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Biology 2e

28.5 Superphylum Ecdysozoa: Nematodes and Tardigrades

Biology 2e28.5 Superphylum Ecdysozoa: Nematodes and Tardigrades
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  1. Preface
  2. The Chemistry of Life
    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. Visual Connection Questions
      7. Review Questions
      8. Critical Thinking Questions
    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. Visual Connection Questions
      8. Review Questions
      9. Critical Thinking 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. Visual Connection Questions
      10. Review Questions
      11. Critical Thinking Questions
  3. The Cell
    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 The Cytoskeleton
      7. 4.6 Connections between Cells and Cellular Activities
      8. Key Terms
      9. Chapter Summary
      10. Visual Connection Questions
      11. Review Questions
      12. Critical Thinking 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. Visual Connection Questions
      9. Review Questions
      10. Critical Thinking 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. Visual Connection Questions
      10. Review Questions
      11. Critical Thinking 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. Visual Connection Questions
      12. Review Questions
      13. Critical Thinking Questions
    5. 8 Photosynthesis
      1. Introduction
      2. 8.1 Overview of Photosynthesis
      3. 8.2 The Light-Dependent Reactions of Photosynthesis
      4. 8.3 Using Light Energy to Make Organic Molecules
      5. Key Terms
      6. Chapter Summary
      7. Visual Connection Questions
      8. Review Questions
      9. Critical Thinking 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. Visual Connection Questions
      9. Review Questions
      10. Critical Thinking 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. Visual Connection Questions
      10. Review Questions
      11. Critical Thinking Questions
  4. Genetics
    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. Visual Connection Questions
      7. Review Questions
      8. Critical Thinking 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. Visual Connection Questions
      8. Review Questions
      9. Critical Thinking Questions
    3. 13 Modern Understandings of Inheritance
      1. Introduction
      2. 13.1 Chromosomal Theory and Genetic Linkage
      3. 13.2 Chromosomal Basis of Inherited Disorders
      4. Key Terms
      5. Chapter Summary
      6. Visual Connection Questions
      7. Review Questions
      8. Critical Thinking 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. Visual Connection Questions
      11. Review Questions
      12. Critical Thinking 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. Visual Connection Questions
      10. Review Questions
      11. Critical Thinking Questions
    6. 16 Gene Expression
      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 Transcription 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. Visual Connection Questions
      12. Review Questions
      13. Critical Thinking 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. Visual Connection Questions
      10. Review Questions
      11. Critical Thinking Questions
  5. Evolutionary Processes
    1. 18 Evolution and the Origin of Species
      1. Introduction
      2. 18.1 Understanding Evolution
      3. 18.2 Formation of New Species
      4. 18.3 Reconnection and Speciation Rates
      5. Key Terms
      6. Chapter Summary
      7. Visual Connection Questions
      8. Review Questions
      9. Critical Thinking 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. Visual Connection Questions
      8. Review Questions
      9. Critical Thinking 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. Visual Connection Questions
      8. Review Questions
      9. Critical Thinking Questions
  6. Biological Diversity
    1. 21 Viruses
      1. Introduction
      2. 21.1 Viral Evolution, Morphology, and Classification
      3. 21.2 Virus Infections 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. Visual Connection Questions
      9. Review Questions
      10. Critical Thinking Questions
    2. 22 Prokaryotes: Bacteria and Archaea
      1. Introduction
      2. 22.1 Prokaryotic Diversity
      3. 22.2 Structure of Prokaryotes: Bacteria and Archaea
      4. 22.3 Prokaryotic Metabolism
      5. 22.4 Bacterial Diseases in Humans
      6. 22.5 Beneficial Prokaryotes
      7. Key Terms
      8. Chapter Summary
      9. Visual Connection Questions
      10. Review Questions
      11. Critical Thinking Questions
    3. 23 Protists
      1. Introduction
      2. 23.1 Eukaryotic Origins
      3. 23.2 Characteristics of Protists
      4. 23.3 Groups of Protists
      5. 23.4 Ecology of Protists
      6. Key Terms
      7. Chapter Summary
      8. Visual Connection Questions
      9. Review Questions
      10. Critical Thinking Questions
    4. 24 Fungi
      1. Introduction
      2. 24.1 Characteristics of Fungi
      3. 24.2 Classifications of Fungi
      4. 24.3 Ecology of Fungi
      5. 24.4 Fungal Parasites and Pathogens
      6. 24.5 Importance of Fungi in Human Life
      7. Key Terms
      8. Chapter Summary
      9. Visual Connection Questions
      10. Review Questions
      11. Critical Thinking Questions
    5. 25 Seedless Plants
      1. Introduction
      2. 25.1 Early Plant Life
      3. 25.2 Green Algae: Precursors of Land Plants
      4. 25.3 Bryophytes
      5. 25.4 Seedless Vascular Plants
      6. Key Terms
      7. Chapter Summary
      8. Visual Connection Questions
      9. Review Questions
      10. Critical Thinking Questions
    6. 26 Seed Plants
      1. Introduction
      2. 26.1 Evolution of Seed Plants
      3. 26.2 Gymnosperms
      4. 26.3 Angiosperms
      5. 26.4 The Role of Seed Plants
      6. Key Terms
      7. Chapter Summary
      8. Visual Connection Questions
      9. Review Questions
      10. Critical Thinking Questions
    7. 27 Introduction to Animal Diversity
      1. Introduction
      2. 27.1 Features of the Animal Kingdom
      3. 27.2 Features Used to Classify Animals
      4. 27.3 Animal Phylogeny
      5. 27.4 The Evolutionary History of the Animal Kingdom
      6. Key Terms
      7. Chapter Summary
      8. Visual Connection Questions
      9. Review Questions
      10. Critical Thinking Questions
    8. 28 Invertebrates
      1. Introduction
      2. 28.1 Phylum Porifera
      3. 28.2 Phylum Cnidaria
      4. 28.3 Superphylum Lophotrochozoa: Flatworms, Rotifers, and Nemerteans
      5. 28.4 Superphylum Lophotrochozoa: Molluscs and Annelids
      6. 28.5 Superphylum Ecdysozoa: Nematodes and Tardigrades
      7. 28.6 Superphylum Ecdysozoa: Arthropods
      8. 28.7 Superphylum Deuterostomia
      9. Key Terms
      10. Chapter Summary
      11. Visual Connection Questions
      12. Review Questions
      13. Critical Thinking Questions
    9. 29 Vertebrates
      1. Introduction
      2. 29.1 Chordates
      3. 29.2 Fishes
      4. 29.3 Amphibians
      5. 29.4 Reptiles
      6. 29.5 Birds
      7. 29.6 Mammals
      8. 29.7 The Evolution of Primates
      9. Key Terms
      10. Chapter Summary
      11. Visual Connection Questions
      12. Review Questions
      13. Critical Thinking Questions
  7. Plant Structure and Function
    1. 30 Plant Form and Physiology
      1. Introduction
      2. 30.1 The Plant Body
      3. 30.2 Stems
      4. 30.3 Roots
      5. 30.4 Leaves
      6. 30.5 Transport of Water and Solutes in Plants
      7. 30.6 Plant Sensory Systems and Responses
      8. Key Terms
      9. Chapter Summary
      10. Visual Connection Questions
      11. Review Questions
      12. Critical Thinking Questions
    2. 31 Soil and Plant Nutrition
      1. Introduction
      2. 31.1 Nutritional Requirements of Plants
      3. 31.2 The Soil
      4. 31.3 Nutritional Adaptations of Plants
      5. Key Terms
      6. Chapter Summary
      7. Visual Connection Questions
      8. Review Questions
      9. Critical Thinking Questions
    3. 32 Plant Reproduction
      1. Introduction
      2. 32.1 Reproductive Development and Structure
      3. 32.2 Pollination and Fertilization
      4. 32.3 Asexual Reproduction
      5. Key Terms
      6. Chapter Summary
      7. Visual Connection Questions
      8. Review Questions
      9. Critical Thinking Questions
  8. Animal Structure and Function
    1. 33 The Animal Body: Basic Form and Function
      1. Introduction
      2. 33.1 Animal Form and Function
      3. 33.2 Animal Primary Tissues
      4. 33.3 Homeostasis
      5. Key Terms
      6. Chapter Summary
      7. Visual Connection Questions
      8. Review Questions
      9. Critical Thinking Questions
    2. 34 Animal Nutrition and the Digestive System
      1. Introduction
      2. 34.1 Digestive Systems
      3. 34.2 Nutrition and Energy Production
      4. 34.3 Digestive System Processes
      5. 34.4 Digestive System Regulation
      6. Key Terms
      7. Chapter Summary
      8. Visual Connection Questions
      9. Review Questions
      10. Critical Thinking Questions
    3. 35 The Nervous System
      1. Introduction
      2. 35.1 Neurons and Glial Cells
      3. 35.2 How Neurons Communicate
      4. 35.3 The Central Nervous System
      5. 35.4 The Peripheral Nervous System
      6. 35.5 Nervous System Disorders
      7. Key Terms
      8. Chapter Summary
      9. Visual Connection Questions
      10. Review Questions
      11. Critical Thinking Questions
    4. 36 Sensory Systems
      1. Introduction
      2. 36.1 Sensory Processes
      3. 36.2 Somatosensation
      4. 36.3 Taste and Smell
      5. 36.4 Hearing and Vestibular Sensation
      6. 36.5 Vision
      7. Key Terms
      8. Chapter Summary
      9. Visual Connection Questions
      10. Review Questions
      11. Critical Thinking Questions
    5. 37 The Endocrine System
      1. Introduction
      2. 37.1 Types of Hormones
      3. 37.2 How Hormones Work
      4. 37.3 Regulation of Body Processes
      5. 37.4 Regulation of Hormone Production
      6. 37.5 Endocrine Glands
      7. Key Terms
      8. Chapter Summary
      9. Visual Connection Questions
      10. Review Questions
      11. Critical Thinking Questions
    6. 38 The Musculoskeletal System
      1. Introduction
      2. 38.1 Types of Skeletal Systems
      3. 38.2 Bone
      4. 38.3 Joints and Skeletal Movement
      5. 38.4 Muscle Contraction and Locomotion
      6. Key Terms
      7. Chapter Summary
      8. Visual Connection Questions
      9. Review Questions
      10. Critical Thinking Questions
    7. 39 The Respiratory System
      1. Introduction
      2. 39.1 Systems of Gas Exchange
      3. 39.2 Gas Exchange across Respiratory Surfaces
      4. 39.3 Breathing
      5. 39.4 Transport of Gases in Human Bodily Fluids
      6. Key Terms
      7. Chapter Summary
      8. Visual Connection Questions
      9. Review Questions
      10. Critical Thinking Questions
    8. 40 The Circulatory System
      1. Introduction
      2. 40.1 Overview of the Circulatory System
      3. 40.2 Components of the Blood
      4. 40.3 Mammalian Heart and Blood Vessels
      5. 40.4 Blood Flow and Blood Pressure Regulation
      6. Key Terms
      7. Chapter Summary
      8. Visual Connection Questions
      9. Review Questions
      10. Critical Thinking Questions
    9. 41 Osmotic Regulation and Excretion
      1. Introduction
      2. 41.1 Osmoregulation and Osmotic Balance
      3. 41.2 The Kidneys and Osmoregulatory Organs
      4. 41.3 Excretion Systems
      5. 41.4 Nitrogenous Wastes
      6. 41.5 Hormonal Control of Osmoregulatory Functions
      7. Key Terms
      8. Chapter Summary
      9. Visual Connection Questions
      10. Review Questions
      11. Critical Thinking Questions
    10. 42 The Immune System
      1. Introduction
      2. 42.1 Innate Immune Response
      3. 42.2 Adaptive Immune Response
      4. 42.3 Antibodies
      5. 42.4 Disruptions in the Immune System
      6. Key Terms
      7. Chapter Summary
      8. Visual Connection Questions
      9. Review Questions
      10. Critical Thinking Questions
    11. 43 Animal Reproduction and Development
      1. Introduction
      2. 43.1 Reproduction Methods
      3. 43.2 Fertilization
      4. 43.3 Human Reproductive Anatomy and Gametogenesis
      5. 43.4 Hormonal Control of Human Reproduction
      6. 43.5 Human Pregnancy and Birth
      7. 43.6 Fertilization and Early Embryonic Development
      8. 43.7 Organogenesis and Vertebrate Formation
      9. Key Terms
      10. Chapter Summary
      11. Visual Connection Questions
      12. Review Questions
      13. Critical Thinking Questions
  9. Ecology
    1. 44 Ecology and the Biosphere
      1. Introduction
      2. 44.1 The Scope of Ecology
      3. 44.2 Biogeography
      4. 44.3 Terrestrial Biomes
      5. 44.4 Aquatic Biomes
      6. 44.5 Climate and the Effects of Global Climate Change
      7. Key Terms
      8. Chapter Summary
      9. Visual Connection Questions
      10. Review Questions
      11. Critical Thinking Questions
    2. 45 Population and Community Ecology
      1. Introduction
      2. 45.1 Population Demography
      3. 45.2 Life Histories and Natural Selection
      4. 45.3 Environmental Limits to Population Growth
      5. 45.4 Population Dynamics and Regulation
      6. 45.5 Human Population Growth
      7. 45.6 Community Ecology
      8. 45.7 Behavioral Biology: Proximate and Ultimate Causes of Behavior
      9. Key Terms
      10. Chapter Summary
      11. Visual Connection Questions
      12. Review Questions
      13. Critical Thinking Questions
    3. 46 Ecosystems
      1. Introduction
      2. 46.1 Ecology of Ecosystems
      3. 46.2 Energy Flow through Ecosystems
      4. 46.3 Biogeochemical Cycles
      5. Key Terms
      6. Chapter Summary
      7. Visual Connection Questions
      8. Review Questions
      9. Critical Thinking Questions
    4. 47 Conservation Biology and Biodiversity
      1. Introduction
      2. 47.1 The Biodiversity Crisis
      3. 47.2 The Importance of Biodiversity to Human Life
      4. 47.3 Threats to Biodiversity
      5. 47.4 Preserving Biodiversity
      6. Key Terms
      7. Chapter Summary
      8. Visual Connection Questions
      9. Review Questions
      10. Critical Thinking Questions
  10. A | The Periodic Table of Elements
  11. B | Geological Time
  12. C | Measurements and the Metric System
  13. Index
By the end of this section, you will be able to do the following:
  • Describe the structural organization of nematodes
  • Describe the importance of Caenorhabditis elegans in research
  • Describe the features of Tardigrades

Superphylum Ecdysozoa

The superphylum Ecdysozoa contains an incredibly large number of species. This is because it contains two of the most diverse animal groups: phylum Nematoda (the roundworms) and phylum Arthropoda (the arthropods). The most prominent distinguishing feature of ecdysozoans is the cuticle—a tough, but flexible exoskeleton that protects these animals from water loss, predators, and other dangers of the external environment. One small phylum within the Ecdysozoa, with exceptional resistance to desiccation and other environmental hazards, is the Tardigrada. The nematodes, tardigrades, and arthropods all belong to the superphylum Ecdysozoa, which is believed to be monophyletic—a clade consisting of all evolutionary descendants from one common ancestor. All members of this superphylum periodically go through a molting process that culminates in ecdysis—the actual shedding of the old exoskeleton. (The term “ecdysis” translates roughly as “take off” or “strip.”) During the molting process, old cuticle is replaced by a new cuticle, which is secreted beneath it, and which will last until the next growth period.

Phylum Nematoda

The Nematoda, like other members of the superphylum Ecdysozoa, are triploblastic and possess an embryonic mesoderm that is sandwiched between the ectoderm and endoderm. They are also bilaterally symmetrical, meaning that a longitudinal section will divide them into right and left sides that are superficially symmetrical. In contrast with flatworms, nematodes are pseudocoelomates and show a tubular morphology and circular cross-section. Nematodes include both free-living and parasitic forms.

In 1914, N.A. Cobb said, “In short, if all the matter in the universe except the nematodes were swept away, our world would still be dimly recognizable, and if, as disembodied spirits, we could then investigate it, we should find its mountains, hills, vales, rivers, lakes and oceans represented by a thin film of nematodes...” To paraphrase Cobb, nematodes are so abundant that if all the non-nematode matter of the biosphere were removed, there would still remain a shadow of the former world outlined by nematodes!1 The phylum Nematoda includes more than 28,000 species with an estimated 16,000 being parasitic in nature. However, nematologists believe there may be over one million unclassified species.

The name Nematoda is derived from the Greek word “Nemos,” which means “thread,” and includes all true roundworms. Nematodes are present in all habitats, typically with each species occurring in great abundance. The free-living nematode, Caenorhabditis elegans, has been extensively used as a model system for many different avenues of biological inquiry in laboratories all over the world.

Morphology

The cylindrical body form of the nematodes is seen in Figure 28.32. These animals have a complete digestive system with a distinct mouth and anus, whereas only one opening is present in the digestive tract of flatworms. The mouth opens into a muscular pharynx and intestine, which leads to a rectum and anal opening at the posterior end. The epidermis can be either a single layer of cells or a syncytium—a multinucleated tissue that in this case is formed by the fusion of many single cells. The cuticle of nematodes is rich in collagen and a polymer called chitin, which forms a protective armor outside the epidermis. The cuticle extends into both ends of the digestive tract, the pharynx, and rectum. In the head, an anterior mouth opening is composed of three (or six) “lips” as well as teeth derived from the cuticle (in some species). Some nematodes may present other modifications of the cuticle such as rings, head shields, or warts. These external rings, however, do not reflect true internal body segmentation, which as we have seen is a hallmark of phylum Annelida. The attachment of the muscles of nematodes differs from that of most animals: they have a longitudinal layer only, and their direct attachment to the dorsal and ventral nerve cords creates a strong muscular contraction that results in a whiplike, almost spastic, body movement.

Photo a shows a worm-shaped nematode next to a capsule-shaped nematode egg. The illustration in part b shows a cross-section of a nematode, which has a mouth at one end and an anus at the other. The mouth connects to a pharynx, then to an intestine. A dorsal nerve runs along the top of the animal and joins ring-like head ganglia at the front end. Testes run alongside the intestine toward the back of the animal.
Figure 28.32 Nematode morphology. Scanning electron micrograph shows (a) the soybean cyst nematode (Heterodera glycines) and a nematode egg. (b) A schematic representation shows the anatomy of a typical nematode. (credit a: modification of work by USDA ARS; scale-bar data from Matt Russell)

Excretory System

In nematodes, specialized excretory systems are not well developed. Nitrogenous wastes, largely in the form of ammonia, are released directly across the body wall. In some nematodes, osmoregulation and salt balance are performed by simple excretory cells or glands that may be connected to paired canals that release wastes through an anterior pore. In marine nematodes, the excretory cells are called renette cells, which are unique to nematodes.

Nervous system

Most nematodes have four longitudinal nerve cords that run along the length of the body in dorsal, ventral, and lateral positions. The ventral nerve cord is better developed than the dorsal and lateral cords. Nonetheless, all nerve cords fuse at the anterior end, to form a pharyngeal nerve ring around the pharynx, which acts as the head ganglion or the “brain” of the roundworm. A similar fusion forms a posterior ganglion at the tail. In C. elegans, the nervous system accounts for nearly one-third of the total number of cells in the animal!

Reproduction

Nematodes employ a variety of reproductive strategies ranging from monoecious to dioecious to parthenogenetic, depending upon the species. C. elegans is a mostly monoecious species with both self-fertilizing hermaphrodites and some males. In the hermaphrodites, ova and sperm develop at different times in the same gonad. Ova are contained in a uterus and amoeboid sperm are contained in a spermatheca ("sperm receptacle"). The uterus has an external opening known as the vulva. The female genital pore is near the middle of the body, whereas the male genital pore is nearer to the tip. In anatomical males, specialized structures called copulatory spicules at the tail of the male keep him in place and open the vulva of the female into which the amoeboid sperm travel into the spermatheca.

Fertilization is internal, and embryonic development starts very soon after fertilization. The embryo is released from the vulva during the gastrulation stage. The embryonic development stage lasts for 14 hours; development then continues through four successive larval stages with molting and ecdysis taking place between each stage—L1, L2, L3, and L4—ultimately leading to the development of a young adult worm. Adverse environmental conditions such as overcrowding or lack of food can result in the formation of an intermediate larval stage known as the dauer larva. An unusual feature of some nematodes is eutely: the body of a given species contains a specific number of cells as the consequence of a rigid developmental pathway.

Everyday Connection

C. elegans: The Model System for Linking Developmental Studies with Genetics

If biologists wanted to research how nicotine dependence develops in the body, how lipids are regulated, or observe the attractant or repellant properties of certain odors, they would clearly need to design three very different experiments. However, they might only need one subject of study: Caenorhabditis elegans. The nematode C. elegans was brought into the focus of mainstream biological research by Dr. Sydney Brenner. Since 1963, Dr. Brenner and scientists worldwide have used this animal as a model system to study many different physiological and developmental mechanisms.

C. elegans is a free-living nematode found in soil. Only about a millimeter long, it can be cultured on agar plates (10,000 worms/plate!), feeding on the common intestinal bacterium Escherichia coli (another long-term resident of biological laboratories worldwide), and therefore can be readily grown and maintained in a laboratory. The biggest asset of this nematode is its transparency, which helps researchers to observe and monitor changes within the animal with ease. It is also a simple organism with about 1,000 cells and a genome of only 20,000 genes. Its chromosomes are organized into five pairs of autosomes plus a pair of sex chromosomes, making it an ideal candidate with which to study genetics. Since every cell can be visualized and identified, this organism is useful for studying cellular phenomena like cell-to-cell interactions, cell-fate determinations, cell division, apoptosis (cell death), and intracellular transport.

Another tremendous asset is the short life cycle of this worm (Figure 28.33). It takes only three days to achieve the “egg to adult to daughter egg”; therefore, the developmental consequences of genetic changes can be quickly identified. The total life span of C. elegans is two to three weeks; hence, age-related phenomena are also easy to observe. There are two sexes in this species: hermaphrodites (XX) and males (XO). However, anatomical males are relatively infrequently obtained from matings between hermaphrodites, since their XO chromosome composition requires meiotic nondisjunction when both parents are XX. Another feature that makes C. elegans an excellent experimental model is that the position and number of the 959 cells present in adult hermaphrodites of this organism is constant. This feature is extremely significant when studying cell differentiation, cell-to-cell communication, and apoptosis. Lastly, C. elegans is also amenable to genetic manipulations using molecular methods, rounding off its usefulness as a model system.

Biologists worldwide have created information banks and groups dedicated to research using C. elegans. Their findings have led, for example, to better understandings of cell communication during development, neuronal signaling, and insight into lipid regulation (which is important in addressing health issues like the development of obesity and diabetes). In recent years, studies have enlightened the medical community with a better understanding of polycystic kidney disease. This simple organism has led biologists to complex and significant findings, growing the field of science in ways that touch the everyday world.

Photo a shows transparent worm about a millimeter in length. Illustration B shows the life cycle of C. elegans, which begins when the egg hatches, releasing a L1 juvenile. After 12 hours the L1 juvenile transforms into an L2 juvenile. After 7 hours the L2 juvenile transforms into an L3 juvenile. After another 7 hours the L3 juvenile transforms into an L4 juvenile. After 14 hours the L4 juvenile transforms into an adult. The hermaphroditic adult mates with another adult to produce fertilized eggs which hatch, completing the cycle.
Figure 28.33 Caenorhabditis elegans. (a) This light micrograph shows the bodies of a group of roundworms. These hermaphrodites consist of exactly 959 cells. (b) The life cycle of C. elegans has four juvenile stages (L1 through L4) and an adult stage. Under ideal conditions, the nematode spends a set amount of time at each juvenile stage, but under stressful conditions, it may enter a dauer state that does not age significantly and is somewhat analogous to the diapausing state of some insects. (credit a: modification of work by “snickclunk”/Flickr: credit b: modification of work by NIDDK, NIH; scale-bar data from Matt Russell)

Parasitic Nematodes

A number of common parasitic nematodes serve as prime examples of parasitism (endoparasitism). These economically and medically important animals exhibit complex life cycles that often involve multiple hosts, and they can have significant medical and veterinary impacts. Here is a partial list of nasty nematodes: Humans may become infected by Dracunculus medinensis, known as guinea worms, when they drink unfiltered water containing copepods (Figure 28.34), an intermediate crustacean host. Hookworms, such as Ancylostoma and Necator, infest the intestines and feed on the blood of mammals, especially of dogs, cats, and humans. Trichina worms (Trichinella) are the causal organism of trichinosis in humans, often resulting from the consumption of undercooked pork; Trichinella can infect other mammalian hosts as well. Ascaris, a large intestinal roundworm, steals nutrition from its human host and may create physical blockage of the intestines. The filarial worms, such as Dirofilaria and Wuchereria, are commonly vectored by mosquitoes, which pass the infective agents among mammals through their blood-sucking activity. One species, Wuchereria bancrofti, infects the lymph nodes of over 120 million people worldwide, usually producing a non-lethal but deforming condition called elephantiasis. In this disease, parts of the body often swell to gigantic proportions due to obstruction of lymphatic drainage, inflammation of lymphatic tissues, and resulting edema. Dirofilaria immitis, a blood-infective parasite, is the notorious dog heartworm species.

 Part A shows a foot with a guinea worm extending from a blister. The end of the worm is wrapped around a stick. Part B shows the life cycle of the guinea worm, which begins when a person drinks unfiltered water containing copepods infected with guinea worm larvae. Larvae, which are released when the copepods die, penetrate the wall of the stomach and intestine. The worms mature and reproduce. Fertilized females migrate to the surface of the skin, where they discharge larvae into the water. Copepods consume the larvae. The copepods are consumed by humans, completing the cycle. About a year after infection, the female worm emerges from the skin.
Figure 28.34 Life cycle of the guinea worm. The guinea worm Dracunculus medinensis infects about 3.5 million people annually, mostly in Africa. (a) Here, the worm is wrapped around a stick so it can be slowly extracted. (b) Infection occurs when people consume water contaminated by infected copepods, but this can easily be prevented by simple filtration systems. (credit: modification of work by CDC)

Phylum Tardigrada

The tardigrades ("slow-steppers") comprise a phylum of inconspicuous little animals living in marine, freshwater, or damp terrestrial environments throughout the world. They are commonly called "water bears" because of their plump bodies and the large claws on their stubby legs. There are over 1,000 species, most of which are less than 1 mm in length. A chitinous cuticle covers the body surface and may be divided into plates (Figure 28.35). Tardigrades are known for their ability to enter a state called cryptobiosis, which provides them with are resistance to multiple environmental challenges, including desiccation, very low temperatures, vacuum, high pressure, and radiation. They can suspend their metabolic activity for years, and survive the loss of up to 99% of their water content. Their remarkable resistance has recently been attributed to unique proteins that replace water in their cells and protect their internal cell structure and their DNA from damage.

The image displays an electron micrgraph of a water bear, or tardigrade.
Figure 28.35 Scanning electronmicrograph of Milnesium tardigradum. (credit: Schokraie E, Warnken U, Hotz-Wagenblatt A, Grohme MA, Hengherr S, et al. (2012) -https://commons.wikimedia.org/w/index.php?curid=22716809)

Morphology and Physiology

Tardigrades have cylindrical bodies, with four pairs of legs terminating in a number of claws. The cuticle is periodically shed, including the cuticular covering of the claws. The first three pairs of legs are used for walking, and the posterior pair for clinging to the substrate. A circular mouth leads to a muscular pharynx and salivary glands. Tardigrades feed on plants, algae, or small animals. Plant cells are pierced with a chitinous stylet and the cellular contents are then sucked into the gut by the muscular pharynx. Bands of single muscle cells are attached to the various points of the epidermis and extend into the legs to provide ambulatory movement. The major body cavity is a hemocoel, but there are no specialized circulatory structures for moving the blood, nor are there specialized respiratory structures. Malpighian tubules in the hemocoel remove metabolic wastes and transport them to the gut. A dorsal brain is connected to a ventral nerve cord with segmental ganglia associated with the appendages. Sensory structures are greatly reduced, but there is a pair of simple eyespots on the head, and sensory cilia or bristles concentrated toward the head end of the animal.

Reproduction

Most tardigrades are dioecious, and males and females each have a single gonad. Mating usually occurs at the time of a molt and fertilization is external. Eggs may be deposited in a molted cuticle or attached to other objects. Development is direct, and the animal may molt a dozen times during its lifetime. In tardigrades, like nematodes, development produces a fixed number of cells, with the actual number of cells being dependent on the species. Further growth occurs by enlarging the cells, not by multiplying them.

Footnotes

  • 1 Stoll, N. R., “This wormy world. 1947,” Journal of Parasitology 85(3) (1999): 392-396.
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