactions and Molecules
      8. 2.1.7: Ions and Ionic Bonds
      9. 2.1.8: Covalent Bonds and Other Bonds and Interactions
      10. 2.1.9: Hydrogen Bonding and Van der Waals Forces
    2. 2.2: Water
      1. 2.2.0: Water’s Polarity
      2. 2.2.1: Water’s States: Gas, Liquid, and Solid
      3. 2.2.2: Water’s High Heat Capacity
      4. 2.2.3: Water’s Heat of Vaporization
      5. 2.2.4: Water’s Solvent Properties
      6. 2.2.5: Water’s Cohesive and Adhesive Properties
      7. 2.2.6: pH, Buffers, Acids, and Bases
    3. 2.3: Carbon
      1. 2.3.0: The Chemical Basis for Life
      2. 2.3.1: Hydrocarbons
      3. 2.3.2: Organic Isomers
      4. 2.3.3: Organic Enantiomers
      5. 2.3.4: Organic Molecules and Functional Groups
  3. 3: Biological Macromolecules
    1. 3.1: Synthesis of Biological Macromolecules
      1. 3.1.0: Types of Biological Macromolecules
      2. 3.1.1: Dehydration Synthesis
      3. 3.1.2: Hydrolysis
    2. 3.2: Carbohydrates
      1. 3.2.0: Carbohydrate Molecules
      2. 3.2.1: Importance of Carbohydrates
    3. 3.3: Lipids
      1. 3.3.0: Lipid Molecules
      2. 3.3.1: Waxes
      3. 3.3.2: Phospholipids
      4. 3.3.3: Steroids
    4. 3.4: Proteins
      1. 3.4.0: Types and Functions of Proteins
      2. 3.4.1: Amino Acids
      3. 3.4.2: Protein Structure
      4. 3.4.3: Denaturation and Protein Folding
    5. 3.5: Nucleic Acids
      1. 3.5.0: DNA and RNA
      2. 3.5.1: The DNA Double Helix
      3. 3.5.2: DNA Packaging
      4. 3.5.3: Types of RNA
  4. 4: Cell Structure
    1. 4.1: Studying Cells
      1. 4.1.0: Cells as the Basic Unit of Life
      2. 4.1.1: Microscopy
      3. 4.1.2: Cell Theory
      4. 4.1.3: Cell Size
    2. 4.2: Prokaryotic Cells
      1. 4.2.0: Characteristics of Prokaryotic Cells
    3. 4.3: Eukaryotic Cells
      1. 4.3.0: Characteristics of Eukaryotic Cells
      2. 4.3.1: The Plasma Membrane and the Cytoplasm
      3. 4.3.2: The Nucleus and Ribosomes
      4. 4.3.3: Mitochondria
      5. 4.3.4: Comparing Plant and Animal Cells
    4. 4.4: The Endomembrane System and Proteins
      1. 4.4.0: Vesicles and Vacuoles
      2. 4.4.1: The Endoplasmic Reticulum
      3. 4.4.2: The Golgi Apparatus
      4. 4.4.3: Lysosomes
      5. 4.4.4: Peroxisomes
    5. 4.5: The Cytoskeleton
      1. 4.5.0: Microfilaments
      2. 4.5.1: Intermediate Filaments and Microtubules
    6. 4.6: Connections between Cells and Cellular Activities
      1. 4.6.0: Extracellular Matrix of Animal Cells
      2. 4.6.1: Intercellular Junctions
  5. 5: Structure and Function of Plasma Membranes
    1. 5.1: Components and Structure
      1. 5.1.0: Components of Plasma Membranes
      2. 5.1.1: Fluid Mosaic Model
      3. 5.1.2: Membrane Fluidity
    2. 5.2: Passive Transport
      1. 5.2.0: The Role of Passive Transport
      2. 5.2.1: Selective Permeability
      3. 5.2.2: Diffusion
      4. 5.2.3: Facilitated transport
      5. 5.2.4: Osmosis
      6. 5.2.5: Tonicity
      7. 5.2.6: Osmoregulation
    3. 5.3: Active Transport
      1. 5.3.0: Electrochemical Gradient
      2. 5.3.1: Primary Active Transport
      3. 5.3.2: Secondary Active Transport
    4. 5.4: Bulk Transport
      1. 5.4.0: Endocytosis
      2. 5.4.1: Exocytosis
  6. 6: Metabolism
    1. 6.1: Energy and Metabolism
      1. 6.1.0: The Role of Energy and Metabolism
      2. 6.1.1: Types of Energy
      3. 6.1.2: Metabolic Pathways
      4. 6.1.3: Metabolism of Carbohydrates
    2. 6.2: Potential, Kinetic, Free, and Activation Energy
      1. 6.2.0: Free Energy
      2. 6.2.1: The First Law of Thermodynamics
      3. 6.2.2: The Second Law of Thermodynamics
      4. 6.2.3: Activation Energy
    3. 6.3: ATP: Adenosine Triphosphate
      1. 6.3.0: ATP: Adenosine Triphosphate
    4. 6.4: Enzymes
      1. 6.4.0: Enzyme Active Site and Substrate Specificity
      2. 6.4.1: Control of Metabolism Through Enzyme Regulation
  7. 7: Cellular Respiration
    1. 7.1: Energy in Living Systems
      1. 7.1.0: Transforming Chemical Energy
      2. 7.1.1: Electrons and Energy
      3. 7.1.2: ATP in Metabolism
    2. 7.2: Glycolysis
      1. 7.2.0: Importance of Glycolysis
      2. 7.2.1: The Energy-Requiring Steps of Glycolysis
      3. 7.2.2: The Energy-Releasing Steps of Glycolysis
      4. 7.2.3: Outcomes of Glycolysis
    3. 7.3: Oxidation of Pyruvate and the Citric Acid Cycle
      1. 7.3.0: Breakdown of Pyruvate
      2. 7.3.1: Acetyl CoA to CO2
      3. 7.3.2: Citric Acid Cycle
    4. 7.4: Oxidative Phosphorylation
      1. 7.4.0: Electron Transport Chain
      2. 7.4.1: Chemiosmosis and Oxidative Phosphorylation
      3. 7.4.2: ATP Yield
    5. 7.5: Metabolism without Oxygen
      1. 7.5.0: Anaerobic Cellular Respiration
    6. 7.6: Connections of Carbohydrate, Protein, and Lipid Metabolic Pathways
      1. 7.6.0: Connecting Other Sugars to Glucose Metabolism
      2. 7.6.1: Connecting Proteins to Glucose Metabolism
      3. 7.6.2: Connecting Lipids to Glucose Metabolism
    7. 7.7: Regulation of Cellular Respiration
      1. 7.7.0: Regulatory Mechanisms for Cellular Respiration
      2. 7.7.1: Control of Catabolic Pathways
  8. 8: Photosynthesis
    1. 8.1: Overview of Photosynthesis
      1. 8.1.0: The Purpose and Process of Photosynthesis
      2. 8.1.1: Main Structures and Summary of Photosynthesis
      3. 8.1.2: The Two Parts of Photosynthesis
    2. 8.2: The Light-Dependent Reactions of Photosynthesis
      1. 8.2.0: Introduction to Light Energy
      2. 8.2.1: Absorption of Light
      3. 8.2.2: Processes of the Light-Dependent Reactions
    3. 8.3: The Light-Independent Reactions of Photosynthesis
      1. 8.3.0: CAM and C4 Photosynthesis
      2. 8.3.1: The Calvin Cycle
      3. 8.3.2: The Carbon Cycle
  9. 9: Cell Communication
    1. 9.1: Signaling Molecules and Cellular Receptors
      1. 9.1.0: Signaling Molecules and Cellular Receptors
      2. 9.1.1: Forms of Signaling
      3. 9.1.2: Types of Receptors
      4. 9.1.3: Signaling Molecules
    2. 9.2: Propagation of the Cellular Signal
      1. 9.2.0: Binding Initiates a Signaling Pathway
      2. 9.2.1: Methods of Intracellular Signaling
    3. 9.3: Response to the Cellular Signal
      1. 9.3.0: Termination of the Signal Cascade
      2. 9.3.1: Cell Signaling and Gene Expression
      3. 9.3.2: Cell Signaling and Cellular Metabolism
      4. 9.3.3: Cell Signaling and Cell Growth
      5. 9.3.4: Cell Signaling and Cell Death
    4. 9.4: Signaling in Single-Celled Organisms
      1. 9.4.0: Signaling in Yeast
      2. 9.4.1: Signaling in Bacteria
  10. 10: Cell Reproduction
    1. 10.1: Cell Division
      1. 10.1.0: The Role of the Cell Cycle
      2. 10.1.1: Genomic DNA and Chromosomes
      3. 10.1.2: Eukaryotic Chromosomal Structure and Compaction
    2. 10.2: The Cell Cycle
      1. 10.2.0: Interphase
      2. 10.2.1: The Mitotic Phase and the G0 Phase
    3. 10.3: Control of the Cell Cycle
      1. 10.3.0: Regulation of the Cell Cycle by External Events
      2. 10.3.1: Regulation of the Cell Cycle at Internal Checkpoints
      3. 10.3.2: Regulator Molecules of the Cell Cycle
    4. 10.4: Cancer and the Cell Cycle
      1. 10.4.0: Proto-oncogenes
      2. 10.4.1: Tumor Suppressor Genes
    5. 10.5: Prokaryotic Cell Division
      1. 10.5.0: Binary Fission
  11. 11: Meiosis and Sexual Reproduction
    1. 11.1: The Process of Meiosis
      1. 11.1.0: Introduction to Meiosis
      2. 11.1.1: Meiosis I
      3. 11.1.2: Meiosis II
      4. 11.1.3: Comparing Meiosis and Mitosis
    2. 11.2: Sexual Reproduction
      1. 11.2.0: Advantages and Disadvantages of Sexual Reproduction
      2. 11.2.1: Life Cycles of Sexually Reproducing Organisms
  12. 12: Mendel's Experiments and Heredity
    1. 12.1: Mendel’s Experiments and the Laws of Probability
      1. 12.1.0: Introduction to Mendelian Inheritance
      2. 12.1.1: Mendel’s Model System
      3. 12.1.2: Mendelian Crosses
      4. 12.1.3: Garden Pea Characteristics Revealed the Basics of Heredity
      5. 12.1.4: Rules of Probability for Mendelian Inheritance
    2. 12.2: Patterns of Inheritance
      1. 12.2.0: Genes as the Unit of Heredity
      2. 12.2.1: Phenotypes and Genotypes
      3. 12.2.2: The Punnett Square Approach for a Monohybrid Cross
      4. 12.2.3: Alternatives to Dominance and Recessiveness
      5. 12.2.4: Sex-Linked Traits
      6. 12.2.5: Lethal Inheritance Patterns
    3. 12.3: Laws of Inheritance
      1. 12.3.0: Mendel's Laws of Heredity
      2. 12.3.1: Mendel's Law of Dominance
      3. 12.3.2: Mendel's Law of Segregation
      4. 12.3.3: Mendel's Law of Independent Assortment
      5. 12.3.4: Genetic Linkage and Violation of the Law of Independent Assortment
      6. 12.3.5: Epistasis
  13. 13: Modern Understandings of Inheritance
    1. 13.1: Chromosomal Theory and Genetic Linkage
      1. 13.1.0: Chromosomal Theory of Inheritance
      2. 13.1.1: Genetic Linkage and Distances
      3. 13.1.2: Identification of Chromosomes and Karyotypes
    2. 13.2: Chromosomal Basis of Inherited Disorders
      1. 13.2.0: Disorders in Chromosome Number
      2. 13.2.1: Chromosomal Structural Rearrangements
      3. 13.2.2: X-Inactivation
  14. 14: DNA Structure and Function
    1. 14.1: Historical Basis of Modern Understanding
      1. 14.1.0: Discovery of DNA
      2. 14.1.1: Modern Applications of DNA
    2. 14.2: DNA Structure and Sequencing
      1. 14.2.0: The Structure and Sequence of DNA
      2. 14.2.1: DNA Sequencing Techniques
    3. 14.3: DNA Replication
      1. 14.3.0: Basics of DNA Replication
      2. 14.3.1: DNA Replication in Prokaryotes
      3. 14.3.2: DNA Replication in Eukaryotes
      4. 14.3.3: Telomere Replication
    4. 14.4: DNA Repair
      1. 14.4.0: DNA Repair
  15. 15: Genes and Proteins
    1. 15.1: The Genetic Code
      1. 15.1.0: The Relationship Between Genes and Proteins
      2. 15.1.1: The Central Dogma: DNA Encodes RNA and RNA Encodes Protein
    2. 15.2: Prokaryotic Transcription
      1. 15.2.0: Transcription in Prokaryotes
      2. 15.2.1: Initiation of Transcription in Prokaryotes
      3. 15.2.2: Elongation and Termination in Prokaryotes
    3. 15.3: Eukaryotic Transcription
      1. 15.3.0: Initiation of Transcription in Eukaryotes
      2. 15.3.1: Elongation and Termination in Eukaryotes
    4. 15.4: RNA Processing in Eukaryotes
      1. 15.4.0: mRNA Processing
      2. 15.4.1: Processing of tRNAs and rRNAs
    5. 15.5: Ribosomes and Protein Synthesis
      1. 15.5.0: The Protein Synthesis Machinery
      2. 15.5.1: The Mechanism of Protein Synthesis
      3. 15.5.2: Protein Folding, Modification, and Targeting
  16. 16: Gene Expression
    1. 16.1: Regulation of Gene Expression
      1. 16.1.0: The Process and Purpose of Gene Expression Regulation
      2. 16.1.1: Prokaryotic versus Eukaryotic Gene Expression
    2. 16.2: Prokaryotic Gene Regulation
      1. 16.2.0: The trp Operon: A Repressor Operon
      2. 16.2.1: Catabolite Activator Protein (CAP): An Activator Regulator
      3. 16.2.2: The lac Operon: An Inducer Operon
    3. 16.3: Eukaryotic Gene Regulation
      1. 16.3.0: The Promoter and the Transcription Machinery
      2. 16.3.1: Transcriptional Enhancers and Repressors
      3. 16.3.2: Epigenetic Control: Regulating Access to Genes within the Chromosome
      4. 16.3.3: RNA Splicing
      5. 16.3.4: The Initiation Complex and Translation Rate
      6. 16.3.5: Regulating Protein Activity and Longevity
    4. 16.4: Regulating Gene Expression in Cell Development
      1. 16.4.0: Gene Expression in Stem Cells
      2. 16.4.1: Cellular Differentiation
      3. 16.4.2: Mechanics of Cellular Differentation
      4. 16.4.3: Establishing Body Axes during Development
      5. 16.4.4: Gene Expression for Spatial Positioning
      6. 16.4.5: Cell Migration in Multicellular Organisms
      7. 16.4.6: Programmed Cell Death
    5. 16.5: Cancer and Gene Regulation
      1. 16.5.0: Altered Gene Expression in Cancer
      2. 16.5.1: Epigenetic Alterations in Cancer
      3. 16.5.2: Cancer and Transcriptional Control
      4. 16.5.3: Cancer and Post-Transcriptional Control
      5. 16.5.4: Cancer and Translational Control
  17. 17: Biotechnology and Genomics
    1. 17.1: Biotechnology
      1. 17.1.0: Biotechnology
      2. 17.1.1: Basic Techniques to Manipulate Genetic Material (DNA and RNA)
      3. 17.1.2: Molecular and Cellular Cloning
      4. 17.1.3: Reproductive Cloning
      5. 17.1.4: Genetic Engineering
      6. 17.1.5: Genetically Modified Organisms (GMOs)
      7. 17.1.6: Biotechnology in Medicine
      8. 17.1.7: Production of Vaccines, Antibiotics, and Hormones
    2. 17.2: Mapping Genomes
      1. 17.2.0: Genetic Maps
      2. 17.2.1: Physical Maps and Integration with Genetic Maps
    3. 17.3: Whole-Genome Sequencing
      1. 17.3.0: Strategies Used in Sequencing Projects
      2. 17.3.1: Use of Whole-Genome Sequences of Model Organisms
      3. 17.3.2: Uses of Genome Sequences
    4. 17.4: Applying Genomics
      1. 17.4.0: Predicting Disease Risk at the Individual Level
      2. 17.4.1: Pharmacogenomics, Toxicogenomics, and Metagenomics
      3. 17.4.2: Genomics and Biofuels
    5. 17.5: Genomics and Proteomics
      1. 17.5.0: Genomics and Proteomics
      2. 17.5.1: Basic Techniques in Protein Analysis
      3. 17.5.2: Cancer Proteomics
  18. 18: Evolution and the Origin of Species
    1. 18.1: Understanding Evolution
      1. 18.1.0: What is Evolution?
      2. 18.1.1: Charles Darwin and Natural Selection
      3. 18.1.2: The Galapagos Finches and Natural Selection
      4. 18.1.3: Processes and Patterns of Evolution
      5. 18.1.4: Evidence of Evolution
      6. 18.1.5: Misconceptions of Evolution
    2. 18.2: Formation of New Species
      1. 18.2.0: The Biological Species Concept
      2. 18.2.1: Reproductive Isolation
      3. 18.2.2: Speciation
      4. 18.2.3: Allopatric Speciation
      5. 18.2.4: Sympatric Speciation
    3. 18.3: Hybrid Zones and Rates of Speciation
      1. 18.3.0: Hybrid Zones
      2. 18.3.1: Varying Rates of Speciation
    4. 18.4: Evolution of Genomes
      1. 18.4.0: Genomic Similiarities between Distant Species
      2. 18.4.1: Genome Evolution
      3. 18.4.2: Whole-Genome Duplication
      4. 18.4.3: Gene Duplications and Divergence
      5. 18.4.4: Noncoding DNA
      6. 18.4.5: Variations in Size and Number of Genes
    5. 18.5: Evidence of Evolution
      1. 18.5.0: The Fossil Record as Evidence for Evolution
      2. 18.5.1: Fossil Formation
      3. 18.5.2: Gaps in the Fossil Record
      4. 18.5.3: Carbon Dating and Estimating Fossil Age
      5. 18.5.4: The Fossil Record and the Evolution of the Modern Horse
      6. 18.5.5: Homologous Structures
      7. 18.5.6: Convergent Evolution
      8. 18.5.7: Vestigial Structures
      9. 18.5.8: Biogeography and the Distribution of Species
  19. 19: The Evolution of Populations
    1. 19.1: Population Evolution
      1. 19.1.0: Defining Population Evolution
      2. 19.1.1: Population Genetics
      3. 19.1.2: Hardy-Weinberg Principle of Equilibrium
    2. 19.2: Population Genetics
      1. 19.2.0: Genetic Variation
      2. 19.2.1: Genetic Drift
      3. 19.2.2: Gene Flow and Mutation
      4. 19.2.3: Nonrandom Mating and Environmental Variance
    3. 19.3: Adaptive Evolution
      1. 19.3.0: Natural Selection and Adaptive Evolution
      2. 19.3.1: Stabilizing, Directional, and Diversifying Selection
      3. 19.3.2: Frequency-Dependent Selection
      4. 19.3.3: Sexual Selection
      5. 19.3.4: No Perfect Organism
  20. 20: Phylogenies and the History of Life
    1. 20.1: Organizing Life on Earth
      1. 20.1.0: Phylogenetic Trees
      2. 20.1.1: Limitations of Phylogenetic Trees
      3. 20.1.2: The Levels of Classification
    2. 20.2: Determining Evolutionary Relationships
      1. 20.2.0: Distinguishing between Similar Traits
      2. 20.2.1: Building Phylogenetic Trees
    3. 20.3: Perspectives on the Phylogenetic Tree
      1. 20.3.0: Limitations to the Classic Model of Phylogenetic Trees
      2. 20.3.1: Horizontal Gene Transfer
      3. 20.3.2: Endosymbiotic Theory and the Evolution of Eukaryotes
      4. 20.3.3: Web, Network, and Ring of Life Models
  21. 21: Viruses
    1. 21.1: Viral Evolution, Morphology, and Classification
      1. 21.1.0: Discovery and Detection of Viruses
      2. 21.1.1: Evolution of Viruses
      3. 21.1.2: Viral Morphology
      4. 21.1.3: Virus Classification
    2. 21.2: Virus Infections and Hosts
      1. 21.2.0: Steps of Virus Infections
      2. 21.2.1: The Lytic and Lysogenic Cycles of Bacteriophages
      3. 21.2.2: Animal Viruses
      4. 21.2.3: Plant Viruses
    3. 21.3: Prevention and Treatment of Viral Infections
      1. 21.3.0: Vaccines and Immunity
      2. 21.3.1: Vaccines and Anti-Viral Drugs for Treatment
    4. 21.4: Prions and Viroids
      1. 21.4.0: Prions and Viroids
  22. 22: Prokaryotes: Bacteria and Archaea
    1. 22.1: Prokaryotic Diversity
      1. 22.1.0: Classification of Prokaryotes
      2. 22.1.1: The Origins of Archaea and Bacteria
      3. 22.1.2: Extremophiles and Biofilms
    2. 22.2: Structure of Prokaryotes
      1. 22.2.0: Basic Structures of Prokaryotic Cells
      2. 22.2.1: Prokaryotic Reproduction
    3. 22.3: Prokaryotic Metabolism
      1. 22.3.0: Energy and Nutrient Requirements for Prokaryotes
      2. 22.3.1: The Role of Prokaryotes in Ecosystems
    4. 22.4: Bacterial Diseases in Humans
      1. 22.4.0: History of Bacterial Diseases
      2. 22.4.1: Biofilms and Disease
      3. 22.4.2: Antibiotics: Are We Facing a Crisis?
      4. 22.4.3: Bacterial Foodborne Diseases
    5. 22.5: Beneficial Prokaryotes
      1. 22.5.0: Symbiosis between Bacteria and Eukaryotes
      2. 22.5.1: Early Biotechnology: Cheese, Bread, Wine, Beer, and Yogurt
      3. 22.5.2: Prokaryotes and Environmental Bioremediation
  23. 23: Protists
    1. 23.1: Eukaryotic Origins
      1. 23.1.0: Early Eukaryotes
      2. 23.1.1: Characteristics of Eukaryotic DNA
      3. 23.1.2: Endosymbiosis and the Evolution of Eukaryotes
      4. 23.1.3: The Evolution of Mitochondria
      5. 23.1.4: The Evolution of Plastids
    2. 23.2: Characteristics of Protists
      1. 23.2.0: Cell Structure, Metabolism, and Motility
      2. 23.2.1: Protist Life Cycles and Habitats
    3. 23.3: Groups of Protists
      1. 23.3.0: Excavata
      2. 23.3.1: Chromalveolata: Alveolates
      3. 23.3.2: Chromalveolata: Stramenopiles
      4. 23.3.3: Rhizaria
      5. 23.3.4: Archaeplastida
      6. 23.3.5: Amoebozoa and Opisthokonta
    4. 23.4: Ecology of Protists
      1. 23.4.0: Protists as Primary Producers, Food Sources, and Symbionts
      2. 23.4.1: Protists as Human Pathogens
      3. 23.4.2: Protists as Plant Pathogens
  24. 24: Fungi
    1. 24.1: Characteristics of Fungi
      1. 24.1.0: Characteristics of Fungi
      2. 24.1.1: Fungi Cell Structure and Function
      3. 24.1.2: Fungi Reproduction
    2. 24.2: Ecology of Fungi
      1. 24.2.0: Fungi Habitat, Decomposition, and Recycling
      2. 24.2.1: Mutualistic Relationships with Fungi and Fungivores
    3. 24.3: Classifications of Fungi
      1. 24.3.0: Chytridiomycota: The Chytrids
      2. 24.3.1: Zygomycota: The Conjugated Fungi
      3. 24.3.2: Ascomycota: The Sac Fungi
      4. 24.3.3: Basidiomycota: The Club Fungi
      5. 24.3.4: Deuteromycota: The Imperfect Fungi
      6. 24.3.5: Glomeromycota
    4. 24.4: Fungal Parasites and Pathogens
      1. 24.4.0: Fungi as Plant, Animal, and Human Pathogens
    5. 24.5: Importance of Fungi in Human Life
      1. 24.5.0: Importance of Fungi in Human Life
  25. 25: Seedless Plants
    1. 25.1: Early Plant Life
      1. 25.1.0: Early Plant Life
      2. 25.1.1: Evolution of Land Plants
      3. 25.1.2: Plant Adaptations to Life on Land
      4. 25.1.3: Sporophytes and Gametophytes in Seedless Plants
      5. 25.1.4: Structural Adaptations for Land in Seedless Plants
      6. 25.1.5: The Major Divisions of Land Plants
    2. 25.2: Green Algae: Precursors of Land Plants
      1. 25.2.0: Streptophytes and Reproduction of Green Algae
      2. 25.2.1: Charales
    3. 25.3: Bryophytes
      1. 25.3.0: Bryophytes
      2. 25.3.1: Liverworts and Hornworts
      3. 25.3.2: Mosses
    4. 25.4: Seedless Vascular Plants
      1. 25.4.0: Seedless Vascular Plants
      2. 25.4.1: Vascular Tissue: Xylem and Phloem
      3. 25.4.2: The Evolution of Roots in Seedless Plants
      4. 25.4.3: Ferns and Other Seedless Vascular Plants
      5. 25.4.4: The Importance of Seedless Vascular Plants
  26. 26: Seed Plants
    1. 26.1: Evolution of Seed Plants
      1. 26.1.0: The Evolution of Seed Plants and Adaptations for Land
      2. 26.1.1: Evolution of Gymnosperms
      3. 26.1.2: Evolution of Angiosperms
    2. 26.2: Gymnosperms
      1. 26.2.0: Characteristics of Gymnosperms
      2. 26.2.1: Life Cycle of a Conifer
      3. 26.2.2: Diversity of Gymnosperms
    3. 26.3: Angiosperms
      1. 26.3.0: Angiosperm Flowers
      2. 26.3.1: Angsiosperm Fruit
      3. 26.3.2: The Life Cycle of an Angiosperm
      4. 26.3.3: Diversity of Angiosperms
    4. 26.4: The Role of Seed Plants
      1. 26.4.0: Herbivory and Pollination
      2. 26.4.1: The Importance of Seed Plants in Human Life
      3. 26.4.2: Biodiversity of Plants
  27. 27: Introduction to Animal Diversity
    1. 27.1: Features of the Animal Kingdom
      1. 27.1.0: Characteristics of the Animal Kingdom
      2. 27.1.1: Complex Tissue Structure
      3. 27.1.2: Animal Reproduction and Development
    2. 27.2: Features Used to Classify Animals
      1. 27.2.0: Animal Characterization Based on Body Symmetry
      2. 27.2.1: Animal Characterization Based on Features of Embryological Development
    3. 27.3: Animal Phylogeny
      1. 27.3.0: Constructing an Animal Phylogenetic Tree
      2. 27.3.1: Molecular Analyses and Modern Phylogenetic Trees
    4. 27.4: The Evolutionary History of the Animal Kingdom
      1. 27.4.0: Pre-Cambrian Animal Life
      2. 27.4.1: The Cambrian Explosion of Animal Life
      3. 27.4.2: Post-Cambrian Evolution and Mass Extinctions
  28. 28: Invertebrates
    1. 28.1: Phylum Porifera
      1. 28.1.0: Phylum Porifera
      2. 28.1.1: Morphology of Sponges
      3. 28.1.2: Physiological Processes in Sponges
    2. 28.2: Phylum Cnidaria
      1. 28.2.0: Phylum Cnidaria
      2. 28.2.1: Class Anthozoa
      3. 28.2.2: Class Scyphozoa
      4. 28.2.3: Class Cubozoa and Class Hydrozoa
    3. 28.3: Superphylum Lophotrochozoa
      1. 28.3.0: Superphylum Lophotrochozoa
      2. 28.3.1: Phylum Platyhelminthes
      3. 28.3.2: Phylum Rotifera
      4. 28.3.3: Phylum Nemertea
      5. 28.3.4: Phylum Mollusca
      6. 28.3.5: Classification of Phylum Mollusca
      7. 28.3.6: Phylum Annelida
    4. 28.4: Superphylum Ecdysozoa
      1. 28.4.0: Superphylum Ecdysozoa
      2. 28.4.1: Phylum Nematoda
      3. 28.4.2: Phylum Arthropoda
      4. 28.4.3: Subphyla of Arthropoda
    5. 28.5: Superphylum Deuterostomia
      1. 28.5.0: Phylum Echinodermata
      2. 28.5.1: Classes of Echinoderms
      3. 28.5.2: Phylum Chordata
  29. 29: Vertebrates
    1. 29.1: Chordates
      1. 29.1.0: Characteristics of Chordata
      2. 29.1.1: Chordates and the Evolution of Vertebrates
      3. 29.1.2: The Evolution of Craniata and Vertebrata
      4. 29.1.3: Characteristics of Vertebrates
    2. 29.2: Fishes
      1. 29.2.0: Agnathans: Jawless Fishes
      2. 29.2.1: Gnathostomes: Jawed Fishes
    3. 29.3: Amphibians
      1. 29.3.0: Characteristics and Evolution of Amphibians
      2. 29.3.1: Modern Amphibians
    4. 29.4: Reptiles
      1. 29.4.0: Characteristics of Amniotes
      2. 29.4.1: Evolution of Amniotes
      3. 29.4.2: Characteristics of Reptiles
      4. 29.4.3: Evolution of Reptiles
      5. 29.4.4: Modern Reptiles
    5. 29.5: Birds
      1. 29.5.0: Characteristics of Birds
      2. 29.5.1: Evolution of Birds
    6. 29.6: Mammals
      1. 29.6.0: Characteristics of Mammals
      2. 29.6.1: Evolution of Mammals
      3. 29.6.2: Living Mammals
    7. 29.7: The Evolution of Primates
      1. 29.7.0: Characteristics and Evolution of Primates
      2. 29.7.1: Early Human Evolution
      3. 29.7.2: Early Hominins
      4. 29.7.3: Genus Homo
  30. 30: Plant Form and Physiology
    1. 30.1: The Plant Body
      1. 30.1.0: Plant Tissues and Organ Systems
    2. 30.2: Stems
      1. 30.2.0: Functions of Stems
      2. 30.2.1: Stem Anatomy
      3. 30.2.2: Primary and Secondary Growth in Stems
      4. 30.2.3: Stem Modifications
    3. 30.3: Roots
      1. 30.3.0: Types of Root Systems and Zones of Growth
      2. 30.3.1: Root Modifications
    4. 30.4: Leaves
      1. 30.4.0: Leaf Structure and Arrangment
      2. 30.4.1: Types of Leaf Forms
      3. 30.4.2: Leaf Structure, Function, and Adaptation
    5. 30.5: Plant Development
      1. 30.5.0: Meristems
      2. 30.5.1: Genetic Control of Flowers
    6. 30.6: Transport of Water and Solutes in Plants
      1. 30.6.0: Water and Solute Potential
      2. 30.6.1: Pressure, Gravity, and Matric Potential
      3. 30.6.2: Movement of Water and Minerals in the Xylem
      4. 30.6.3: Transportation of Photosynthates in the Phloem
    7. 30.7: Plant Sensory Systems and Responses
      1. 30.7.0: Plant Responses to Light
      2. 30.7.1: The Phytochrome System and Red Light Response
      3. 30.7.2: Blue Light Response
      4. 30.7.3: Plant Responses to Gravity
      5. 30.7.4: Auxins, Cytokinins, and Gibberellins
      6. 30.7.5: Abscisic Acid, Ethylene, and Nontraditional Hormones
      7. 30.7.6: Plant Responses to Wind and Touch
    8. 30.8: Plant Defense Mechanisms
      1. 30.8.0: Plant Defenses Against Herbivores
      2. 30.8.1: Plant Defenses Against Pathogens
  31. 31: Soil and Plant Nutrition
    1. 31.1: Nutritional Requirements of Plants
      1. 31.1.0: Plant Nutrition
      2. 31.1.1: The Chemical Composition of Plants
      3. 31.1.2: Essential Nutrients for Plants
    2. 31.2: The Soil
      1. 31.2.0: Soil Composition
      2. 31.2.1: Soil Formation
      3. 31.2.2: Physical Properties of Soil
    3. 31.3: Nutritional Adaptations of Plants
      1. 31.3.0: Nitrogen Fixation: Root and Bacteria Interactions
      2. 31.3.1: Mycorrhizae: The Symbiotic Relationship between Fungi and Roots
      3. 31.3.2: Nutrients from Other Sources
  32. 32: Plant Reproduction
    1. 32.1: Plant Reproductive Development and Structure
      1. 32.1.0: Plant Reproductive Development and Structure
      2. 32.1.1: Sexual Reproduction in Gymnosperms
      3. 32.1.2: Sexual Reproduction in Angiosperms
    2. 32.2: Pollination and Fertilization
      1. 32.2.0: Pollination and Fertilization
      2. 32.2.1: Pollination by Insects
      3. 32.2.2: Pollination by Bats, Birds, Wind, and Water
      4. 32.2.3: Double Fertilization in Plants
      5. 32.2.4: Development of the Seed
      6. 32.2.5: Development of Fruit and Fruit Types
      7. 32.2.6: Fruit and Seed Dispersal
    3. 32.3: Asexual Reproduction
      1. 32.3.0: Asexual Reproduction in Plants
      2. 32.3.1: Natural and Artificial Methods of Asexual Reproduction in Plants
      3. 32.3.2: Plant Life Spans
  33. 33: The Animal Body: Basic Form and Function
    1. 33.1: Animal Form and Function
      1. 33.1.0: Characteristics of the Animal Body
      2. 33.1.1: Body Plans
      3. 33.1.2: Limits on Animal Size and Shape
      4. 33.1.3: Limiting Effects of Diffusion on Size and Development
      5. 33.1.4: Animal Bioenergetics
      6. 33.1.5: Animal Body Planes and Cavities
    2. 33.2: Animal Primary Tissues
      1. 33.2.0: Epithelial Tissues
      2. 33.2.1: Connective Tissues: Loose, Fibrous, and Cartilage
      3. 33.2.2: Connective Tissues: Bone, Adipose, and Blood
      4. 33.2.3: Muscle Tissues and Nervous Tissues
    3. 33.3: Homeostasis
      1. 33.3.0: Homeostatic Process
      2. 33.3.1: Control of Homeostasis
      3. 33.3.2: Homeostasis: Thermoregulation
      4. 33.3.3: Heat Conservation and Dissipation
  34. 34: Animal Nutrition and the Digestive System
    1. 34.1: Digestive Systems
      1. 34.1.0: Digestive Systems
      2. 34.1.1: Herbivores, Omnivores, and Carnivores
      3. 34.1.2: Invertebrate Digestive Systems
      4. 34.1.3: Vertebrate Digestive Systems
      5. 34.1.4: Digestive System: Mouth and Stomach
      6. 34.1.5: Digestive System: Small and Large Intestines
    2. 34.2: Nutrition and Energy Production
      1. 34.2.0: Food Requirements and Essential Nutrients
      2. 34.2.1: Food Energy and ATP
    3. 34.3: Digestive System Processes
      1. 34.3.0: Ingestion
      2. 34.3.1: Digestion and Absorption
      3. 34.3.2: Elimination
    4. 34.4: Digestive System Regulation
      1. 34.4.0: Neural Responses to Food
      2. 34.4.1: Hormonal Responses to Food
  35. 35: The Nervous System
    1. 35.1: Neurons and Glial Cells
      1. 35.1.0: Neurons and Glial Cells
      2. 35.1.1: Neurons
      3. 35.1.2: Glia
    2. 35.2: How Neurons Communicate
      1. 35.2.0: Nerve Impulse Transmission within a Neuron: Resting Potential
      2. 35.2.1: Nerve Impulse Transmission within a Neuron: Action Potential
      3. 35.2.2: Synaptic Transmission
      4. 35.2.3: Signal Summation
      5. 35.2.4: Synaptic Plasticity
    3. 35.3: The Nervous System
      1. 35.3.0: The Nervous System
    4. 35.4: The Central Nervous System
      1. 35.4.0: Brain: Cerebral Cortex and Brain Lobes
      2. 35.4.1: Brain: Midbrain and Brain Stem
      3. 35.4.2: Spinal Cord
    5. 35.5: The Peripheral Nervous System
      1. 35.5.0: Autonomic Nervous System
      2. 35.5.1: Sensory-Somatic Nervous System
    6. 35.6: Nervous System Disorders
      1. 35.6.0: Neurodegenerative Disorders
      2. 35.6.1: Neurodevelopmental Disorders: Autism and ADHD
      3. 35.6.2: Neurodevelopmental Disorders: Mental Illnesses
      4. 35.6.3: Other Neurological Disorders
  36. 36: Sensory Systems
    1. 36.1: Sensory Processes
      1. 36.1.0: Reception
      2. 36.1.1: Transduction and Perception
    2. 36.2: Somatosensation
      1. 36.2.0: Somatosensory Receptors
      2. 36.2.1: Integration of Signals from Mechanoreceptors
      3. 36.2.2: Thermoreception
    3. 36.3: Taste and Smell
      1. 36.3.0: Tastes and Odors
      2. 36.3.1: Reception and Transduction
    4. 36.4: Hearing and Vestibular Sensation
      1. 36.4.0: Sound
      2. 36.4.1: Reception of Sound
      3. 36.4.2: Transduction of Sound
      4. 36.4.3: The Vestibular System
      5. 36.4.4: Balance and Determining Equilibrium
    5. 36.5: Vision
      1. 36.5.0: Light
      2. 36.5.1: Anatomy of the Eye
      3. 36.5.2: Transduction of Light
      4. 36.5.3: Visual Processing
  37. 37: The Endocrine System
    1. 37.1: Types of Hormones
      1. 37.1.0: Hormone Functions
      2. 37.1.1: Lipid-Derived, Amino Acid-Derived, and Peptide Hormones
    2. 37.2: How Hormones Work
      1. 37.2.0: How Hormones Work
      2. 37.2.1: Intracellular Hormone Receptors
      3. 37.2.2: Plasma Membrane Hormone Receptors
    3. 37.3: Regulation of Body Processes
      1. 37.3.0: Hormonal Regulation of the Excretory System
      2. 37.3.1: Hormonal Regulation of the Reproductive System
      3. 37.3.2: Hormonal Regulation of Metabolism
      4. 37.3.3: Hormonal Control of Blood Calcium Levels
      5. 37.3.4: Hormonal Regulation of Growth
      6. 37.3.5: Hormonal Regulation of Stress
    4. 37.4: Regulation of Hormone Production
      1. 37.4.0: Humoral, Hormonal, and Neural Stimuli
    5. 37.5: Endocrine Glands
      1. 37.5.0: Hypothalamic-Pituitary Axis
      2. 37.5.1: Thyroid Gland
      3. 37.5.2: Parathyroid Glands
      4. 37.5.3: Adrenal Glands
      5. 37.5.4: Pancreas
      6. 37.5.5: Pineal Gland and Gonads
      7. 37.5.6: Organs with Secondary Endocrine Functions
  38. 38: The Musculoskeletal System
    1. 38.1: Types of Skeletal Systems
      1. 38.1.0: Functions of the Musculoskeletal System
      2. 38.1.1: Types of Skeletal Systems
      3. 38.1.2: Human Axial Skeleton
      4. 38.1.3: Human Appendicular Skeleton
    2. 38.2: Bone
      1. 38.2.0: Bone
      2. 38.2.1: Cell Types in Bones
      3. 38.2.2: Bone Development
      4. 38.2.3: Growth of Bone
      5. 38.2.4: Bone Remodeling and Repair
    3. 38.3: Joints and Skeletal Movement
      1. 38.3.0: Classification of Joints on the Basis of Structure and Function
      2. 38.3.1: Movement at Synovial Joints
      3. 38.3.2: Types of Synovial Joints
      4. 38.3.3: Bone and Joint Disorders
    4. 38.4: Muscle Contraction and Locomotion
      1. 38.4.0: Structure and Function of the Muscular System
      2. 38.4.1: Skeletal Muscle Fibers
      3. 38.4.2: Sliding Filament Model of Contraction
      4. 38.4.3: ATP and Muscle Contraction
      5. 38.4.4: Regulatory Proteins
      6. 38.4.5: Excitation–Contraction Coupling
      7. 38.4.6: Control of Muscle Tension
  39. 39: The Respiratory System
    1. 39.1: Systems of Gas Exchange
      1. 39.1.0: The Respiratory System and Direct Diffusion
      2. 39.1.1: Skin, Gills, and Tracheal Systems
      3. 39.1.2: Amphibian and Bird Respiratory Systems
      4. 39.1.3: Mammalian Systems and Protective Mechanisms
    2. 39.2: Gas Exchange across Respiratory Surfaces
      1. 39.2.0: Gas Pressure and Respiration
      2. 39.2.1: Basic Principles of Gas Exchange
      3. 39.2.2: Lung Volumes and Capacities
      4. 39.2.3: Gas Exchange across the Alveoli
    3. 39.3: Breathing
      1. 39.3.0: The Mechanics of Human Breathing
      2. 39.3.1: Types of Breathing
      3. 39.3.2: The Work of Breathing
      4. 39.3.3: Dead Space: V/Q Mismatch
    4. 39.4: Transport of Gases in Human Bodily Fluids
      1. 39.4.0: Transport of Oxygen in the Blood
      2. 39.4.1: Transport of Carbon Dioxide in the Blood
  40. 40: The Circulatory System
    1. 40.1: Overview of the Circulatory System
      1. 40.1.0: The Role of the Circulatory System
      2. 40.1.1: Open and Closed Circulatory Systems
      3. 40.1.2: Types of Circulatory Systems in Animals
    2. 40.2: Components of the Blood
      1. 40.2.0: The Role of Blood in the Body
      2. 40.2.1: Red Blood Cells
      3. 40.2.2: White Blood Cells
      4. 40.2.3: Platelets and Coagulation Factors
      5. 40.2.4: Plasma and Serum
    3. 40.3: Mammalian Heart and Blood Vessels
      1. 40.3.0: Structures of the Heart
      2. 40.3.1: Arteries, Veins, and Capillaries
      3. 40.3.2: The Cardiac Cycle
    4. 40.4: Blood Flow and Blood Pressure Regulation
      1. 40.4.0: Blood Flow Through the Body
      2. 40.4.1: Blood Pressure
  41. 41: Osmotic Regulation and the Excretory System
    1. 41.1: Osmoregulation and Osmotic Balance
      1. 41.1.0: Introduction to Osmoregulation
      2. 41.1.1: Transport of Electrolytes across Cell Membranes
      3. 41.1.2: Concept of Osmolality and Milliequivalent
      4. 41.1.3: Osmoregulators and Osmoconformers
    2. 41.2: Nitrogenous Wastes
      1. 41.2.0: Nitrogenous Waste in Terrestrial Animals: The Urea Cycle
      2. 41.2.1: Nitrogenous Waste in Birds and Reptiles: Uric Acid
    3. 41.3: Excretion Systems
      1. 41.3.0: Contractile Vacuoles in Microorganisms
      2. 41.3.1: Flame Cells of Planaria and Nephridia of Worms
      3. 41.3.2: Malpighian Tubules of Insects
    4. 41.4: Human Osmoregulatory and Excretory Systems
      1. 41.4.0: Kidney Structure
      2. 41.4.1: Nephron: The Functional Unit of the Kidney
      3. 41.4.2: Kidney Function and Physiology
    5. 41.5: Hormonal Control of Osmoregulatory Functions
      1. 41.5.0: Epinephrine and Norepinephrine
      2. 41.5.1: Other Hormonal Controls for Osmoregulation
  42. 42: The Immune System
    1. 42.1: Innate Immune Response
      1. 42.1.0: Innate Immune Response
      2. 42.1.1: Physical and Chemical Barriers
      3. 42.1.2: Pathogen Recognition
      4. 42.1.3: Natural Killer Cells
      5. 42.1.4: The Complement System
    2. 42.2: Adaptive Immune Response
      1. 42.2.0: Antigen-presenting Cells: B and T cells
      2. 42.2.1: Humoral Immune Response
      3. 42.2.2: Cell-Mediated Immunity
      4. 42.2.3: Cytotoxic T Lymphocytes and Mucosal Surfaces
      5. 42.2.4: Immunological Memory
      6. 42.2.5: Regulating Immune Tolerance
    3. 42.3: Antibodies
      1. 42.3.0: Antibody Structure
      2. 42.3.1: Antibody Functions
    4. 42.4: Disruptions in the Immune System
      1. 42.4.0: Immunodeficiency
      2. 42.4.1: Hypersensitivities
  43. 43: Animal Reproduction and Development
    1. 43.1: Reproduction Methods
      1. 43.1.0: Methods of Reproducing
      2. 43.1.1: Types of Sexual and Asexual Reproduction
      3. 43.1.2: Sex Determination
    2. 43.2: Fertilization
      1. 43.2.0: External and Internal Fertilization
      2. 43.2.1: The Evolution of Reproduction
    3. 43.3: Human Reproductive Anatomy and Gametogenesis
      1. 43.3.0: Male Reproductive Anatomy
      2. 43.3.1: Female Reproductive Anatomy
      3. 43.3.2: Gametogenesis (Spermatogenesis and Oogenesis)
    4. 43.4: Hormonal Control of Human Reproduction
      1. 43.4.0: Male Hormones
      2. 43.4.1: Female Hormones
    5. 43.5: Fertilization and Early Embryonic Development
      1. 43.5.0: Fertilization
      2. 43.5.1: Cleavage, the Blastula Stage, and Gastrulation
    6. 43.6: Organogenesis and Vertebrate Formation
      1. 43.6.0: Organogenesis
      2. 43.6.1: Vertebrate Axis Formation
    7. 43.7: Human Pregnancy and Birth
      1. 43.7.0: Human Gestation
      2. 43.7.1: Labor and Birth
      3. 43.7.2: Contraception and Birth Control
      4. 43.7.3: Infertility
  44. 44: Ecology and the Biosphere
    1. 44.1: The Scope of Ecology
      1. 44.1.0: Introduction to Ecology
      2. 44.1.1: Organismal Ecology and Population Ecology
      3. 44.1.2: Community Ecology and Ecosystem Ecology
    2. 44.2: Biogeography
      1. 44.2.0: Biogeography
      2. 44.2.1: Energy Sources
      3. 44.2.2: Temperature and Water
      4. 44.2.3: Inorganic Nutrients and Other Factors
      5. 44.2.4: Abiotic Factors Influencing Plant Growth
    3. 44.3: Terrestrial Biomes
      1. 44.3.0: What constitutes a biome?
      2. 44.3.1: Tropical Wet Forest and Savannas
      3. 44.3.2: Subtropical Deserts and Chaparral
      4. 44.3.3: Temperate Grasslands
      5. 44.3.4: Temperate Forests
      6. 44.3.5: Boreal Forests and Arctic Tundra
    4. 44.4: Aquatic Biomes
      1. 44.4.0: Abiotic Factors Influencing Aquatic Biomes
      2. 44.4.1: Marine Biomes
      3. 44.4.2: Estuaries: Where the Ocean Meets Fresh Water
      4. 44.4.3: Freshwater Biomes
    5. 44.5: Climate and the Effects of Global Climate Change
      1. 44.5.0: Climate and Weather
      2. 44.5.1: Causes of Global Climate Change
      3. 44.5.2: Evidence of Global Climate Change
      4. 44.5.3: Past and Present Effects of Climate Change
  45. 45: Population and Community Ecology
    1. 45.1: Population Demography
      1. 45.1.0: Population Demography
      2. 45.1.1: Population Size and Density
      3. 45.1.2: Species Distribution
      4. 45.1.3: The Study of Population Dynamics
    2. 45.2: Environmental Limits to Population Growth
      1. 45.2.0: Exponential Population Growth
      2. 45.2.1: Logistic Population Growth
      3. 45.2.2: Density-Dependent and Density-Independent Population Regulation
    3. 45.3: Life History Patterns
      1. 45.3.0: Life History Patterns and Energy Budgets
      2. 45.3.1: Theories of Life History
    4. 45.4: Human Population Growth
      1. 45.4.0: Human Population Growth
      2. 45.4.1: Overcoming Density-Dependent Regulation
      3. 45.4.2: Age Structure, Population Growth, and Economic Development
    5. 45.5: Community Ecology
      1. 45.5.0: The Role of Species within Communities
      2. 45.5.1: Predation, Herbivory, and the Competitive Exclusion Principle
      3. 45.5.2: Symbiosis
      4. 45.5.3: Ecological Succession
    6. 45.6: Innate Animal Behavior
      1. 45.6.0: Introduction to Animal Behavior
      2. 45.6.1: Movement and Migration
      3. 45.6.2: Animal Communication and Living in Groups
      4. 45.6.3: Altruism and Populations
      5. 45.6.4: Mating Systems and Sexual Selection
    7. 45.7: Learned Animal Behavior
      1. 45.7.0: Simple Learned Behaviors
      2. 45.7.1: Conditioned Behavior
      3. 45.7.2: Cognitive Learning and Sociobiology
  46. 46: Ecosystems
    1. 46.1: Ecology of Ecosystems
      1. 46.1.0: Ecosystem Dynamics
      2. 46.1.1: Food Chains and Food Webs
      3. 46.1.2: Studying Ecosystem Dynamics
      4. 46.1.3: Modeling Ecosystem Dynamics
    2. 46.2: Energy Flow through Ecosystems
      1. 46.2.0: Strategies for Acquiring Energy
      2. 46.2.1: Productivity within Trophic Levels
      3. 46.2.2: Transfer of Energy between Trophic Levels
      4. 46.2.3: Ecological Pyramids
      5. 46.2.4: Biological Magnification
    3. 46.3: Biogeochemical Cycles
      1. 46.3.0: Biogeochemical Cycles
      2. 46.3.1: The Water (Hydrologic) Cycle
      3. 46.3.2: The Carbon Cycle
      4. 46.3.3: The Nitrogen Cycle
      5. 46.3.4: The Phosphorus Cycle
      6. 46.3.5: The Sulfur Cycle
  47. 47: Conservation Biology and Biodiversity
    1. 47.1: The Biodiversity Crisis
      1. 47.1.0: Loss of Biodiversity
      2. 47.1.1: Types of Biodiversity
      3. 47.1.2: Biodiversity Change through Geological Time
      4. 47.1.3: The Pleistocene Extinction
      5. 47.1.4: Present-Time Extinctions
    2. 47.2: The Importance of Biodiversity to Human Life
      1. 47.2.0: Human Health and Biodiversity
      2. 47.2.1: Agricultural Diversity
      3. 47.2.2: Managing Fisheries
    3. 47.3: Threats to Biodiversity
      1. 47.3.0: Habitat Loss and Sustainability
      2. 47.3.1: Overharvesting
      3. 47.3.2: Exotic Species
      4. 47.3.3: Climate Change and Biodiversity
    4. 47.4: Preserving Biodiversity
      1. 47.4.0: Measuring Biodiversity
      2. 47.4.1: Changing Human Behavior in Response to Biodiversity Loss
      3. 47.4.2: Ecological Restoration

15.4: RNA Processing in Eukaryotes

15.4.1: mRNA Processing

Eukaryotic pre-mRNA receives a 5' cap and a 3' poly (A) tail before introns are removed and the mRNA is considered ready for translation.

Learning Objective

Outline the steps of pre-mRNA processing

Key Points

  • A 7-methylguanosine cap is added to the 5' end of the pre-mRNA while elongation is still in progress. The 5' cap protects the nascent mRNA from degradation and assists in ribosome binding during translation.
  • A poly (A) tail is added to the 3' end of the pre-mRNA once elongation is complete. The poly (A) tail protects the mRNA from degradation, aids in the export of the mature mRNA to the cytoplasm, and is involved in binding proteins involved in initiating translation.
  • Introns are removed from the pre-mRNA before the mRNA is exported to the cytoplasm.

Key Terms

intron

a portion of a split gene that is included in pre-RNA transcripts but is removed during RNA processing and rapidly degraded

moiety

a specific segment of a molecule

spliceosome

a dynamic complex of RNA and protein subunits that removes introns from precursor mRNA

Pre-mRNA Processing

The eukaryotic pre-mRNA undergoes extensive processing before it is ready to be translated. The additional steps involved in eukaryotic mRNA maturation create a molecule with a much longer half-life than a prokaryotic mRNA. Eukaryotic mRNAs last for several hours, whereas the typical E. coli mRNA lasts no more than five seconds.

Pre-mRNAs are first coated in RNA-stabilizing proteins; these protect the pre-mRNA from degradation while it is processed and exported out of the nucleus. The three most important steps of pre-mRNA processing are the addition of stabilizing and signaling factors at the 5' and 3' ends of the molecule, and the removal of intervening sequences that do not specify the appropriate amino acids. In rare cases, the mRNA transcript can be "edited" after it is transcribed.

5' Capping

While the pre-mRNA is still being synthesized, a 7-methylguanosine cap is added to the 5' end of the growing transcript by a 5'-to-5' phosphate linkage. This moiety protects the nascent mRNA from degradation. In addition, initiation factors involved in protein synthesis recognize the cap to help initiate translation by ribosomes.

5' cap structure

5' cap structure

Capping of the pre-mRNA involves the addition of 7-methylguanosine (m7G) to the 5' end. The cap protects the 5' end of the primary RNA transcript from attack by ribonucleases and is recognized by eukaryotic initiation factors involved in assembling the ribosome on the mature mRNA prior to initiating translation.

3' Poly-A Tail

While RNA Polymerase II is still transcribing downstream of the proper end of a gene, the pre-mRNA is cleaved by an endonuclease-containing protein complex between an AAUAAA consensus sequence and a GU-rich sequence. This releases the functional pre-mRNA from the rest of the transcript, which is still attached to the RNA Polymerase. An enzyme called poly (A) polymerase (PAP) is part of the same protein complex that cleaves the pre-mRNA and it immediately adds a string of approximately 200 A nucleotides, called the poly (A) tail, to the 3' end of the just-cleaved pre-mRNA. The poly (A) tail protects the mRNA from degradation, aids in the export of the mature mRNA to the cytoplasm, and is involved in binding proteins involved in initiating translation.

Poly (A) Polymerase adds a 3' poly (A) tail to the pre-mRNA.

Poly (A) Polymerase adds a 3' poly (A) tail to the pre-mRNA.

The pre-mRNA is cleaved off the rest of the growing transcript before RNA Polymerase II has stopped transcribing. This cleavage is done by an endonuclease-containing protein complex that binds to an AAUAAA sequence upstream of the cleavage site and to a GU-rich sequence downstream of the cut site. Immediately after the cleavage, Poly (A) Polymerase (PAP), which is also part of the protein complex, catalyzes the addition of up to 200 A nucleotides to the 3' end of the just-cleaved pre-mRNA.

Pre-mRNA Splicing

Eukaryotic genes are composed of exons, which correspond to protein-coding sequences (ex-on signifies that they are expressed), and intervening sequences called introns (int-ron denotes their intervening role), which may be involved in gene regulation, but are removed from the pre-mRNA during processing. Intron sequences in mRNA do not encode functional proteins.

Discovery of Introns

The discovery of introns came as a surprise to researchers in the 1970s who expected that pre-mRNAs would specify protein sequences without further processing, as they had observed in prokaryotes. The genes of higher eukaryotes very often contain one or more introns. While these regions may correspond to regulatory sequences, the biological significance of having many introns or having very long introns in a gene is unclear. It is possible that introns slow down gene expression because it takes longer to transcribe pre-mRNAs with lots of introns. Alternatively, introns may be nonfunctional sequence remnants left over from the fusion of ancient genes throughout evolution. This is supported by the fact that separate exons often encode separate protein subunits or domains. For the most part, the sequences of introns can be mutated without ultimately affecting the protein product.

Intron Processing

All introns in a pre-mRNA must be completely and precisely removed before protein synthesis. If the process errs by even a single nucleotide, the reading frame of the rejoined exons would shift, and the resulting protein would be dysfunctional. The process of removing introns and reconnecting exons is called splicing. Introns are removed and degraded while the pre-mRNA is still in the nucleus. Splicing occurs by a sequence-specific mechanism that ensures introns will be removed and exons rejoined with the accuracy and precision of a single nucleotide. The splicing of pre-mRNAs is conducted by complexes of proteins and RNA molecules called spliceosomes .

Pre-mRNA splicing

Pre-mRNA splicing

Pre-mRNA splicing involves the precise removal of introns from the primary RNA transcript. The splicing process is catalyzed by large complexes called spliceosomes. Each spliceosome is composed of five subunits called snRNPs. The spliceseome's actions result in the splicing together of the two exons and the release of the intron in a lariat form.

Each spliceosome is composed of five subunits called snRNPs (for small nuclear ribonucleoparticles, and pronounced "snurps".) Each snRNP is itself a complex of proteins and a special type of RNA found only in the nucleus called snRNAs (small nuclear RNAs). Spliceosomes recognize sequences at the 5' end of the intron because introns always start with the nucleotides GU and they recognize sequences at the 3' end of the intron because they always end with the nucleotides AG. The spliceosome cleaves the pre-mRNA's sugar phosphate backbone at the G that starts the intron and then covalently attaches that G to an internal A nucleotide within the intron. Then the spliceosme connects the 3' end of the first exon to the 5' end of the following exon, cleaving the 3' end of the intron in the process. This results in the splicing together of the two exons and the release of the intron in a lariat form.

Mechanism of pre-mRNA splicing.

Mechanism of pre-mRNA splicing.

The snRNPs of the spliceosome were left out of this figure, but it shows the sites within the intron whose interactions are catalyzed by the spliceosome. Initially, the conserved G which starts an intron is cleaved from the 3' end of the exon upstream to it and the G is covalently attached to an internal A within the intron. Then the 3' end of the just-released exon is joined to the 5' end of the next exon, cleaving the bond that attaches the 3' end of the intron to its adjacent exon. This both joins the two exons and removes the intron in lariat form.

15.4.2: Processing of tRNAs and rRNAs

rRNA and tRNA are structural molecules that aid in protein synthesis but are not themselves translated into protein.

Learning Objective

Describe how pre-rRNAs and pre-tRNAs are processed into mature rRNAs and tRNAs.

Key Points

  • Ribosomal RNA (rRNA) is a structural molecule that makes up over half of the mass of a ribosome and aids in protein synthesis.
  • Transfer RNA (tRNA) recognizes a codon on mRNA and brings the appropriate amino acid to that site.
  • rRNAs are processed from larger pre-rRNAs by trimming the larger rRNAs down and methylating some of the nucleotides.
  • tRNAs are processed from pre-tRNAs by trimming both ends of the pre-tRNA, adding a CCA trinucleotide to the 3' end, if needed, removing any introns present, and chemically modified 12 nucleotides on average per tRNA.

Key Term

anticodon

a sequence of three nucleotides in transfer RNA that binds to the complementary triplet (codon) in messenger RNA, specifying an amino acid during protein synthesis

Processing of tRNAs and rRNAs

The tRNAs and rRNAs are structural molecules that have roles in protein synthesis; however, these RNAs are not themselves translated. In eukaryotes, pre-rRNAs are transcribed, processed, and assembled into ribosomes in the nucleolus, while pre-tRNAs are transcribed and processed in the nucleus and then released into the cytoplasm where they are linked to free amino acids for protein synthesis.

Ribosomal RNA (rRNA)

The four rRNAs in eukaryotes are first transcribed as two long precursor molecules. One contains just the pre-rRNA that will be processed into the 5S rRNA; the other spans the 28S, 5.8S, and 18S rRNAs. Enzymes then cleave the precursors into subunits corresponding to each rRNA. In bacteria, there are only three rRNAs and all are transcribed in one long precursor molecule that is cleaved into the individual rRNAs. Some of the bases of pre-rRNAs are methylated for added stability. Mature rRNAs make up 50-60% of each ribosome. Some of a ribosome's RNA molecules are purely structural, whereas others have catalytic or binding activities.

The eukaryotic ribosome is composed of two subunits: a large subunit (60S) and a small subunit (40S). The 60S subunit is composed of the 28S rRNA, 5.8S rRNA, 5S rRNA, and 50 proteins. The 40S subunit is composed of the 18S rRNA and 33 proteins. The bacterial ribosome is composed of two similar subunits, with slightly different components. The bacterial large subunit is called the 50S subunit and is composed of the 23S rRNA, 5S rRNA, and 31 proteins, while the bacterial small subunit is called the 30S subunit and is composed of the 16S rRNA and 21 proteins.

The two subunits join to constitute a functioning ribosome that is capable of creating proteins.

Transfer RNA (tRNA)

Each different tRNA binds to a specific amino acid and transfers it to the ribosome. Mature tRNAs take on a three-dimensional structure through intramolecular basepairing to position the amino acid binding site at one end and the anticodon in an unbasepaired loop of nucleotides at the other end. The anticodon is a three-nucleotide sequence, unique to each different tRNA, that interacts with a messenger RNA (mRNA) codon through complementary base pairing.

There are different tRNAs for the 21 different amino acids. Most amino acids can be carried by more than one tRNA.

.

Structure of tRNA

Structure of tRNA

This is a space-filling model of a tRNA molecule that adds the amino acid phenylalanine to a growing polypeptide chain. The anticodon AAG binds the codon UUC on the mRNA. The amino acid phenylalanine is attached to the other end of the tRNA.

In all organisms, tRNAs are transcribed in a pre-tRNA form that requires multiple processing steps before the mature tRNA is ready for use in translation. In bacteria, multiple tRNAs are often transcribed as a single RNA. The first step in their processing is the digestion of the RNA to release individual pre-tRNAs. In archaea and eukaryotes, each pre-tRNA is transcribed as a separate transcript.

The processing to convert the pre-tRNA to a mature tRNA involves five steps.

1. The 5' end of the pre-tRNA, called the 5' leader sequence, is cleaved off.

2. The 3' end of the pre-tRNA is cleaved off.

3. In all eukaryote pre-tRNAs, but in only some bacterial and archaeal pre-tRNAs, a CCA sequence of nucleotides is added to the 3' end of the pre-tRNA after the original 3' end is trimmed off. Some bacteria and archaea pre-tRNAs already have the CCA encoded in their transcript immediately upstream of the 3' cleavage site, so they don't need to add one. The CCA at the 3' end of the mature tRNA will be the site at which the tRNA's amino acid will be added.

4. Multiple nucleotides in the pre-tRNA are chemically modified, altering their nitorgen bases. On average about 12 nucleotides are modified per tRNA. The most common modifications are the conversion of adenine (A) to pseudouridine (ψ), the conversion of adenine to inosine (I), and the conversion of uridine to dihydrouridine (D). But over 100 other modifications can occur.

5. A significant number of eukaryotic and archaeal pre-tRNAs have introns that have to be spliced out. Introns are rarer in bacterial pre-tRNAs, but do occur occasionally and are spliced out.

After processing, the mature pre-tRNA is ready to have its cognate amino acid attached. The cognate amino acid for a tRNA is the one specified by its anticodon. Attaching this amino acid is called charging the tRNA. In eukaryotes, the mature tRNA is generated in the nucleus, and then exported to the cytoplasm for charging.

Processing of a pre-tRNA.

Processing of a pre-tRNA.

A typical pre-tRNA undergoing processing steps to generate a mature tRNA ready to have its cognate amino acid attached. Nucleotides that are cleaved away are shown in green. Chemically-modified nucleotides are in yellow, as is the CAA trinucleotide that is added to the 3' end of the pre-tRNA during processing. The anticodon nucleotides are shown in a lighter shade of red.

Attributions

  • mRNA Processing
    • "OpenStax College, Biology. October 16, 2013." http://openstaxcollege.org/textbooks/biology. OpenStax CNX CC BY-SA 3.0.
    • "RNA splicing." http://en.wikipedia.org/wiki/RNA_splicing. Wikipedia CC BY-SA 3.0.
    • "spliceosome." http://en.wiktionary.org/wiki/spliceosome. Wiktionary CC BY-SA 3.0.
    • "OpenStax College, Biology. October 16, 2013." http://cnx.org/content/m44532/latest/?collection=col11448/latest. OpenStax CNX CC BY 3.0.
    • "intron." http://en.wiktionary.org/wiki/intron. Wiktionary CC BY-SA 3.0.
    • "moiety." http://en.wiktionary.org/wiki/moiety. Wiktionary CC BY-SA 3.0.
    • "Original figure by Ross Whitwam. Licensed CC BY-SA 4.0." Ross Whitwam CC BY-SA 3.0.
    • "RNA Splicing." http://en.wikipedia.org/wiki/RNA_splicing. WikiPedia CC BY-SA 3.0.
    • "OpenStax College, RNA Processing in Eukaryotes. October 16, 2013." http://cnx.org/content/m44532/latest/Figure_15_04_02.png. OpenStax CNX CC BY 3.0.
    • "An Introduction to Molecular Biology/Transcription of RNA and its modification." http://en.wikibooks.org/wiki/An_Introduction_to_Molecular_Biology/Transcription_of_RNA_and_its_modification%23Splicing_of_RNA. Wikibooks CC BY-SA 3.0.
  • Processing of tRNAs and rRNAs
    • "OpenStax College, Biology. October 16, 2013." http://openstaxcollege.org/textbooks/biology. OpenStax CNX CC BY-SA 3.0.
    • "OpenStax College, Biology. October 16, 2013." http://cnx.org/content/m44532/latest/?collection=col11448/latest. OpenStax CNX CC BY 3.0.
    • "anticodon." http://en.wiktionary.org/wiki/anticodon. Wiktionary CC BY-SA 3.0.
    • "Structural Biochemistry/Nucleic Acid/RNA/Ribosomal RNA (rRNA)." http://en.wikibooks.org/wiki/Structural_Biochemistry/Nucleic_Acid/RNA/Ribosomal_RNA_(rRNA). Wikibooks CC BY-SA 3.0.
    • "Structural Biochemistry/Cell Organelles/Ribosome." http://en.wikibooks.org/wiki/Structural_Biochemistry/Cell_Organelles/Ribosome. Wikibooks CC BY-SA 3.0.
    • "Structural Biochemistry/Nucleic Acid/RNA/Transfer RNA (tRNA)." http://en.wikibooks.org/wiki/Structural_Biochemistry/Nucleic_Acid/RNA/Transfer_RNA_(tRNA). Wikibooks CC BY-SA 3.0.
    • "Original figure by Ross Whitwam. Licensed CC BY-SA 4.0." Ross Whitwam CC BY-SA 3.0.
    • "OpenStax College, RNA Processing in Eukaryotes. October 16, 2013." http://cnx.org/content/m44532/latest/Figure_15_04_03.jpg. OpenStax CNX CC BY 3.0.
    • "Structural Biochemistry/Nucleic Acid/RNA/Transfer RNA (tRNA)." http://en.wikibooks.org/wiki/Structural_Biochemistry/Nucleic_Acid/RNA/Transfer_RNA_(tRNA). Wikibooks CC BY-SA 3.0.

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