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Boundless Biology: 5.2: Passive Transport

Boundless Biology
5.2: Passive Transport
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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

5.2: Passive Transport

5.2.1: The Role of Passive Transport

Passive transport, such as diffusion and osmosis, moves materials of small molecular weight across membranes.

Learning Objective

Indicate the manner in which various materials cross the cell membrane

Key Points

  • Plasma membranes are selectively permeable; if they were to lose this selectivity, the cell would no longer be able to sustain itself.
  • In passive transport, substances simply move from an area of higher concentration to an area of lower concentration, which does not require the input of energy.
  • Concentration gradient, size of the particles that are diffusing, and temperature of the system affect the rate of diffusion.
  • Some materials diffuse readily through the membrane, but others require specialized proteins, such as channels and transporters, to carry them into or out of the cell.

Key Terms

concentration gradient

A concentration gradient is present when a membrane separates two different concentrations of molecules.

passive transport

A movement of biochemicals and other atomic or molecular substances across membranes that does not require an input of chemical energy.

permeable

Of or relating to substance, substrate, membrane or material that absorbs or allows the passage of fluids.

Introduction: Passive Transport

Plasma membranes must allow or prevent certain substances from entering or leaving a cell. In other words, plasma membranes are selectively permeable; they allow some substances to pass through, but not others. If they were to lose this selectivity, the cell would no longer be able to sustain itself, and it would be destroyed. Some cells require larger amounts of specific substances than other cells; they must have a way of obtaining these materials from extracellular fluids. This may happen passively, as certain materials move back and forth, or the cell may have special mechanisms that facilitate transport. Some materials are so important to a cell that it spends some of its energy (hydrolyzing adenosine triphosphate (ATP)) to obtain these materials. Red blood cells use some of their energy to do this. All cells spend the majority of their energy to maintain an imbalance of sodium and potassium ions between the interior and exterior of the cell.

The most direct forms of membrane transport are passive. Passive transport is a naturally-occurring phenomenon and does not require the cell to exert any of its energy to accomplish the movement. In passive transport, substances move from an area of higher concentration to an area of lower concentration . A physical space in which there is a range of concentrations of a single substance is said to have a concentration gradient.

Passive Transport

Passive Transport

Diffusion is a type of passive transport. Diffusion through a permeable membrane moves a substance from an area of high concentration (extracellular fluid, in this case) down its concentration gradient (into the cytoplasm).

The passive forms of transport, diffusion and osmosis, move materials of small molecular weight across membranes. Substances diffuse from areas of high concentration to areas of lower concentration; this process continues until the substance is evenly distributed in a system. In solutions containing more than one substance, each type of molecule diffuses according to its own concentration gradient, independent of the diffusion of other substances. Many factors can affect the rate of diffusion, including, but not limited to, concentration gradient, size of the particles that are diffusing, and temperature of the system.

In living systems, diffusion of substances in and out of cells is mediated by the plasma membrane. Some materials diffuse readily through the membrane, but others are hindered; their passage is made possible by specialized proteins, such as channels and transporters. The chemistry of living things occurs in aqueous solutions; balancing the concentrations of those solutions is an ongoing problem. In living systems, diffusion of some substances would be slow or difficult without membrane proteins that facilitate transport.

5.2.2: Selective Permeability

The hydrophobic and hydrophilic regions of plasma membranes aid the diffusion of some molecules and hinder the diffusion of others.

Learning Objective

Describe how membrane permeability, concentration gradient, and molecular properties affect biological diffusion rates.

Key Points

  • The interior and exterior surfaces of the plasma membrane are not identical, which adds to the selective permeability of the membrane.
  • Fat soluble substances are able to pass easily to the hydrophobic interior of the plasma membrane and diffuse into the cell.
  • Polar molecules and charged molecules do not diffuse easily through the lipid core of the plasma membrane and must be transported across by proteins, sugars, or amino acids.

Key Terms

amphiphilic

Having one surface consisting of hydrophilic amino acids and the opposite surface consisting of hydrophobic (or lipophilic) ones.

polar

a separation of electric charge leading to a molecule or its chemical groups having an electric dipole

Selective Permeability

Plasma membranes are asymmetric: the interior of the membrane is not identical to the exterior of the membrane. In fact, there is a considerable difference between the array of phospholipids and proteins between the two leaflets that form a membrane . On the interior of the membrane, some proteins serve to anchor the membrane to fibers of the cytoskeleton. There are peripheral proteins on the exterior of the membrane that bind elements of the extracellular matrix. Carbohydrates, attached to lipids or proteins, are also found on the exterior surface of the plasma membrane. These carbohydrate complexes help the cell bind substances that the cell needs in the extracellular fluid. This adds considerably to the selective nature of plasma membranes.

Asymmetry in Plasma Membranes

Asymmetry in Plasma Membranes

The exterior surface of the plasma membrane is not identical to the interior surface of the same membrane.

Recall that plasma membranes are amphiphilic; that is, they have hydrophilic and hydrophobic regions. This characteristic helps the movement of some materials through the membrane and hinders the movement of others. Lipid-soluble material with a low molecular weight can easily slip through the hydrophobic lipid core of the membrane. Substances such as the fat-soluble vitamins A, D, E, and K readily pass through the plasma membranes in the digestive tract and other tissues. Fat-soluble drugs and hormones also gain easy entry into cells and are readily transported into the body's tissues and organs. Molecules of oxygen and carbon dioxide have no charge and so pass through membranes by simple diffusion.

Polar substances present problems for the membrane. While some polar molecules connect easily with the outside of a cell, they cannot readily pass through the lipid core of the plasma membrane. Additionally, while small ions could easily slip through the spaces in the mosaic of the membrane, their charge prevents them from doing so. Ions such as sodium, potassium, calcium, and chloride must have special means of penetrating plasma membranes. Simple sugars and amino acids also need help with transport across plasma membranes, achieved by various transmembrane proteins (channels).

5.2.3: Diffusion

Diffusion is a process of passive transport in which molecules move from an area of higher concentration to one of lower concentration.

Learning Objective

Describe diffusion and the factors that affect how materials move across the cell membrane.

Key Points

  • Substances diffuse according to their concentration gradient; within a system, different substances in the medium will each diffuse at different rates according to their individual gradients.
  • After a substance has diffused completely through a space, removing its concentration gradient, molecules will still move around in the space, but there will be no net movement of the number of molecules from one area to another, a state known as dynamic equilibrium.
  • Several factors affect the rate of diffusion of a solute including the mass of the solute, the temperature of the environment, the solvent density, and the distance traveled.

Key Terms

concentration gradient

A concentration gradient is present when a membrane separates two different concentrations of molecules.

diffusion

The passive movement of a solute across a permeable membrane

Example

  • When someone is cooking food in a kitchen, the smell begins to waft through the house, and eventually everyone can tell what's for dinner! This is due to the diffusion of odor molecules through the air, from an area of high concentration (the kitchen) to areas of low concentration (your upstairs bedroom).

Diffusion

Diffusion is a passive process of transport. A single substance tends to move from an area of high concentration to an area of low concentration until the concentration is equal across a space . You are familiar with diffusion of substances through the air. For example, think about someone opening a bottle of ammonia in a room filled with people. The ammonia gas is at its highest concentration in the bottle; its lowest concentration is at the edges of the room. The ammonia vapor will diffuse, or spread away, from the bottle; gradually, more and more people will smell the ammonia as it spreads. Materials move within the cell's cytosol by diffusion, and certain materials move through the plasma membrane by diffusion. Diffusion expends no energy. On the contrary, concentration gradients are a form of potential energy, dissipated as the gradient is eliminated.

Diffusion

Diffusion

Diffusion through a permeable membrane moves a substance from an area of high concentration (extracellular fluid, in this case) down its concentration gradient (into the cytoplasm).

Each separate substance in a medium, such as the extracellular fluid, has its own concentration gradient independent of the concentration gradients of other materials. In addition, each substance will diffuse according to that gradient. Within a system, there will be different rates of diffusion of the different substances in the medium.

Factors That Affect Diffusion

Molecules move constantly in a random manner at a rate that depends on their mass, their environment, and the amount of thermal energy they possess, which in turn is a function of temperature. This movement accounts for the diffusion of molecules through whatever medium in which they are localized. A substance will tend to move into any space available to it until it is evenly distributed throughout it. After a substance has diffused completely through a space removing its concentration gradient, molecules will still move around in the space, but there will be no net movement of the number of molecules from one area to another. This lack of a concentration gradient in which there is no net movement of a substance is known as dynamic equilibrium. While diffusion will go forward in the presence of a concentration gradient of a substance, several factors affect the rate of diffusion:

  • Extent of the concentration gradient: The greater the difference in concentration, the more rapid the diffusion. The closer the distribution of the material gets to equilibrium, the slower the rate of diffusion becomes.
  • Mass of the molecules diffusing: Heavier molecules move more slowly; therefore, they diffuse more slowly. The reverse is true for lighter molecules.
  • Temperature: Higher temperatures increase the energy and therefore the movement of the molecules, increasing the rate of diffusion. Lower temperatures decrease the energy of the molecules, thus decreasing the rate of diffusion.
  • Solvent density: As the density of a solvent increases, the rate of diffusion decreases. The molecules slow down because they have a more difficult time getting through the denser medium. If the medium is less dense, diffusion increases. Because cells primarily use diffusion to move materials within the cytoplasm, any increase in the cytoplasm's density will inhibit the movement of the materials. An example of this is a person experiencing dehydration. As the body's cells lose water, the rate of diffusion decreases in the cytoplasm, and the cells' functions deteriorate. Neurons tend to be very sensitive to this effect. Dehydration frequently leads to unconsciousness and possibly coma because of the decrease in diffusion rate within the cells.
  • Solubility: As discussed earlier, nonpolar or lipid-soluble materials pass through plasma membranes more easily than polar materials, allowing a faster rate of diffusion.
  • Surface area and thickness of the plasma membrane: Increased surface area increases the rate of diffusion, whereas a thicker membrane reduces it.
  • Distance travelled: The greater the distance that a substance must travel, the slower the rate of diffusion. This places an upper limitation on cell size. A large, spherical cell will die because nutrients or waste cannot reach or leave the center of the cell. Therefore, cells must either be small in size, as in the case of many prokaryotes, or be flattened, as with many single-celled eukaryotes.

A variation of diffusion is the process of filtration. In filtration, material moves according to its concentration gradient through a membrane; sometimes the rate of diffusion is enhanced by pressure, causing the substances to filter more rapidly. This occurs in the kidney where blood pressure forces large amounts of water and accompanying dissolved substances, or solutes, out of the blood and into the renal tubules. The rate of diffusion in this instance is almost totally dependent on pressure. One of the effects of high blood pressure is the appearance of protein in the urine, which is "squeezed through" by the abnormally high pressure.

5.2.4: Facilitated transport

Facilitated diffusion is a process by which molecules are transported across the plasma membrane with the help of membrane proteins.

Learning Objective

Explain why and how passive transport occurs

Key Points

  • A concentration gradient exists that would allow ions and polar molecules to diffuse into the cell, but these materials are repelled by the hydrophobic parts of the cell membrane.
  • Facilitated diffusion uses integral membrane proteins to move polar or charged substances across the hydrophobic regions of the membrane.
  • Channel proteins can aid in the facilitated diffusion of substances by forming a hydrophilic passage through the plasma membrane through which polar and charged substances can pass.
  • Channel proteins can be open at all times, constantly allowing a particular substance into or out of the cell, depending on the concentration gradient; or they can be gated and can only be opened by a particular biological signal.
  • Carrier proteins aid in facilitated diffusion by binding a particular substance, then altering their shape to bring that substance into or out of the cell.

Key Terms

membrane protein

Proteins that are attached to, or associated with the membrane of a cell or an organelle.

facilitated diffusion

The spontaneous passage of molecules or ions across a biological membrane passing through specific transmembrane integral proteins.

Example

  • Channel-mediated facilitated diffusion functions much like a bridge over a river that must raise and lower in order to allow boats to pass. When the bridge is lowered, boats cannot pass through to the other side of the river. Similarly, a gated channel protein often remains closed, not allowing substances into the cell until it receives a signal (like the binding of an ion) to open. When this signal is received, the bridge (gate) opens, allowing the boats (substance) to pass through the bridge and into the other side of the river (cell).

Facilitated transport

Facilitated transport is a type of passive transport. Unlike simple diffusion where materials pass through a membrane without the help of proteins, in facilitated transport, also called facilitated diffusion, materials diffuse across the plasma membrane with the help of membrane proteins. A concentration gradient exists that would allow these materials to diffuse into the cell without expending cellular energy. However, these materials are ions or polar molecules that are repelled by the hydrophobic parts of the cell membrane. Facilitated transport proteins shield these materials from the repulsive force of the membrane, allowing them to diffuse into the cell.

The material being transported is first attached to protein or glycoprotein receptors on the exterior surface of the plasma membrane. This allows the material that is needed by the cell to be removed from the extracellular fluid. The substances are then passed to specific integral proteins that facilitate their passage. Some of these integral proteins are collections of beta-pleated sheets that form a channel through the phospholipid bilayer. Others are carrier proteins which bind with the substance and aid its diffusion through the membrane.

Channels

The integral proteins involved in facilitated transport are collectively referred to as transport proteins; they function as either channels for the material or carriers. In both cases, they are transmembrane proteins. Channels are specific for the substance that is being transported. Channel proteins have hydrophilic domains exposed to the intracellular and extracellular fluids; they additionally have a hydrophilic channel through their core that provides a hydrated opening through the membrane layers . Passage through the channel allows polar compounds to avoid the nonpolar central layer of the plasma membrane that would otherwise slow or prevent their entry into the cell. Aquaporins are channel proteins that allow water to pass through the membrane at a very high rate.

Channel Proteins in Facilitated Transport

Channel Proteins in Facilitated Transport

Facilitated transport moves substances down their concentration gradients. They may cross the plasma membrane with the aid of channel proteins.

Channel proteins are either open at all times or they are "gated," which controls the opening of the channel. The attachment of a particular ion to the channel protein may control the opening or other mechanisms or substances may be involved. In some tissues, sodium and chloride ions pass freely through open channels, whereas in other tissues, a gate must be opened to allow passage. An example of this occurs in the kidney, where both forms of channels are found in different parts of the renal tubules. Cells involved in the transmission of electrical impulses, such as nerve and muscle cells, have gated channels for sodium, potassium, and calcium in their membranes. Opening and closing of these channels changes the relative concentrations on opposing sides of the membrane of these ions, resulting in the facilitation of electrical transmission along membranes (in the case of nerve cells) or in muscle contraction (in the case of muscle cells).

Carrier Proteins

Another type of protein embedded in the plasma membrane is a carrier protein. This protein binds a substance and, in doing so, triggers a change of its own shape, moving the bound molecule from the outside of the cell to its interior; depending on the gradient, the material may move in the opposite direction . Carrier proteins are typically specific for a single substance. This adds to the overall selectivity of the plasma membrane. The exact mechanism for the change of shape is poorly understood. Proteins can change shape when their hydrogen bonds are affected, but this may not fully explain this mechanism. Each carrier protein is specific to one substance, and there are a finite number of these proteins in any membrane. This can cause problems in transporting enough of the material for the cell to function properly.

Carrier Proteins

Carrier Proteins

Some substances are able to move down their concentration gradient across the plasma membrane with the aid of carrier proteins. Carrier proteins change shape as they move molecules across the membrane.

An example of this process occurs in the kidney. Glucose, water, salts, ions, and amino acids needed by the body are filtered in one part of the kidney. This filtrate, which includes glucose, is then reabsorbed in another part of the kidney. Because there are only a finite number of carrier proteins for glucose, if more glucose is present than the proteins can handle, the excess is not transported; it is excreted from the body in the urine. In a diabetic individual, this is described as "spilling glucose into the urine." A different group of carrier proteins called glucose transport proteins, or GLUTs, are involved in transporting glucose and other hexose sugars through plasma membranes within the body.

Channel and carrier proteins transport material at different rates. Channel proteins transport much more quickly than do carrier proteins. Channel proteins facilitate diffusion at a rate of tens of millions of molecules per second, whereas carrier proteins work at a rate of a thousand to a million molecules per second.

5.2.5: Osmosis

Osmosis is the movement of water across a membrane from an area of low solute concentration to an area of high solute concentration.

Learning Objective

Describe the process of osmosis and explain how concentration gradient affects osmosis

Key Points

  • Osmosis occurs according to the concentration gradient of water across the membrane, which is inversely proportional to the concentration of solutes.
  • Osmosis occurs until the concentration gradient of water goes to zero or until the hydrostatic pressure of the water balances the osmotic pressure.
  • Osmosis occurs when there is a concentration gradient of a solute within a solution, but the membrane does not allow diffusion of the solute.

Key Terms

solute

Any substance that is dissolved in a liquid solvent to create a solution

osmosis

The net movement of solvent molecules from a region of high solvent potential to a region of lower solvent potential through a partially permeable membrane

semipermeable membrane

A type of biological membrane that will allow certain molecules or ions to pass through it by diffusion and occasionally by specialized facilitated diffusion

Osmosis and Semipermeable Membranes

Osmosis is the movement of water through a semipermeable membrane according to the concentration gradient of water across the membrane, which is inversely proportional to the concentration of solutes. Semipermeable membranes, also termed selectively permeable membranes or partially permeable membranes, allow certain molecules or ions to pass through by diffusion.

While diffusion transports materials across membranes and within cells, osmosis transports only water across a membrane. The semipermeable membrane limits the diffusion of solutes in the water. Not surprisingly, the aquaporin proteins that facilitate water movement play a large role in osmosis, most prominently in red blood cells and the membranes of kidney tubules.

Mechanism of Osmosis

Osmosis is a special case of diffusion. Water, like other substances, moves from an area of high concentration to one of low concentration. An obvious question is what makes water move at all? Imagine a beaker with a semipermeable membrane separating the two sides or halves. On both sides of the membrane the water level is the same, but there are different concentrations of a dissolved substance, or solute, that cannot cross the membrane (otherwise the concentrations on each side would be balanced by the solute crossing the membrane). If the volume of the solution on both sides of the membrane is the same but the concentrations of solute are different, then there are different amounts of water, the solvent, on either side of the membrane. If there is more solute in one area, then there is less water; if there is less solute in one area, then there must be more water.

To illustrate this, imagine two full glasses of water. One has a single teaspoon of sugar in it, whereas the second one contains one-quarter cup of sugar. If the total volume of the solutions in both cups is the same, which cup contains more water? Because the large amount of sugar in the second cup takes up much more space than the teaspoon of sugar in the first cup, the first cup has more water in it.

Osmosis

Osmosis

In osmosis, water always moves from an area of higher water concentration to one of lower concentration. In the diagram shown, the solute cannot pass through the selectively permeable membrane, but the water can.

Returning to the beaker example, recall that it has a mixture of solutes on either side of the membrane. A principle of diffusion is that the molecules move around and will spread evenly throughout the medium if they can. However, only the material capable of passing through the membrane will diffuse through it. In this example, the solute cannot diffuse through the membrane, but the water can. Water has a concentration gradient in this system. Thus, water will diffuse down its concentration gradient, crossing the membrane to the side where it is less concentrated. This diffusion of water through the membrane—osmosis—will continue until the concentration gradient of water goes to zero or until the hydrostatic pressure of the water balances the osmotic pressure. In the beaker example, this means that the level of fluid in the side with a higher solute concentration will go up .

5.2.6: Tonicity

Tonicity, which is directly related to the osmolarity of a solution, affects osmosis by determining the direction of water flow.

Learning Objective

Define tonicity and describe its relevance to osmosis

Key Points

  • Osmolarity describes the total solute concentration of a solution; solutions with a low solute concentration have a low osmolarity, while those with a high osmolarity have a high solute concentration.
  • Water moves from the side of the membrane with lower osmolarity (and more water) to the side with higher osmolarity (and less water).
  • In a hypotonic solution, the extracellular fluid has a lower osmolarity than the fluid inside the cell; water enters the cell.
  • In a hypertonic solution, the extracellular fluid has a higher osmolarity than the fluid inside the cell; water leaves the cell.
  • In an isotonic solution, the extracellular fluid has the same osmolarity as the cell; there will be no net movement of water into or out of the cell.

Key Terms

isotonic

having the same osmotic pressure

hypertonic

having a greater osmotic pressure than another

hypotonic

Having a lower osmotic pressure than another; a cell in this environment causes water to enter the cell, causing it to swell.

osmolarity

The osmotic concentration of a solution, normally expressed as osmoles of solute per litre of solution.

Example

  • Tonicity is the reason why salt water fish cannot live in fresh water and vice versa. A salt water fish's cells have evolved to have a very high solute concentration to match the high osmolarity of the salt water they live in. If you place a salt water fish in fresh water, which has a low osmolarity, water in the environment will flow into the cells of the fish, eventually causing them to burst and killing the fish.

Tonicity

Tonicity describes how an extracellular solution can change the volume of a cell by affecting osmosis. A solution's tonicity often directly correlates with the osmolarity of the solution. Osmolarity describes the total solute concentration of the solution. A solution with low osmolarity has a greater number of water molecules relative to the number of solute particles; a solution with high osmolarity has fewer water molecules with respect to solute particles. In a situation in which solutions of two different osmolarities are separated by a membrane permeable to water, though not to the solute, water will move from the side of the membrane with lower osmolarity (and more water) to the side with higher osmolarity (and less water). This effect makes sense if you remember that the solute cannot move across the membrane, and thus the only component in the system that can move—the water—moves along its own concentration gradient. An important distinction that concerns living systems is that osmolarity measures the number of particles (which may be molecules) in a solution. Therefore, a solution that is cloudy with cells may have a lower osmolarity than a solution that is clear if the second solution contains more dissolved molecules than there are cells.

Hypotonic Solutions

Three terms—hypotonic, isotonic, and hypertonic—are used to relate the osmolarity of a cell to the osmolarity of the extracellular fluid that contains the cells . In a hypotonic situation, the extracellular fluid has lower osmolarity than the fluid inside the cell, and water enters the cell. (In living systems, the point of reference is always the cytoplasm, so the prefix hypo- means that the extracellular fluid has a lower concentration of solutes, or a lower osmolarity, than the cell cytoplasm. ) It also means that the extracellular fluid has a higher concentration of water in the solution than does the cell. In this situation, water will follow its concentration gradient and enter the cell, causing the cell to expand.

Changes in Cell Shape Due to Dissolved Solutes

Changes in Cell Shape Due to Dissolved Solutes

Osmotic pressure changes the shape of red blood cells in hypertonic, isotonic, and hypotonic solutions.

Hypertonic Solutions

As for a hypertonic solution, the prefix hyper- refers to the extracellular fluid having a higher osmolarity than the cell's cytoplasm; therefore, the fluid contains less water than the cell does. Because the cell has a relatively higher concentration of water, water will leave the cell, and the cell will shrink.

Isotonic Solutions

In an isotonic solution, the extracellular fluid has the same osmolarity as the cell. If the osmolarity of the cell matches that of the extracellular fluid, there will be no net movement of water into or out of the cell, although water will still move in and out.

Blood cells and plant cells in hypertonic, isotonic, and hypotonic solutions take on characteristic appearances. Cells in an isotonic solution retain their shape. Cells in a hypotonic solution swell as water enters the cell, and may burst if the concentration gradient is large enough between the inside and outside of the cell. Cells in a hypertonic solution shrink as water exits the cell, becoming shriveled.

5.2.7: Osmoregulation

Osmoregulation is the process by which living things regulate the effects of osmosis in order to protect cellular integrity.

Learning Objective

Describe the methods by which organisms regulate the effects of osmosis on their cells

Key Points

  • In a hypotonic environment, water enters a cell, causing it to swell and burst (lyse).
  • In a hypertonic environment, water leaves a cell, causing it to shrink and die.
  • In plants, in a hypotonic environment, the inflow of water produces turgor pressure, which stiffens the cell walls of the plant, while in a hypertonic environment, turgor pressure is lost, and the plant wilts.
  • Some protists that lack cell walls have contractile vacuoles, which collect excess water from the cell and pump it out, keeping the cell from lysing.
  • Freshwater fish living in a hypotonic environment take in salt through their gills and excrete diluted urine to rid themselves of excess water; saltwater fish living in a hypertonic environment secrete salt through their gills and excrete highly concentrated urine.
  • In vertebrates, osmoreceptors monitor the concentration of solutes in the blood, and albumin controls osmotic pressure in tissues.

Key Terms

osmoregulation

the homeostatic regulation of osmotic pressure in the body in order to maintain a constant water content

hypertonic

having a greater osmotic pressure than another

hypotonic

Having a lower osmotic pressure than another; a cell in this environment causes water to enter the cell, causing it to swell.

Tonicity in Living Systems

Tonicity is the ability of a solution to exert an osmotic pressure upon a membrane. There are three types of tonicity: hypotonic, hypertonic, and isotonic. In a hypotonic environment, water enters a cell, and the cell swells. In a hypertonic solution, water leaves a cell and the cell shrinks. In an isotonic condition, the relative concentrations of solute and solvent are equal on both sides of the membrane. There is no net water movement; therefore, there is no change in the size of the cell. If either the hypo- or hyper- condition goes to excess, the cell's functions become compromised, and the cell may be destroyed.

A red blood cell will burst, or lyse, when it swells beyond the plasma membrane's capability to expand. The membrane resembles a mosaic with discrete spaces between the molecules comprising it. If the cell swells and the spaces between the lipids and proteins become too large, the cell will break apart.

In contrast, when excessive amounts of water leave a red blood cell, the cell shrinks, or crenates. This has the effect of concentrating the solutes left in the cell, making the cytosol denser and interfering with diffusion within the cell. The cell's ability to function will be compromised and may also result in the death of the cell.

Turgor Pressure and Tonicity in a Plant Cell

Turgor Pressure and Tonicity in a Plant Cell

The turgor pressure within a plant cell depends on the tonicity of the solution in which it is bathed.

Osmoregulation

Various living things have ways of controlling the effects of osmosis—a mechanism called osmoregulation. Some organisms, such as plants, fungi, bacteria, and some protists, have cell walls that surround the plasma membrane and prevent cell lysis in a hypotonic solution. The plasma membrane can only expand to the limit of the cell wall, so the cell will not lyse. In fact, in plants, the cellular environment is always slightly hypotonic to the cytoplasm, and water will always enter a cell if water is available. This inflow of water produces turgor pressure, which stiffens the cell walls of the plant. In nonwoody plants, turgor pressure supports the plant. Conversely, if the plant is not watered, the extracellular fluid will become hypertonic, causing water to leave the cell . In this condition, the cell does not shrink because the cell wall is not flexible. However, the cell membrane detaches from the wall and constricts the cytoplasm. This is called plasmolysis. Plants lose turgor pressure in this condition and wilt .

Turgor Pressure and Plasmolysis

Turgor Pressure and Plasmolysis

Without adequate water, the plant on the left has lost turgor pressure, visible in its wilting; the turgor pressure is restored by watering it (right).

Tonicity is a concern for all living things. For example, paramecia and amoebas, which are protists that lack cell walls, have contractile vacuoles. This vesicle collects excess water from the cell and pumps it out, keeping the cell from lysing as it takes on water from its environment .

Contractile Vacuoles

Contractile Vacuoles

A paramecium's contractile vacuole, here visualized using bright field light microscopy at 480x magnification, continuously pumps water out of the organism's body to keep it from bursting in a hypotonic medium.

Many marine invertebrates have internal salt levels matched to their environments, making them isotonic with the water in which they live. Fish, however, must spend approximately five percent of their metabolic energy maintaining osmotic homeostasis. Freshwater fish live in an environment that is hypotonic to their cells. These fish actively take in salt through their gills and excrete diluted urine to rid themselves of excess water. Saltwater fish live in the reverse environment, which is hypertonic to their cells, and they secrete salt through their gills and excrete highly concentrated urine.

In vertebrates, the kidneys regulate the amount of water in the body. Osmoreceptors are specialized cells in the brain that monitor the concentration of solutes in the blood. If the levels of solutes increase beyond a certain range, a hormone is released that retards water loss through the kidney and dilutes the blood to safer levels. Animals also have high concentrations of albumin (produced by the liver) in their blood. This protein is too large to pass easily through plasma membranes and is a major factor in controlling the osmotic pressures applied to tissues.

Attributions

  • The Role of Passive Transport
    • "Boundless." http://www.boundless.com/. Boundless Learning CC BY-SA 3.0.
    • "passive transport." http://en.wikipedia.org/wiki/passive%20transport. Wikipedia CC BY-SA 3.0.
    • "concentration gradient." http://en.wikipedia.org/wiki/concentration%20gradient. Wikipedia CC BY-SA 3.0.
    • "permeable." http://en.wiktionary.org/wiki/permeable. Wiktionary CC BY-SA 3.0.
    • "OpenStax College, Biology. October 16, 2013." http://cnx.org/content/m44417/latest/?collection=col11448/latest. OpenStax CNX CC BY 3.0.
    • "OpenStax College, Passive Transport. October 16, 2013." http://cnx.org/content/m44417/latest/Figure_05_02_02.jpg. OpenStax CNX CC BY 3.0.
  • Selective Permeability
    • "Boundless." http://www.boundless.com/. Boundless Learning CC BY-SA 3.0.
    • "amphiphilic." http://en.wiktionary.org/wiki/amphiphilic. Wiktionary CC BY-SA 3.0.
    • "polar." http://en.wiktionary.org/wiki/polar. Wiktionary CC BY-SA 3.0.
    • "OpenStax College, Biology. October 16, 2013." http://cnx.org/content/m44417/latest/?collection=col11448/latest. OpenStax CNX CC BY 3.0.
    • "OpenStax College, Passive Transport. October 16, 2013." http://cnx.org/content/m44417/latest/Figure_05_02_01.jpg. OpenStax CNX CC BY 3.0.
  • Diffusion
    • "Boundless." http://www.boundless.com/. Boundless Learning CC BY-SA 3.0.
    • "concentration gradient." http://en.wikipedia.org/wiki/concentration%20gradient. Wikipedia CC BY-SA 3.0.
    • "OpenStax College, Biology. October 16, 2013." http://cnx.org/content/m44417/latest/?collection=col11448/latest. OpenStax CNX CC BY 3.0.
    • "OpenStax College, Passive Transport. October 16, 2013." http://cnx.org/content/m44417/latest/Figure_05_02_02.jpg. OpenStax CNX CC BY 3.0.
  • Facilitated transport
    • "OpenStax College, Biology. October 16, 2013." http://openstaxcollege.org/textbooks/biology. OpenStax CNX CC BY-SA 3.0.
    • "membrane protein." http://en.wikipedia.org/wiki/membrane%20protein. Wikipedia CC BY-SA 3.0.
    • "facilitated diffusion." http://en.wikipedia.org/wiki/facilitated%20diffusion. Wikipedia CC BY-SA 3.0.
    • "OpenStax College, Biology. October 16, 2013." http://cnx.org/content/m44417/latest/?collection=col11448/latest. OpenStax CNX CC BY 3.0.
    • "OpenStax College, Passive Transport. October 16, 2013." http://cnx.org/content/m44417/latest/Figure_05_02_04.jpg. OpenStax CNX CC BY 3.0.
    • "OpenStax College, Passive Transport. October 16, 2013." http://cnx.org/content/m44417/latest/Figure_05_02_05.jpg. OpenStax CNX CC BY 3.0.
  • Osmosis
    • "Boundless." http://www.boundless.com/. Boundless Learning CC BY-SA 3.0.
    • "osmosis." http://en.wiktionary.org/wiki/osmosis. Wiktionary CC BY-SA 3.0.
    • "OpenStax College, Biology. October 16, 2013." http://cnx.org/content/m44417/latest/?collection=col11448/latest. OpenStax CNX CC BY 3.0.
    • "solute." http://en.wiktionary.org/wiki/solute. Wiktionary CC BY-SA 3.0.
    • "OpenStax College, Passive Transport. October 16, 2013." http://cnx.org/content/m44417/latest/Figure_05_02_06.jpg. OpenStax CNX CC BY 3.0.
  • Tonicity
    • "Boundless." http://www.boundless.com/. Boundless Learning CC BY-SA 3.0.
    • "hypertonic." http://en.wiktionary.org/wiki/hypertonic. Wiktionary CC BY-SA 3.0.
    • "OpenStax College, Biology. October 16, 2013." http://cnx.org/content/m44417/latest/?collection=col11448/latest. OpenStax CNX CC BY 3.0.
    • "osmolarity." http://en.wiktionary.org/wiki/osmolarity. Wiktionary CC BY-SA 3.0.
    • "hypotonic." http://en.wiktionary.org/wiki/hypotonic. Wiktionary CC BY-SA 3.0.
    • "isotonic." http://en.wiktionary.org/wiki/isotonic. Wiktionary CC BY-SA 3.0.
    • "OpenStax College, Passive Transport. October 16, 2013." http://cnx.org/content/m44417/latest/Figure_05_02_07.png. OpenStax CNX CC BY 3.0.
  • Osmoregulation
    • "Boundless." http://www.boundless.com/. Boundless Learning CC BY-SA 3.0.
    • "hypotonic." http://en.wiktionary.org/wiki/hypotonic. Wiktionary CC BY-SA 3.0.
    • "tonicity." http://en.wiktionary.org/wiki/tonicity. Wiktionary CC BY-SA.
    • "osmoregulation." http://en.wiktionary.org/wiki/osmoregulation. Wiktionary CC BY-SA 3.0.
    • "OpenStax College, Biology. October 16, 2013." http://cnx.org/content/m44417/latest/?collection=col11448/latest. OpenStax CNX CC BY 3.0.
    • "hypertonic." http://en.wiktionary.org/wiki/hypertonic. Wiktionary CC BY-SA 3.0.
    • "OpenStax College, Passive Transport. October 16, 2013." http://cnx.org/content/m44417/latest/Figure_05_02_10.jpg. OpenStax CNX CC BY 3.0.
    • "OpenStax College, Passive Transport. October 16, 2013." http://cnx.org/content/m44417/latest/Figure_05_02_08.jpg. OpenStax CNX CC BY 3.0.
    • "OpenStax College, Passive Transport. October 16, 2013." http://cnx.org/content/m44417/latest/Figure_05_02_09.jpg. OpenStax CNX CC BY 3.0.

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