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Biology 2e

28.7 Superphylum Deuterostomia

Biology 2e28.7 Superphylum Deuterostomia
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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 distinguishing characteristics of echinoderms
  • Describe the distinguishing characteristics of chordates

The phyla Echinodermata and Chordata (the phylum that includes humans) both belong to the superphylum Deuterostomia. Recall that protostomes and deuterostomes differ in certain aspects of their embryonic development, and they are named based on which opening of the archenteron (primitive gut tube) develops first. The word deuterostome comes from the Greek word meaning “mouth second,” indicating that the mouth develops as a secondary structure opposite the location of the blastopore, which becomes the anus. In protostomes (“mouth first”), the first embryonic opening becomes the mouth, and the second opening becomes the anus.

There are a series of other developmental characteristics that differ between protostomes and deuterostomes, including the type of early cleavage (embryonic cell division) and the mode of formation of the coelom of the embryo: Protosomes typically exhibit spiral mosaic cleavage whereas deuterostomes exhibit radial regulative cleavage. In deuterostomes, the endodermal lining of the archenteron usually forms buds called coelomic pouches that expand and ultimately obliterate the embryonic blastocoel (the cavity within the blastula and early gastrula) to become the embryonic mesoderm, the third germ layer. This happens when the mesodermal pouches become separated from the invaginating endodermal layer forming the archenteron, then expand and fuse to form the coelomic cavity. The resulting coelom is termed an enterocoelom. The archenteron develops into the alimentary canal, and a mouth opening is formed by invagination of ectoderm at the pole opposite the blastopore of the gastrula. The blastopore forms the anus of the alimentary system in the juvenile and adult forms. Cleavage in most deuterostomes is also indeterminant, meaning that the developmental fates of early embryonic cells are not decided at that point of embryonic development (this is why we could potentially clone most deuterostomes, including ourselves).

The deuterostomes consist of two major clades—the Chordata and the Ambulacraria. The Chordata include the vertebrates and two invertebrate subphyla, the urochordates and the cephalochordates. The Ambulacraria include the echinoderms and the hemichordates, which were once considered to be a chordate subphylum (Figure 28.46). The two clades, in addition to being deuterostomes, have some other interesting features in common. As we have seen, the vast majority of invertebrate animals do not possess a defined bony vertebral endoskeleton, or a bony cranium. However, one of the most ancestral groups of deuterostome invertebrates, the Echinodermata, do produce tiny skeletal “bones” called ossicles that make up a true endoskeleton, or internal skeleton, covered by an epidermis. The Hemichordata (acorn worms and pterobranchs) will not be covered here, but share with the echinoderms a three-part (tripartite) coelom, similar larval forms, and a derived metanephridium that rids the animals of nitrogenous wastes. They also share pharyngeal slits with the chordates (Figure 28.46). In addition, hemichordates have a dorsal nerve cord in the midline of the epidermis, but lack a neural tube, a true notochord and the endostyle and post-anal tail characteristic of chordates.

Part a shows the deuterostome taxa.  Deuterostomes lead to Ambulacraria and Chordates.  Ambulacraria lead to Echinoderms and Hemichordates.  Chordates lead to Cephalochordates, Urochordates, and Vertebrates.  Part b shows a hemichordates with a worm-like, helical body;  its pharyngeal slits appear on the top part of the helix.  And it also shows an Urochordate, a flattened creature with its pharyngeal slites across its flat front.
Figure 28.46 Ambulacraria and Chordata. (a) The major deuterostome taxa. (b) pharyngeal slits in hemichordates and urochordates. (credit a MAC; credit b modification of Gill Slits By Own work by Zebra.element [Public domain], via Wikimedia Commons)

Phylum Echinodermata

Echinodermata are named after their “prickly skin” (from the Greek “echinos” meaning “prickly” and “dermos” meaning “skin”). This phylum is a collection of about 7,000 described living species of exclusively marine, bottom-dwelling organisms. Sea stars (Figure 28.47), sea cucumbers, sea urchins, sand dollars, and brittle stars are all examples of echinoderms.

Morphology and Anatomy

Despite the adaptive value of bilaterality for most free-living cephalized animals, adult echinoderms exhibit pentaradial symmetry (with “arms” typically arrayed in multiples of five around a central axis). Echinoderms have an endoskeleton made of calcareous ossicles (small bony plates), covered by the epidermis. For this reason, it is an endoskeleton like our own, not an exoskeleton like that of arthropods. The ossicles may be fused together, embedded separately in the connective tissue of the dermis, or be reduced to minute spicules of bone as in sea cucumbers. The spines for which the echinoderms are named are connected to some of the plates. The spines may be moved by small muscles, but they can also be locked into place for defense. In some species, the spines are surrounded by tiny stalked claws called pedicellaria, which help keep the animal's surface clean of debris, protect papulae used in respiration, and sometimes aid in food capture.

The endoskeleton is produced by dermal cells, which also produce several kinds of pigments, imparting vivid colors to these animals. In sea stars, fingerlike projections (papillae) of dermal tissue extend through the endoskeleton and function as gills. Some cells are glandular, and may produce toxins. Each arm or section of the animal contains several different structures: for example, digestive glands, gonads, and the tube feet that are unique to the echinoderms. In echinoderms like sea stars, every arm bears two rows of tube feet on the oral side, running along an external ambulacral groove. These tube feet assist in locomotion, feeding, and chemical sensations, as well as serve to attach some species to the substratum.

The illustration shows a sea star, which has a mouth on the bottom and an anus on top, both in the middle of the star. The disk-shaped stomach is sandwiched between the mouth and anus. Two tubes radiate from the stomach to each arm, and many small digestive glands connect to these tubes. Beneath the stomach is a central ring canal that also connects to tubes that extend into each arm. Tube feet are attached to these tubes. Each tube foot resembles a medicine dropper, with a bulb-shaped ampulla at the top and an extension called a podium at the bottom. The bottom of the podium protrudes from the bottom of the starfish. There are many podia along the length of the arm, which allow the sea star to latch onto objects and walk. A structure called a madreporite connects to the central ring, and protrudes from the upper surface of the sea star, next to the anus.
Figure 28.47 Anatomy of a sea star. This diagram of a sea star shows the pentaradial pattern typical of adult echinoderms, and the water vascular system that is their defining characteristic.

Water Vascular and Hemal Systems

Echinoderms have a unique ambulacral (water vascular) system, derived from part of the coelom, or “body cavity.” The water vascular system consists of a central ring canal and radial canals that extend along each arm. Each radial canal is connected to a double row of tube feet, which project through holes in the endoskeleton, and function as tactile and ambulatory structures. These tube feet can extend or retract based on the volume of water present in the system of that arm, allowing the animal to move and also allowing it to capture prey with their suckerlike action. Individual tube feet are controlled by bulblike ampullae. Seawater enters the system through an aboral madreporite (opposite the oral area where the mouth is located) and passes to the ring canal through a short stone canal. Water circulating through these structures facilitates gaseous exchange and provides a hydrostatic source for locomotion and prey manipulation. A hemal system, consisting of oral, gastric, and aboral rings, as well as other vessels running roughly parallel to the water vascular system, circulates nutrients. Transport of nutrients and gases is shared by the water vascular and hemal systems in addition to the visceral body cavity that surrounds the major organs.

Nervous System

The nervous system in these animals is a relatively simple, comprising a circumoral nerve ring at the center and five radial nerves extending outward along the arms. In addition, several networks of nerves are located in different parts of the body. However, structures analogous to a brain or large ganglia are not present in these animals. Depending on the group, echinoderms may have well-developed sensory organs for touch and chemoreception (e.g., within the tube feet and on tentacles at the tips of the arms), as well as photoreceptors and statocysts.

Digestive and Excretory Systems

A mouth, located on the oral (ventral) side, opens through a short esophagus to a large, baglike stomach. The so-called “cardiac” stomach can be everted through the mouth during feeding (for example, when a starfish everts its stomach into a bivalve prey item to digest the animal—alive—within its own shell!) There are masses of digestive glands (pyloric caeca) in each arm, running dorsally along the arms and overlying the reproductive glands below them. After passing through the pyloric caeca in each arm, the digested food is channeled to a small anus, if one exists.

Podocytes—cells specialized for ultrafiltration of bodily fluids—are present near the center of the echinoderm disc, at the junction of the water vascular and hemal systems. These podocytes are connected by an internal system of canals to the madreporite, where water enters the stone canal. The adult echinoderm typically has a spacious and fluid-filled coelom. Cilia aid in circulating the fluid within the body cavity, and lead to the fluid-filled papulae, where the exchange of oxygen and carbon dioxide takes place, as well as the secretion of nitrogenous waste such as ammonia, by diffusion.

Reproduction

Echinoderms are dioecious, but males and females are indistinguishable apart from their gametes. Males and females release their gametes into water at the same time and fertilization is external. The early larval stages of all echinoderms (e.g., the bipinnaria of asteroid echinoderms such as sea stars) have bilateral symmetry, although each class of echinoderms has its own larval form. The radially symmetrical adult forms from a cluster of cells in the larva. Sea stars, brittle stars, and sea cucumbers may also reproduce asexually by fragmentation, as well as regenerate body parts lost in trauma, even when over 75 percent of their body mass is lost!

Classes of Echinoderms

This phylum is divided into five extant classes: Asteroidea (sea stars), Ophiuroidea (brittle stars), Echinoidea (sea urchins and sand dollars), Crinoidea (sea lilies or feather stars), and Holothuroidea (sea cucumbers) (Figure 28.48).

The most well-known echinoderms are members of class Asteroidea, or sea stars. They come in a large variety of shapes, colors, and sizes, with more than 1,800 species known so far. The key characteristic of sea stars that distinguishes them from other echinoderm classes includes thick arms that extend from a central disk from which various body organs branch into the arms. At the end of each arm are simple eye spots and tentacles that serve as touch receptors. Sea stars use their rows of tube feet not only for gripping surfaces but also for grasping prey. Most sea stars are carnivores and their major prey are in the phylum Mollusca. By manipulating its tube feet, a sea star can open molluscan shells. Sea stars have two stomachs, one of which can protrude through their mouths and secrete digestive juices into or onto prey, even before ingestion. A sea star eating a clam can partially open the shell, and then evert its stomach into the shell, introducing digestive enzymes into the interior of the mollusk. This process can both weaken the strong adductor (closing) muscles of a bivalve and begin the process of digestion.

Link to Learning

Explore the sea star’s body plan up close, watch one move across the sea floor, and see it devour a mussel.

Brittle stars belong to the class Ophiuroidea ("snake-tails"). Unlike sea stars, which have plump arms, brittle stars have long, thin, flexible arms that are sharply demarcated from the central disk. Brittle stars move by lashing out their arms or wrapping them around objects and pulling themselves forward. Their arms are also used for grasping prey. The water vascular system in ophiuroids is not used for locomotion.

Sea urchins and sand dollars are examples of Echinoidea ("prickly"). These echinoderms do not have arms, but are hemispherical or flattened with five rows of tube feet that extend through five rows of pores in a continuous internal shell called a test. Their tube feet are used to keep the body surface clean. Skeletal plates around the mouth are organized into a complex multipart feeding structure called "Aristotle's lantern." Most echinoids graze on algae, but some are suspension feeders, and others may feed on small animals or organic detritus—the fragmentary remains of plants or animals.

Sea lilies and feather stars are examples of Crinoidea. Sea lilies are sessile, with the body attached to a stalk, but the feather stars can actively move about using leglike cirri that emerge from the aboral surface. Both types of crinoid are suspension feeders, collecting small food organisms along the ambulacral grooves of their feather-like arms. The "feathers" consisted of branched arms lined with tube feet. The tube feet are used to move captured food toward the mouth. There are only about 600 extant species of crinoids, but they were far more numerous and abundant in ancient oceans. Many crinoids are deep-water species, but feather stars typically inhabit shallow areas, especially in substropical and tropical waters.

Sea cucumbers of class Holothuroidea exhibit an extended oral-aboral axis. These are the only echinoderms that demonstrate “functional” bilateral symmetry as adults, because the extended oral-aboral axis compels the animal to lie horizontally rather than stand vertically. The tube feet are reduced or absent, except on the side on which the animal lies. They have a single gonad and the digestive tract is more typical of a bilaterally symmetrical animal. A pair of gill-like structures called respiratory trees branch from the posterior gut; muscles around the cloaca pump water in and out of these trees. There are clusters of tentacles around the mouth. Some sea cucumbers feed on detritus, while others are suspension feeders, sifting out small organisms with their oral tentacles. Some species of sea cucumbers are unique among the echinoderms in that cells containing hemoglobin circulate in the coelomic fluid, the water vascular system and/or the hemal system.

The sea star in photo a is red and white, with a thick squat body and protruding spikes. The brittle star in part b is brown with a flat, pentagon-shaped body. Thin striped legs extend from each point of the pentagon. Photo c shows a sea urchin with a round, black body and very long, thin, black spines. Photo d shows a sea lily that has appendages resembling branches of a spruce tree. Photo e shows a log-shaped sea cucumber with spikes extending from its body.
Figure 28.48 Classes of echinoderms. Different members of Echinodermata include the (a) sea star of class Asteroidea, (b) the brittle star of class Ophiuroidea, (c) the sea urchins of class Echinoidea, (d) the sea lilies belonging to class Crinoidea, and (e) sea cucumbers, representing class Holothuroidea. (credit a: modification of work by Adrian Pingstone; credit b: modification of work by Joshua Ganderson; credit c: modification of work by Samuel Chow; credit d: modification of work by Sarah Depper; credit e: modification of work by Ed Bierman)

Phylum Chordata

Animals in the phylum Chordata share five key features that appear at some stage of their development: a notochord, a dorsal hollow nerve cord, pharyngeal slits, a post-anal tail, and an endostyle/thyroid gland that secretes iodinated hormones. In some groups, some of these traits are present only during embryonic development. In addition to containing vertebrate classes, the phylum Chordata contains two clades of “invertebrates”: Urochordata (tunicates, salps, and larvaceans) and Cephalochordata (lancelets). Most tunicates live on the ocean floor and are suspension feeders. Lancelets are suspension feeders that feed on phytoplankton and other microorganisms. The invertebrate chordates will be discussed more extensively in the following chapter.

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