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Chordates - Advanced Douglas Wilkin, Ph.D. Jennifer Blanchette Say Thanks to the Authors Click http://www.ck12.org/saythanks (No sign in required) To access a customizable version of this book, as well as other interactive content, visit www.ck12.org CK-12 Foundation is a non-profit organization with a mission to reduce the cost of textbook materials for the K-12 market both in the U.S. and worldwide. Using an open-source, collaborative, and web-based compilation model, CK-12 pioneers and promotes the creation and distribution of high-quality, adaptive online textbooks that can be mixed, modified and printed (i.e., the FlexBook® textbooks). Copyright © 2016 CK-12 Foundation, www.ck12.org The names “CK-12” and “CK12” and associated logos and the terms “FlexBook®” and “FlexBook Platform®” (collectively “CK-12 Marks”) are trademarks and service marks of CK-12 Foundation and are protected by federal, state, and international laws. Any form of reproduction of this book in any format or medium, in whole or in sections must include the referral attribution link http://www.ck12.org/saythanks (placed in a visible location) in addition to the following terms. Except as otherwise noted, all CK-12 Content (including CK-12 Curriculum Material) is made available to Users in accordance with the Creative Commons Attribution-Non-Commercial 3.0 Unported (CC BY-NC 3.0) License (http://creativecommons.org/ licenses/by-nc/3.0/), as amended and updated by Creative Commons from time to time (the “CC License”), which is incorporated herein by this reference. Complete terms can be found at http://www.ck12.org/about/ terms-of-use. Printed: January 3, 2016 AUTHORS Douglas Wilkin, Ph.D. Jennifer Blanchette www.ck12.org C HAPTER Chapter 1. Chordates - Advanced 1 Chordates - Advanced • Discuss the major characteristics of chordates, especially those that distinguish chordates from other phyla. What’s a chordate? Most chordates are animals with backbones. These range from small fish to giant whales and from playful dogs to ferocious cats. They are not nearly as diverse as the invertebrates. But chordates do have many amazing species. The white tiger (Panthera tigris) is a chordate. The tiger is also from the class Mammalia, order Carnivora, and family Felidae, meaning it is a meat-eating cat. 1 www.ck12.org Characteristics of Chordates The animal phylum Chordata includes the animals that we are by far the most familiar with: ourselves! We and all other chordate species, including some quite primitive looking organisms, share several important features that indicate our close evolutionary relationships. All of the phyla we have studied so far have consisted entirely of invertebrate animals. The phylum Chordata includes both nonvertebrate and vertebrate groups. Some groups within the chordates are considered to be very “young” compared to other chordates and to many of the phyla that we have discussed in previous chapters. Birds, for example, have only been around for about 146 million years. Compare that to some arthropods, such as horseshoe crabs, that have existed in basically the same form for over 500 million years. In this lesson we will examine the specific features that define chordates. We will then consider the different taxonomic groups within the phylum and how they evolved, keeping an emphasis on the nonvertebrate chordates. Chordata is a large and diverse phylum that includes roughly 60,000 species. The examples of chordate species shown in the Figure 1.1 illustrate their diversity and vast range in size. Chordates can be anywhere from about one cm (the width of your index finger) to over 30 meters. FIGURE 1.1 Different species of chordates, illustrating their diversity and vast range in size. (a) A tunicate, approximately one inch in length. Notice the tiny fish (also a chordate) swimming in front of the tunicate. (b) A white rhinoceros, approximately 6600 pounds (3000 kg). (c) A kangaroo and joey. (d) A great white shark. (e) A double-crested cormorant bird species. (f) A blue whale. Tunicates, for example, are small marine invertebrate members of the phylum that are generally one to a few centimeters long. In contrast, some mammalian species can be quite large, either in length (for example, blue whales can extend up to 30 meters in length) or in weight and girth (such as elephants, which can weigh up to roughly 20,000 lbs). Chordates can be found all over the planet, with species inhabiting marine, freshwater, terrestrial, and aerial environments. They are found from the equator to the poles and everywhere in between. Members of the phylum Chordata share the following characteristics (those traits specific to the phylum Chordata will be in bold): • • • • • • • • • • • 2 Three germ layers (triploblasts). Bilateral symmetry. Segmented body. Coelom. Complete digestive system. Closed circulatory system. Centralized nervous system. Tail (post-anal). Notochord. Nerve cord. Pharyngeal slits. www.ck12.org Chapter 1. Chordates - Advanced As you read this list, you may be thinking, “humans don’t have tails or slits in their pharynx!” But, in fact, we do. Actually, to be more precise, we did. All chordates possess a tail and pharyngeal slits at some point in their lives, and humans are no exception. Early on in human development, the embryo has both a tail and pharyngeal slits, both of which are lost during the course of development. A human embryo with a prominent tail is shown in the Figure 1.2. FIGURE 1.2 A five-week old (gestational age), ectopic human embryo. Notice the prominent tail that extends beyond the anal region. This embryo is ectopic, meaning it did not successfully implant within the uterine cavity. This complication is almost always fatal to the embryo and is often dangerous to the mother. Chordates, like species in several other phyla such as annelids and arthropods, are bilaterally symmetrical triploblasts. You will recall that bilaterally symmetrical animals can be divided into two equal, mirror-image halves when cut down the middle of the anterior-posterior (front-back) axis. Triploblasts are organisms that develop from three basic cell layers, called germ layers, in the embryo, which are called the endoderm, the mesoderm, and the ectoderm. The endoderm forms the gut of the animal, the ectoderm forms the outer surface and the nervous system, and the mesoderm forms the coelom and many of the internal organs including the muscles and circulatory system. Like the annelids and arthropods, most chordates have at least partially segmented bodies. The segments are often highly specialized into distinct body regions and do not resemble the repeating segments of annelids. In addition, chordates are deuterostomes, meaning that they have a coelom (fluid-filled body cavity) that arises from internalized pouches of tissue in the developing embryo and an anus that is formed from the embryonic blastopore. Chordate species have a complete digestive system, and most species also have a closed circulatory system that relies on a ventral (located on the bottom side of the organism’s interior) heart to pump blood throughout the body in enclosed blood vessels. The chordate nervous system is usually highly centralized, with a distinct anterior brain that, in some groups (including humans), is enclosed in a solid protective case called a cranium, or skull. The four most prominent characteristics that distinguish chordates from species in all other phyla are the presence of a post-anal tail, a notochord, a dorsal, hollow nerve cord, and pharyngeal slits (also called gill slits). The organization of these features within the body plan of a chordate is illustrated in the Figure 1.3. The last three of these features will be discussed in more detail in the following sections. 3 www.ck12.org FIGURE 1.3 The Body Plan of a Typical Chordate. The body plan of a chordate includes a postanal tail, a notochord, a dorsal, hollow nerve cord, and pharyngeal slits. Notochord The most important thing to remember about a notochord is that it is not a nerve cord or a backbone (column of vertebrae). A backbone is the distinguishing feature of the subphylum Vertebrata within the phylum Chordata (this subphylum will be discussed in the next section). It is easy to confuse the notochord with either of these because of its name and its function. The notochord is a long, rod-like support structure that usually extends the length of the animal. As shown in the Figure 1.3, the notochord is situated between the gut (it is dorsal to the gut) and the dorsal nerve cord (it is ventral to the nerve cord). Chordates are defined as having a notochord at some point during their lives, but they do not all retain the notochord as adults. In some groups it is completely lost, in others (generally nonvertebrate chordate species) it is retained and functions to support the body of the animal during locomotion. In adult vertebrates, the notochord is essentially replaced by the backbone, however, the notochord should not be considered a primitive backbone. It is a separate structure entirely. All chordates have a notochord during embryonic development when it plays an important role in specifying the organization of the developing embryo. For example, the presence of the notochord communicates to nearby cells that they should differentiate in certain ways based on their position relative to the notochord. Nerve Cord A dorsal (top side), hollow nerve cord derived from ectodermal tissue is another characteristic that distinguishes chordates from other animals. Most other bilateral animal phyla with a nervous system have a solid, ventral (bottom side) nerve cord. Unlike the notochord, essentially all vertebrate chordates retain the nerve cord as adults. The nerve cord ultimately develops into the brain and spinal cord in most adult vertebrates. It is the core of the centralized nervous system. Pharyngeal Slits Like the notochord, pharyngeal slits are a feature shared by all developing chordate embryos but not by all chordate adults. A pharynx is the region of the digestive tract that functions to connect the mouth to the esophagus. In many animals, including humans, it is also a junction between the respiratory system and the digestive system, connecting the mouth (digestive opening) and nasal passages to the bronchial tubes of the lungs. The pharyngeal slits are formed from invaginations, or pouches, in the lining of the pharynx. The developing pharyngeal pouches eventually form openings through the pharyngeal wall called slits. This is shown in the Figure 1.4. The slits function in feeding and also in respiration in some species. Water flows through the slits as a result of the action of beating hair-like projections, called cilia, that line the slits, and food particles are trapped for ingestion. In most aquatic species, the pharyngeal slits ultimately develop into the gills of the adult organism, and they are used to absorb oxygen from the incoming water. In non-aquatic species, the pouches/slits develop into other structures connected to the pharyngeal region including components of the middle ear and the tonsils in humans. Now that we have examined the overall characteristics of chordates, let’s consider how species within the phylum 4 www.ck12.org Chapter 1. Chordates - Advanced FIGURE 1.4 A diagram of a cross-section through a chordate pharynx. The developing pharyngeal pouches on the right side are labeled 1 through 4. As the pouches extend to the outer edge of the pharynx, they will eventually extend through the pharyngeal wall to form slits, as shown in the uppermost pouch on the left. are classified with regard to their evolutionary history. Vocabulary • blastopore: The opening of an embryo’s central cavity in the early stage of development. • cilia (singular, cilium): Short hairlike projection, similar to flagella, that allow some cells to move. • coelom: A fluid-filled cavity formed within the mesoderm; it forms between the digestive cavity and the body wall. • cranium: The skull of a vertebrate. • deuterostome: An animal in which the first opening formed during development (the blastopore) becomes the anus. • nerve cord: A single hollow tract of nervous tissue; it forms the central nervous system of chordates and develops into the spinal cord and brain in vertebrates. • notochord: A rod-shaped, semi-rigid support structure that forms between the dorsal nerve cord and the gut of an animal. • pharyngeal slits: Openings between the pharynx, or throat, and the outside. • triploblast: An animal with three germ layers: an endoderm, a mesoderm, and an ectoderm. Summary • Chordates are bilaterally symmetrical triploblasts with true coeloms and segmented bodies. • Like echinoderms, chordates are deuterostomes. • The four most prominent characteristics that distinguish chordates from species in all other phyla are the presence of a post-anal tail, a notochord, a dorsal, hollow nerve cord, and pharyngeal slits (also called gill slits). • The notochord is a long, rod-like support structure that usually extends the length of the animal. • While a notochord and pharyngeal slits might not be present in all adult chordates, all vertebrate chordates retain the nerve cord as adults. 5 www.ck12.org Practice Use this resource to answer the questions that follow. • Chordate at http://www.britannica.com/EBchecked/topic/114462/chordate . 1. How do the pharyngeal slits function in tunicates and cephalochordates? How does this differ from vertebrates? 2. How do chordates reproduce? 3. What function do notocords serve in locomotion? Review 1. 2. 3. 4. 5. What are the four features that distinguish chordates from other phyla? As a human develops, which chordate characteristics do we lose as we turn into adults? What important role does the notochord play in embryonic development? How does the nerve cord of chordates differ from the nervous systems of other animals? What function do the pharyngeal slits serve? References 1. Tunicate: Chika Watanabe; Rhinoceros: Brian Snelson; Kangaroo: Subhash Chandra; Shark: Terry Goss; Cormorant: Linda Tanner; Whale: Courtesy of US National Oceanic and Atmospheric Administration. Tunicate: http://www.flickr.com/photos/chikawatanabe/61157859/; Rhinoceros: http://www.flickr.com/photos/exfordy /2961323775/; Kangaroo: http://www.flickr.com/photos/ysc/2179798100/; Shark: http://commons.wikimedi a.org/wiki/File:White_shark.jpg; Cormorant: http://www.flickr.com/photos/goingslo/2306021399/in/photostr eam/; Whale: http://www.nmfs.noaa.gov/pr/species/mammals/cetaceans/bluewhale.htm . Tunicate, Rhinoceros, Kangaroo, Shark, Cormorant: CC BY 2.0; Whale: Public Domain 2. Ed Uthman, MD. http://commons.wikimedia.org/wiki/File:Tubal_Pregnancy_with_embryo.jpg . Public Domain 3. Mariana Ruiz Villarreal (LadyofHats) for CK-12 Foundation. Body plan of a typical chordate . CC BY-NC 3.0 4. Uwe Gille. http://en.wikipedia.org/wiki/Image:Kiemenbogen.jpg . CC-BY-SA 6