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  1. Preface
  2. Unit 1
    1. 1 The Study of Life
      1. Introduction
      2. 1.1 The Science of Biology
      3. 1.2 Themes and Concepts of Biology
      4. Key Terms
      5. Chapter Summary
      6. Review Questions
      7. Critical Thinking Questions
      8. Test Prep for AP® Courses
    2. 2 The Chemical Foundation of Life
      1. Introduction
      2. 2.1 Atoms, Isotopes, Ions, and Molecules: The Building Blocks
      3. 2.2 Water
      4. 2.3 Carbon
      5. Key Terms
      6. Chapter Summary
      7. Review Questions
      8. Critical Thinking Questions
      9. Test Prep for AP® Courses
      10. Science Practice Challenge Questions
    3. 3 Biological Macromolecules
      1. Introduction
      2. 3.1 Synthesis of Biological Macromolecules
      3. 3.2 Carbohydrates
      4. 3.3 Lipids
      5. 3.4 Proteins
      6. 3.5 Nucleic Acids
      7. Key Terms
      8. Chapter Summary
      9. Review Questions
      10. Critical Thinking Questions
      11. Test Prep for AP® Courses
      12. Science Practice Challenge Questions
  3. Unit 2
    1. 4 Cell Structure
      1. Introduction
      2. 4.1 Studying Cells
      3. 4.2 Prokaryotic Cells
      4. 4.3 Eukaryotic Cells
      5. 4.4 The Endomembrane System and Proteins
      6. 4.5 Cytoskeleton
      7. 4.6 Connections between Cells and Cellular Activities
      8. Key Terms
      9. Chapter Summary
      10. Review Questions
      11. Critical Thinking Questions
      12. Test Prep for AP® Courses
      13. Science Practice Challenge Questions
    2. 5 Structure and Function of Plasma Membranes
      1. Introduction
      2. 5.1 Components and Structure
      3. 5.2 Passive Transport
      4. 5.3 Active Transport
      5. 5.4 Bulk Transport
      6. Key Terms
      7. Chapter Summary
      8. Review Questions
      9. Critical Thinking Questions
      10. Test Prep for AP® Courses
      11. Science Practice Challenge Questions
    3. 6 Metabolism
      1. Introduction
      2. 6.1 Energy and Metabolism
      3. 6.2 Potential, Kinetic, Free, and Activation Energy
      4. 6.3 The Laws of Thermodynamics
      5. 6.4 ATP: Adenosine Triphosphate
      6. 6.5 Enzymes
      7. Key Terms
      8. Chapter Summary
      9. Review Questions
      10. Critical Thinking Questions
      11. Test Prep for AP® Courses
      12. Science Practice Challenge Questions
    4. 7 Cellular Respiration
      1. Introduction
      2. 7.1 Energy in Living Systems
      3. 7.2 Glycolysis
      4. 7.3 Oxidation of Pyruvate and the Citric Acid Cycle
      5. 7.4 Oxidative Phosphorylation
      6. 7.5 Metabolism without Oxygen
      7. 7.6 Connections of Carbohydrate, Protein, and Lipid Metabolic Pathways
      8. 7.7 Regulation of Cellular Respiration
      9. Key Terms
      10. Chapter Summary
      11. Review Questions
      12. Critical Thinking Questions
      13. Test Prep for AP® Courses
      14. Science Practice Challenge Questions
    5. 8 Photosynthesis
      1. Introduction
      2. 8.1 Overview of Photosynthesis
      3. 8.2 The Light-Dependent Reaction of Photosynthesis
      4. 8.3 Using Light to Make Organic Molecules
      5. Key Terms
      6. Chapter Summary
      7. Review Questions
      8. Critical Thinking Questions
      9. Test Prep for AP® Courses
      10. Science Practice Challenge Questions
    6. 9 Cell Communication
      1. Introduction
      2. 9.1 Signaling Molecules and Cellular Receptors
      3. 9.2 Propagation of the Signal
      4. 9.3 Response to the Signal
      5. 9.4 Signaling in Single-Celled Organisms
      6. Key Terms
      7. Chapter Summary
      8. Review Questions
      9. Critical Thinking Questions
      10. Test Prep for AP® Courses
      11. Science Practice Challenge Questions
    7. 10 Cell Reproduction
      1. Introduction
      2. 10.1 Cell Division
      3. 10.2 The Cell Cycle
      4. 10.3 Control of the Cell Cycle
      5. 10.4 Cancer and the Cell Cycle
      6. 10.5 Prokaryotic Cell Division
      7. Key Terms
      8. Chapter Summary
      9. Review Questions
      10. Critical Thinking Questions
      11. Test Prep for AP® Courses
      12. Science Practice Challenge Questions
  4. Unit 3
    1. 11 Meiosis and Sexual Reproduction
      1. Introduction
      2. 11.1 The Process of Meiosis
      3. 11.2 Sexual Reproduction
      4. Key Terms
      5. Chapter Summary
      6. Review Questions
      7. Critical Thinking Questions
      8. Test Prep for AP® Courses
      9. Science Practice Challenge Questions
    2. 12 Mendel's Experiments and Heredity
      1. Introduction
      2. 12.1 Mendel’s Experiments and the Laws of Probability
      3. 12.2 Characteristics and Traits
      4. 12.3 Laws of Inheritance
      5. Key Terms
      6. Chapter Summary
      7. Review Questions
      8. Critical Thinking Questions
      9. Test Prep for AP® Courses
      10. Science Practice Challenge Questions
    3. 13 Modern Understandings of Inheritance
      1. Introduction
      2. 13.1 Chromosomal Theory and Genetic Linkages
      3. 13.2 Chromosomal Basis of Inherited Disorders
      4. Key Terms
      5. Chapter Summary
      6. Review Questions
      7. Critical Thinking Questions
      8. Test Prep for AP® Courses
      9. Science Practice Challenge Questions
    4. 14 DNA Structure and Function
      1. Introduction
      2. 14.1 Historical Basis of Modern Understanding
      3. 14.2 DNA Structure and Sequencing
      4. 14.3 Basics of DNA Replication
      5. 14.4 DNA Replication in Prokaryotes
      6. 14.5 DNA Replication in Eukaryotes
      7. 14.6 DNA Repair
      8. Key Terms
      9. Chapter Summary
      10. Review Questions
      11. Critical Thinking Questions
      12. Test Prep for AP® Courses
      13. Science Practice Challenge Questions
    5. 15 Genes and Proteins
      1. Introduction
      2. 15.1 The Genetic Code
      3. 15.2 Prokaryotic Transcription
      4. 15.3 Eukaryotic Transcription
      5. 15.4 RNA Processing in Eukaryotes
      6. 15.5 Ribosomes and Protein Synthesis
      7. Key Terms
      8. Chapter Summary
      9. Review Questions
      10. Critical Thinking Questions
      11. Test Prep for AP® Courses
      12. Science Practice Challenge Questions
    6. 16 Gene Regulation
      1. Introduction
      2. 16.1 Regulation of Gene Expression
      3. 16.2 Prokaryotic Gene Regulation
      4. 16.3 Eukaryotic Epigenetic Gene Regulation
      5. 16.4 Eukaryotic Transcriptional Gene Regulation
      6. 16.5 Eukaryotic Post-transcriptional Gene Regulation
      7. 16.6 Eukaryotic Translational and Post-translational Gene Regulation
      8. 16.7 Cancer and Gene Regulation
      9. Key Terms
      10. Chapter Summary
      11. Review Questions
      12. Critical Thinking Questions
      13. Test Prep for AP® Courses
      14. Science Practice Challenge Questions
    7. 17 Biotechnology and Genomics
      1. Introduction
      2. 17.1 Biotechnology
      3. 17.2 Mapping Genomes
      4. 17.3 Whole-Genome Sequencing
      5. 17.4 Applying Genomics
      6. 17.5 Genomics and Proteomics
      7. Key Terms
      8. Chapter Summary
      9. Review Questions
      10. Critical Thinking Questions
      11. Test Prep for AP® Courses
      12. Science Practice Challenge Questions
  5. Unit 4
    1. 18 Evolution and Origin of Species
      1. Introduction
      2. 18.1 Understanding Evolution
      3. 18.2 Formation of New Species
      4. 18.3 Reconnection and Rates of Speciation
      5. Key Terms
      6. Chapter Summary
      7. Review Questions
      8. Critical Thinking Questions
      9. Test Prep for AP® Courses
      10. Science Practice Challenge Questions
    2. 19 The Evolution of Populations
      1. Introduction
      2. 19.1 Population Evolution
      3. 19.2 Population Genetics
      4. 19.3 Adaptive Evolution
      5. Key Terms
      6. Chapter Summary
      7. Review Questions
      8. Critical Thinking Questions
      9. Test Prep for AP® Courses
      10. Science Practice Challenge Questions
    3. 20 Phylogenies and the History of Life
      1. Introduction
      2. 20.1 Organizing Life on Earth
      3. 20.2 Determining Evolutionary Relationships
      4. 20.3 Perspectives on the Phylogenetic Tree
      5. Key Terms
      6. Chapter Summary
      7. Review Questions
      8. Critical Thinking Questions
      9. Test Prep for AP® Courses
      10. Science Practice Challenge Questions
  6. Unit 5
    1. 21 Viruses
      1. Introduction
      2. 21.1 Viral Evolution, Morphology, and Classification
      3. 21.2 Virus Infection and Hosts
      4. 21.3 Prevention and Treatment of Viral Infections
      5. 21.4 Other Acellular Entities: Prions and Viroids
      6. Key Terms
      7. Chapter Summary
      8. Review Questions
      9. Critical Thinking Questions
      10. Test Prep for AP® Courses
      11. Science Practice Challenge Questions
    2. 22 Prokaryotes: Bacteria and Archaea
      1. Introduction
      2. 22.1 Prokaryotic Diversity
      3. 22.2 Structure of Prokaryotes
      4. 22.3 Prokaryotic Metabolism
      5. 22.4 Bacterial Diseases in Humans
      6. 22.5 Beneficial Prokaryotes
      7. Key Terms
      8. Chapter Summary
      9. Review Questions
      10. Critical Thinking Questions
      11. Test Prep for AP® Courses
      12. Science Practice Challenge Questions
  7. Unit 6
    1. 23 Plant Form and Physiology
      1. Introduction
      2. 23.1 The Plant Body
      3. 23.2 Stems
      4. 23.3 Roots
      5. 23.4 Leaves
      6. 23.5 Transport of Water and Solutes in Plants
      7. 23.6 Plant Sensory Systems and Responses
      8. Key Terms
      9. Chapter Summary
      10. Review Questions
      11. Critical Thinking Questions
      12. Test Prep for AP® Courses
      13. Science Practice Challenge Questions
  8. Unit 7
    1. 24 The Animal Body: Basic Form and Function
      1. Introduction
      2. 24.1 Animal Form and Function
      3. 24.2 Animal Primary Tissues
      4. 24.3 Homeostasis
      5. Key Terms
      6. Chapter Summary
      7. Review Questions
      8. Critical Thinking Questions
      9. Test Prep for AP® Courses
    2. 25 Animal Nutrition and the Digestive System
      1. Introduction
      2. 25.1 Digestive Systems
      3. 25.2 Nutrition and Energy Production
      4. 25.3 Digestive System Processes
      5. 25.4 Digestive System Regulation
      6. Key Terms
      7. Chapter Summary
      8. Review Questions
      9. Critical Thinking Questions
      10. Test Prep for AP® Courses
      11. Science Practice Challenge Questions
    3. 26 The Nervous System
      1. Introduction
      2. 26.1 Neurons and Glial Cells
      3. 26.2 How Neurons Communicate
      4. 26.3 The Central Nervous System
      5. 26.4 The Peripheral Nervous System
      6. 26.5 Nervous System Disorders
      7. Key Terms
      8. Chapter Summary
      9. Review Questions
      10. Critical Thinking Questions
      11. Test Prep for AP® Courses
      12. Science Practice Challenge Questions
    4. 27 Sensory Systems
      1. Introduction
      2. 27.1 Sensory Processes
      3. 27.2 Somatosensation
      4. 27.3 Taste and Smell
      5. 27.4 Hearing and Vestibular Sensation
      6. 27.5 Vision
      7. Key Terms
      8. Chapter Summary
      9. Review Questions
      10. Critical Thinking Questions
      11. Science Practice Challenge Questions
    5. 28 The Endocrine System
      1. Introduction
      2. 28.1 Types of Hormones
      3. 28.2 How Hormones Work
      4. 28.3 Regulation of Body Processes
      5. 28.4 Regulation of Hormone Production
      6. 28.5 Endocrine Glands
      7. Key Terms
      8. Chapter Summary
      9. Review Questions
      10. Critical Thinking Questions
      11. Test Prep for AP® Courses
      12. Science Practice Challenge Questions
    6. 29 The Musculoskeletal System
      1. Introduction
      2. 29.1 Types of Skeletal Systems
      3. 29.2 Bone
      4. 29.3 Joints and Skeletal Movement
      5. 29.4 Muscle Contraction and Locomotion
      6. Key Terms
      7. Chapter Summary
      8. Review Questions
      9. Critical Thinking Questions
      10. Science Practice Challenge Questions
    7. 30 The Respiratory System
      1. Introduction
      2. 30.1 Systems of Gas Exchange
      3. 30.2 Gas Exchange across Respiratory Surfaces
      4. 30.3 Breathing
      5. 30.4 Transport of Gases in Human Bodily Fluids
      6. Key Terms
      7. Chapter Summary
      8. Review Questions
      9. Critical Thinking Questions
      10. Test Prep for AP® Courses
      11. Science Practice Challenge Questions
    8. 31 The Circulatory System
      1. Introduction
      2. 31.1 Overview of the Circulatory System
      3. 31.2 Components of the Blood
      4. 31.3 Mammalian Heart and Blood Vessels
      5. 31.4 Blood Flow and Blood Pressure Regulation
      6. Key Terms
      7. Chapter Summary
      8. Review Questions
      9. Critical Thinking Questions
      10. Test Prep for AP® Courses
      11. Science Practice Challenge Questions
    9. 32 Osmotic Regulation and Excretion
      1. Introduction
      2. 32.1 Osmoregulation and Osmotic Balance
      3. 32.2 The Kidneys and Osmoregulatory Organs
      4. 32.3 Excretion Systems
      5. 32.4 Nitrogenous Wastes
      6. 32.5 Hormonal Control of Osmoregulatory Functions
      7. Key Terms
      8. Chapter Summary
      9. Review Questions
      10. Critical Thinking Questions
      11. Test Prep for AP® Courses
    10. 33 The Immune System
      1. Introduction
      2. 33.1 Innate Immune Response
      3. 33.2 Adaptive Immune Response
      4. 33.3 Antibodies
      5. 33.4 Disruptions in the Immune System
      6. Key Terms
      7. Chapter Summary
      8. Review Questions
      9. Critical Thinking Questions
      10. Test Prep for AP® Courses
      11. Science Practice Challenge Questions
    11. 34 Animal Reproduction and Development
      1. Introduction
      2. 34.1 Reproduction Methods
      3. 34.2 Fertilization
      4. 34.3 Human Reproductive Anatomy and Gametogenesis
      5. 34.4 Hormonal Control of Human Reproduction
      6. 34.5 Fertilization and Early Embryonic Development
      7. 34.6 Organogenesis and Vertebrate Formation
      8. 34.7 Human Pregnancy and Birth
      9. Key Terms
      10. Chapter Summary
      11. Review Questions
      12. Critical Thinking Questions
      13. Test Prep for AP® Courses
      14. Science Practice Challenge Questions
  9. Unit 8
    1. 35 Ecology and the Biosphere
      1. Introduction
      2. 35.1 The Scope of Ecology
      3. 35.2 Biogeography
      4. 35.3 Terrestrial Biomes
      5. 35.4 Aquatic Biomes
      6. 35.5 Climate and the Effects of Global Climate Change
      7. Key Terms
      8. Chapter Summary
      9. Review Questions
      10. Critical Thinking Questions
      11. Test Prep for AP® Courses
      12. Science Practice Challenge Questions
    2. 36 Population and Community Ecology
      1. Introduction
      2. 36.1 Population Demography
      3. 36.2 Life Histories and Natural Selection
      4. 36.3 Environmental Limits to Population Growth
      5. 36.4 Population Dynamics and Regulation
      6. 36.5 Human Population Growth
      7. 36.6 Community Ecology
      8. 36.7 Behavioral Biology: Proximate and Ultimate Causes of Behavior
      9. Key Terms
      10. Chapter Summary
      11. Review Questions
      12. Critical Thinking Questions
      13. Test Prep for AP® Courses
      14. Science Practice Challenge Questions
    3. 37 Ecosystems
      1. Introduction
      2. 37.1 Ecology for Ecosystems
      3. 37.2 Energy Flow through Ecosystems
      4. 37.3 Biogeochemical Cycles
      5. Key Terms
      6. Chapter Summary
      7. Review Questions
      8. Critical Thinking Questions
      9. Test Prep for AP® Courses
      10. Science Practice Challenge Questions
    4. 38 Conservation Biology and Biodiversity
      1. Introduction
      2. 38.1 The Biodiversity Crisis
      3. 38.2 The Importance of Biodiversity to Human Life
      4. 38.3 Threats to Biodiversity
      5. 38.4 Preserving Biodiversity
      6. Key Terms
      7. Chapter Summary
      8. Review Questions
      9. Critical Thinking Questions
      10. Test Prep for AP® Courses
  10. A | The Periodic Table of Elements
  11. B | Geological Time
  12. C | Measurements and the Metric System
  13. Index

In this section, you will explore the following questions:

  • What are the parts of a typical leaf?
  • What is the internal structure and function of a leaf?
  • What are differences between simple leaves and compound leaves? Similarities?
  • What are examples of modified leaves?

Connection for AP® Courses

Plants have specialized leaves adapted to their particular environments. For example, the leaves of plants growing in tropical rainforests have a larger surface area than cacti growing in the desert or in very cold conditions, whose smaller surface area minimizes water loss through transpiration. A waxy cuticle covers the surface of all plant species to reduce the rate of water loss from the leaf surface. Other leaves may have small hairs called trichomes on the leaf surface; not only do trichomes also help reduce water loss; they also protect the leaf from herbivory by restricting insect movements or by storing toxic or bad-tasting chemicals.

The information presented in this module does not align to the content and AP® Learning Objectives outlined in the AP® Curriculum Framework. The role of stomata, guard cells, and trichomes in transpiration are explored in the Transport of Water and Solutes in Plants module.

Leaves are the main sites for photosynthesis: the process by which plants synthesize food. Most leaves are usually green, due to the presence of chlorophyll in the leaf cells. However, some leaves may have different colors, caused by other plant pigments that mask the green chlorophyll.

The thickness, shape, and size of leaves are adapted to the environment. Each variation helps a plant species maximize its chances of survival in a particular habitat. Usually, the leaves of plants growing in tropical rainforests have larger surface areas than those of plants growing in deserts or very cold conditions, which are likely to have a smaller surface area to minimize water loss.

Structure of a Typical Leaf

Each leaf typically has a leaf blade called the lamina, which is also the widest part of the leaf. Some leaves are attached to the plant stem by a petiole. Leaves that do not have a petiole and are directly attached to the plant stem are called sessile leaves. Small green appendages usually found at the base of the petiole are known as stipules. Most leaves have a midrib, which travels the length of the leaf and branches to each side to produce veins of vascular tissue. The edge of the leaf is called the margin. Figure 23.21 shows the structure of a typical eudicot leaf.

 Illustration shows the parts of a leaf. The petiole is the stem of the leaf. The midrib is a vessel that extends from the petiole to the leaf tip. Veins branch from the midrib. The lamina is the wide, flat part of the leaf. The margin is the edge of the leaf.
Figure 23.21 Deceptively simple in appearance, a leaf is a highly efficient structure.

Within each leaf, the vascular tissue forms veins. The arrangement of veins in a leaf is called the venation pattern. Monocots and dicots differ in their patterns of venation (Figure 23.22). Monocots have parallel venation; the veins run in straight lines across the length of the leaf without converging at a point. In dicots, however, the veins of the leaf have a net-like appearance, forming a pattern known as reticulate venation. One extant plant, the Ginkgo biloba, has dichotomous venation where the veins fork.

 Part A photo shows the broad, sword-shaped leaves of a tulip. Parallel veins run up the leaves. Part B photo shows a teardrop-shaped linden leaf that has veins radiating out from the midrib. Smaller veins radiate out from these. Right photo shows a fan-shaped ginkgo leaf, which has veins radiating out from the petiole.
Figure 23.22 (a) Tulip (Tulipa), a monocot, has leaves with parallel venation. The netlike venation in this (b) linden (Tilia cordata) leaf distinguishes it as a dicot. The (c) Ginkgo biloba tree has dichotomous venation. (credit a photo: modification of work by “Drewboy64”/Wikimedia Commons; credit b photo: modification of work by Roger Griffith; credit c photo: modification of work by "geishaboy500"/Flickr; credit abc illustrations: modification of work by Agnieszka Kwiecień)

Leaf Arrangement

The arrangement of leaves on a stem is known as phyllotaxy. The number and placement of a plant’s leaves will vary depending on the species, with each species exhibiting a characteristic leaf arrangement. Leaves are classified as either alternate, spiral, or opposite. Plants that have only one leaf per node have leaves that are said to be either alternate—meaning the leaves alternate on each side of the stem in a flat plane—or spiral, meaning the leaves are arrayed in a spiral along the stem. In an opposite leaf arrangement, two leaves arise at the same point, with the leaves connecting opposite each other along the branch. If there are three or more leaves connected at a node, the leaf arrangement is classified as whorled.

Leaf Form

Leaves may be simple or compound (Figure 23.23). In a simple leaf, the blade is either completely undivided—as in the banana leaf—or it has lobes, but the separation does not reach the midrib, as in the maple leaf. In a compound leaf, the leaf blade is completely divided, forming leaflets, as in the locust tree. Each leaflet may have its own stalk, but is attached to the rachis. A palmately compound leaf resembles the palm of a hand, with leaflets radiating outwards from one point Examples include the leaves of poison ivy, the buckeye tree, or the familiar houseplant Schefflera sp. (common name “umbrella plant”). Pinnately compound leaves take their name from their feather-like appearance; the leaflets are arranged along the midrib, as in rose leaves (Rosa sp.), or the leaves of hickory, pecan, ash, or walnut trees.

Photo (a) shows the large-leaves of a potted banana plant growing from a single stem; (b) shows a horse chestnut plant, which has five leaves radiating from the petiole as fingers radiate from the palm of a hand; (c) shows a scrub hickory plant with feather-shaped leaves opposing each other along the stem, and a single leaf at the end of the stem. (d) shows a honey locust with five pairs of stem-like veins connected to the midrib. Tiny leaflets grow from the veins.
Figure 23.23 Leaves may be simple or compound. In simple leaves, the lamina is continuous. The (a) banana plant (Musa sp.) has simple leaves. In compound leaves, the lamina is separated into leaflets. Compound leaves may be palmate or pinnate. In (b) palmately compound leaves, such as those of the horse chestnut (Aesculus hippocastanum), the leaflets branch from the petiole. In (c) pinnately compound leaves, the leaflets branch from the midrib, as on a scrub hickory (Carya floridana). The (d) honey locust has double compound leaves, in which leaflets branch from the veins. (credit a: modification of work by "BazzaDaRambler"/Flickr; credit b: modification of work by Roberto Verzo; credit c: modification of work by Eric Dion; credit d: modification of work by Valerie Lykes)

Leaf Structure and Function

The outermost layer of the leaf is the epidermis; it is present on both sides of the leaf and is called the upper and lower epidermis, respectively. Botanists call the upper side the adaxial surface (or adaxis) and the lower side the abaxial surface (or abaxis). The epidermis helps in the regulation of gas exchange. It contains stomata (Figure 23.24): openings through which the exchange of gases takes place. Two guard cells surround each stoma, regulating its opening and closing.

Photo (a) shows small oval-like stomata scattered on the bumpy surface of a leaf that is magnified 500 times; (b) is a close-up of a stoma showing the thick lip-like guard cells either side of an opening. Photo (a) and (b) are scanning electron micrographs. Photo (c) is a light micrograph of a leaf cross section that shows a large air space underneath two guard cells. The air space is surrounded by large oval and egg-shaped cells.
Figure 23.24 Visualized at 500x with a scanning electron microscope, several stomata are clearly visible on (a) the surface of this sumac (Rhus glabra) leaf. At 5,000x magnification, the guard cells of (b) a single stoma from lyre-leaved sand cress (Arabidopsis lyrata) have the appearance of lips that surround the opening. In this (c) light micrograph cross-section of an A. lyrata leaf, the guard cell pair is visible along with the large, sub-stomatal air space in the leaf. (credit: modification of work by Robert R. Wise; part c scale-bar data from Matt Russell)

The epidermis is usually one cell layer thick; however, in plants that grow in very hot or very cold conditions, the epidermis may be several layers thick to protect against excessive water loss from transpiration. A waxy layer known as the cuticle covers the leaves of all plant species. The cuticle reduces the rate of water loss from the leaf surface. Other leaves may have small hairs (trichomes) on the leaf surface. Trichomes help to deter herbivory by restricting insect movements, or by storing toxic or bad-tasting compounds; they can also reduce the rate of transpiration by blocking air flow across the leaf surface (Figure 23.25).

Photo (a) shows a plant with many fuzzy white hairs growing from its surface. Scanning electron micrograph (b) shows branched tree-like hairs emerging from the surface of a leaf. The trunk of each hair is about 250 microns tall. Branches are somewhat shorter. Scanning electron micrograph (c) shows many multi-pronged hairs about 100 microns long that look like sea anemones scattered across a leaf surface.
Figure 23.25 Trichomes give leaves a fuzzy appearance as in this (a) sundew (Drosera sp.). Leaf trichomes include (b) branched trichomes on the leaf of Arabidopsis lyrata and (c) multibranched trichomes on a mature Quercus marilandica leaf. (credit a: John Freeland; credit b, c: modification of work by Robert R. Wise; scale-bar data from Matt Russell)

Below the epidermis of dicot leaves are layers of cells known as the mesophyll, or “middle leaf.” The mesophyll of most leaves typically contains two arrangements of parenchyma cells: the palisade parenchyma and spongy parenchyma (Figure 23.26). The palisade parenchyma (also called the palisade mesophyll) has column-shaped, tightly packed cells, and may be present in one, two, or three layers. Below the palisade parenchyma are loosely arranged cells of an irregular shape. These are the cells of the spongy parenchyma (or spongy mesophyll). The air space found between the spongy parenchyma cells allows gaseous exchange between the leaf and the outside atmosphere through the stomata. In aquatic plants, the intercellular spaces in the spongy parenchyma help the leaf float. Both layers of the mesophyll contain many chloroplasts. Guard cells are the only epidermal cells to contain chloroplasts.

 Part A is a leaf cross section illustration. A flat layer of rectangular cells make up the upper and lower epidermis. A cuticle layer protects the outside of both epidermal layers. A stomatal pore in the lower epidermis allows carbon dioxide to enter and oxygen to leave. Oval guard cells surround the pore. Sandwiched between the upper and lower epidermis is the mesophyll. The upper part of the mesophyll is comprised of columnar cells called palisade parenchyma. The lower part of the mesophyll is made up of loosely packed spongy parenchyma. Part B is a scanning electron micrograph of a leaf in which all the layers described above are visible. Palisade cells are about 50 microns tall and 10 microns wide and are covered with tiny bumps, which are the chloroplasts. Spongy cells smaller and irregularly shaped. Several large bumps about 20 microns across project from the lower surface of the leaf.
Figure 23.26 In the (a) leaf drawing, the central mesophyll is sandwiched between an upper and lower epidermis. The mesophyll has two layers: an upper palisade layer comprised of tightly packed, columnar cells, and a lower spongy layer, comprised of loosely packed, irregularly shaped cells. Stomata on the leaf underside allow gas exchange. A waxy cuticle covers all aerial surfaces of land plants to minimize water loss. These leaf layers are clearly visible in the (b) scanning electron micrograph. The numerous small bumps in the palisade parenchyma cells are chloroplasts. Chloroplasts are also present in the spongy parenchyma, but are not as obvious. The bumps protruding from the lower surface of the leave are glandular trichomes, which differ in structure from the stalked trichomes in Figure 23.25. (credit b: modification of work by Robert R. Wise)

Like the stem, the leaf contains vascular bundles composed of xylem and phloem (Figure 23.27). The xylem consists of tracheids and vessels, which transport water and minerals to the leaves. The phloem transports the photosynthetic products from the leaf to the other parts of the plant. A single vascular bundle, no matter how large or small, always contains both xylem and phloem tissues.

 The scanning electron micrograph shows an oval vascular bundle. Small phloem cells make up the bottom of the bundle, and larger xylem cells make up the top. The bundle is surrounded by a ring of larger cells.
Figure 23.27 This scanning electron micrograph shows xylem and phloem in the leaf vascular bundle from the lyre-leaved sand cress (Arabidopsis lyrata). (credit: modification of work by Robert R. Wise; scale-bar data from Matt Russell)

Leaf Adaptations

Coniferous plant species that thrive in cold environments, like spruce, fir, and pine, have leaves that are reduced in size and needle-like in appearance. These needle-like leaves have sunken stomata and a smaller surface area: two attributes that aid in reducing water loss. In hot climates, plants such as cacti have leaves that are reduced to spines, which in combination with their succulent stems, help to conserve water. Many aquatic plants have leaves with wide lamina that can float on the surface of the water, and a thick waxy cuticle on the leaf surface that repels water.

Link to Learning

Watch “The Pale Pitcher Plant” episode of the video series Plants Are Cool, Too, a Botanical Society of America video about a carnivorous plant species found in Louisiana.

How do pale pitcher plants (Sarracinia alata) make sure that insects do not escape after consuming the nectar that attracted them?
  1. The insects are skewered on spikes and thorns that rim the funnel-shaped leaf.
  2. The insects ingest narcotics secreted by the leaf and fall into the funnel-shaped leaf.
  3. The insects are poisoned by lethal compounds in the nectar.
  4. The insects are immobilized by sticky substances on the rim of the funnel-shaped leaf.

Evolution Connection

Plant Adaptations in Resource-Deficient Environments

Roots, stems, and leaves are structured to ensure that a plant can obtain the required sunlight, water, soil nutrients, and oxygen resources. Some remarkable adaptations have evolved to enable plant species to thrive in less than ideal habitats, where one or more of these resources is in short supply.

In tropical rainforests, light is often scarce, since many trees and plants grow close together and block much of the sunlight from reaching the forest floor. Many tropical plant species have exceptionally broad leaves to maximize the capture of sunlight. Other species are epiphytes: plants that grow on other plants that serve as a physical support. Such plants are able to grow high up in the canopy atop the branches of other trees, where sunlight is more plentiful. Epiphytes live on rain and minerals collected in the branches and leaves of the supporting plant. Bromeliads (members of the pineapple family), ferns, and orchids are examples of tropical epiphytes (Figure 23.28). Many epiphytes have specialized tissues that enable them to efficiently capture and store water.

Photo shows long, thin brown leaves of Spanish moss hanging down from the branches of a large oak tree.
Figure 23.28 One of the most well known bromeliads is Spanish moss (Tillandsia usneoides), seen here in an oak tree. (credit: Kristine Paulus)

Some plants have special adaptations that help them to survive in nutrient-poor environments. Carnivorous plants, such as the Venus flytrap and the pitcher plant (Figure 23.29), grow in bogs where the soil is low in nitrogen. In these plants, leaves are modified to capture insects. The insect-capturing leaves may have evolved to provide these plants with a supplementary source of much-needed nitrogen.

 Left photo shows modified leaves of a Venus flytrap. The two leaves resemble the upper and lower part of the mouth, and are red on the interior. Hair-like appendages, like teeth, frame each modified leaf, so that when the leaves close, the insect will be trapped. Right photo shows three modified leaves of the pitcher plant, which are green tubes with red specks and have a red rim forming the top opening.
Figure 23.29 The (a) Venus flytrap has modified leaves that can capture insects. When an unlucky insect touches the trigger hairs inside the leaf, the trap suddenly closes. The opening of the (b) pitcher plant is lined with a slippery wax. Insects crawling on the lip slip and fall into a pool of water in the bottom of the pitcher, where they are digested by bacteria. The plant then absorbs the smaller molecules. (credit a: modification of work by Peter Shanks; credit b: modification of work by Tim Mansfield)

Many swamp plants have adaptations that enable them to thrive in wet areas, where their roots grow submerged underwater. In these aquatic areas, the soil is unstable and little oxygen is available to reach the roots. Trees such as mangroves (Rhizophora sp.) growing in coastal waters produce aboveground roots that help support the tree (Figure 23.30). Some species of mangroves, as well as cypress trees, have pneumatophores: upward-growing roots containing pores and pockets of tissue specialized for gas exchange. Wild rice is an aquatic plant with large air spaces in the root cortex. The air-filled tissue—called aerenchyma—provides a path for oxygen to diffuse down to the root tips, which are embedded in oxygen-poor bottom sediments.

How do pale pitcher plants (Sarracinia alata) make sure that insects do not escape after consuming the nectar that attracted them?
  1. The insects are skewered on spikes and thorns that rim the funnel-shaped leaf.
  2. The insects ingest narcotics secreted by the leaf and fall into the funnel-shaped leaf.
  3. The insects are poisoned by lethal compounds in the nectar.
  4. The insects are immobilized by sticky substances on the rim of the funnel-shaped leaf.
 Photo A shows mangrove trees with roots extending into the water. Part B shows cypress trees growing in the water, with upward-growing roots between the trees. Part C is a scanning electron micrograph showing a cross section of wild rice. The cells radiate from the center like spokes on a bicycle wheel, and are interspersed by large spaces that hold air.
Figure 23.30 The branches of (a) mangrove trees develop aerial roots, which descend to the ground and help to anchor the trees. (b) Cypress trees and some mangrove species have upward-growing roots called pneumatophores that are involved in gas exchange. Aquatic plants such as (c) wild rice have large spaces in the root cortex called aerenchyma, visualized here using scanning electron microscopy. (credit a: modification of work by Roberto Verzo; credit b: modification of work by Duane Burdick; credit c: modification of work by Robert R. Wise)

Link to Learning

Watch Venus Flytraps: Jaws of Death, an extraordinary BBC close-up of the Venus flytrap in action.

Why do many ornamental plants that thrive indoors originate on the floor of tropical rainforest, where they grow under the canopy of trees?
  1. Growing under the rainforest’s canopy made these plants adapt to less water and nutrients.
  2. With their narrow leaves, these plants are adapted to grow in low light.
  3. With their broad leaves, these plants are adapted to grow in low light, like that usually found indoors.
  4. Growing under the rainforest’s canopy provides the plants with more water and nutrients, which they also need as indoor plants.
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