28.5: Superphylum Deuterostomia

28.5.1: Phylum Echinodermata

Learning Objective

Describe the characteristics of echinodermata

Key Points

• Echinoderms live exclusively in marine systems; they are widely divergent, with over 7,000 known species in the phylum.
• Echinoderms have pentaradial symmetry and a calcareous endoskeleton that may possess pigment cells that give them a wide range of colors, as well as cells that possess toxins.
• Echinoderms have a water vascular system composed of a central ring of canals that extend along each arm, through which water circulates for gaseous exchange and nutrition.
• Echinoderms have a very simple nervous system, comprised of a nerve ring at the center and five radial nerves extending outward along the arms; there is no structure resembling a brain.
• There are two sexes in echinoderms, which each release their eggs and sperm into the water; here, the sperm will fertilize the eggs.
• Echinoderms can reproduce asexually by regeneration.

Key Terms

water vascular system

a hydraulic system used by echinoderms, such as sea stars and sea urchins, for locomotion, food and waste transportation, and respiration

pentaradial symmetry

a variant of radial symmetry that arranges roughly equal parts around a central axis at orientations of 72° apart

podocyte

cells that filter the bodily fluids in echinoderms

madreporite

a lightcolored calcerous opening used to filter water into the water vascular system of echinoderms

ampulla

the dilated end of a duct

Phylum Echinodermata

Echinodermata are so named owing to their spiny skin (from the Greek "echinos" meaning "spiny" and "dermos" meaning "skin"). This phylum is a collection of about 7,000 described living species. Echinodermata are exclusively marine organisms. Sea stars, sea cucumbers, sea urchins, sand dollars, and brittle stars are all examples of echinoderms . To date, no freshwater or terrestrial echinoderms are known.

Sea stars

Star stars are among the most familiar of marine invertebrates. They are members of the phylum Echinodermata.

Morphology and Anatomy

Adult echinoderms exhibit pentaradial symmetry and have a calcareous endoskeleton made of ossicles, although the early larval stages of all echinoderms have bilateral symmetry . The endoskeleton is developed by epidermal cells and may possess pigment cells that give vivid colors to these animals, as well as cells laden with toxins. Echinoderms possess a simple digestive system which varies according to the animal's diet. Starfish are mostly carnivorous and have a mouth, oesophagus, two-part pyloric stomach with a pyloric duct leading to the intestine and rectum, with the anus located in the center of the aboral body surface. In many species, the large cardiac stomach can be everted and digest food outside the body. Gonads are present in each arm. In echinoderms such as sea stars, every arm bears two rows of tube feet on the oral side which help in attachment to the substratum. These animals possess a true coelom that is modified into a unique circulatory system called a water vascular system. The more notably distinct trait, which most echinoderms have, is their remarkable powers of regeneration of tissue, organs, limbs, and, in some cases, complete regeneration from a single limb.

Sea cucumbers

Sea cucumbers are a member of the phylum Echinodermata which are found on the sea floor worldwide. As with all echinoderms, sea cucumbers have an endoskeleton just below the skin, calcified structures that are usually reduced to isolated microscopic ossicles joined by connective tissue.

Water Vascular System

Echinoderms possess a unique ambulacral or water vascular system, consisting of a central ring canal and radial canals that extend along each arm . Water circulates through these structures and facilitates gaseous exchange as well as nutrition, predation, and locomotion. The water vascular system also projects from holes in the skeleton in the form of tube feet. These tube feet can expand or contract based on the volume of water (hydrostatic pressure) present in the system of that arm.

The water vascular system of the sea star

1. Pyloric stomach 2. Intestine 3. Rectal gland 4. Stone canal 5. Madreporite 6. Pyloric duct 7. Pyloric cecum 8. Cardiac stomach 9. Gonad 10. Ambulacral plates 11. Ampullae

The madreporite is a light-colored, calcerous opening used to filter water into the water vascular system of echinoderms. Acting as a pressure-equalizing valve, it is visible as a small red or yellow button-like structure (similar to a small wart) on the aboral surface of the central disk of a sea star. Close up, it is visibly structured, resembling a "madrepore" colony. From this, it derives its name. Water enters the madreporite on the aboral side of the echinoderm. From there, it passes into the stone canal, which moves water into the ring canal. The ring canal connects the radial canals (there are five in a pentaradial animal), and the radial canals move water into the ampullae, which have tube feet through which the water moves. By moving water through the unique water vascular system, the echinoderm can move and force open mollusk shells during feeding.

Other Body Systems

The nervous system in these animals is a relatively simple structure with a nerve ring at the center and five radial nerves extending outward along the arms. Structures analogous to a brain or derived from fusion of ganglia are not present in these animals.

Podocytes, cells specialized for ultrafiltration of bodily fluids, are present near the center of echinoderms. These podocytes are connected by an internal system of canals to the madreporite.

Echinoderms are sexually dimorphic and release their eggs and sperm cells into water; fertilization is external. In some species, the larvae divide asexually and multiply before they reach sexual maturity. Echinoderms may also reproduce asexually, as well as regenerate body parts lost in trauma.

28.5.2: Classes of Echinoderms

Learning Objective

Differentiate among the classes of echinoderms

Key Points

• Sea stars have thick arms called ambulacra that are used for gripping surfaces and grabbing hold of prey.
• Brittle stars have thin arms that wrap around prey or objects to pull themselves forward.
• Sea urchins and sand dollars embody flattened discs that do not have arms, but do have rows of tube feet they use for movement.
• Sea cucumbers demonstrate "functional" bilateral symmetry as adults because they actually lie horizontally rather than stand vertically.
• Sea lilies and feather stars are suspension feeders.

Key Terms

ossicle

a small bone (or bony structure), especially one of the three of the middle ear

fissiparous

of cells that reproduce through fission, splitting into two

ambulacrum

a row of pores for the protrusion of appendages such as tube feet.

Classes of Echinoderms

The phylum echinoderms 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).

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 (ambulacra; singular: ambulacrum) that extend from a central disk where organs penetrate into the arms. Sea stars use their tube feet not only for gripping surfaces, but also for grasping prey. Sea stars have two stomachs, one of which can protrude through their mouths and secrete digestive juices into or onto prey, even before ingestion. This process can essentially liquefy the prey, making digestion easier.

Brittle stars belong to the class Ophiuroidea. Unlike sea stars, which have plump arms, brittle stars have long, thin 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. Of all echinoderms, the Ophiuroidea may have the strongest tendency toward 5-segment radial (pentaradial) symmetry. Ophiuroids are generally scavengers or detritivores. Small organic particles are moved into the mouth by the tube feet. Ophiuroids may also prey on small crustaceans or worms. Some brittle stars, such as the six-armed members of the family Ophiactidae, are fissiparous (divide though fission), with the disk splitting in half. Regrowth of both the lost part of the disk and the arms occur, yielding an animal with three large arms and three small arms during the period of growth.

Sea urchins and sand dollars are examples of Echinoidea . These echinoderms do not have arms, but are hemispherical or flattened with five rows of tube feet that help them in slow movement; tube feet are extruded through pores of a continuous internal shell called a test. Like other echinoderms, sea urchins are bilaterans. Their early larvae have bilateral symmetry, but they develop fivefold symmetry as they mature. This is most apparent in the "regular" sea urchins, which have roughly spherical bodies, with five equally-sized parts radiating out from their central axes. Several sea urchins, however, including the sand dollars, are oval in shape, with distinct front and rear ends, giving them a degree of bilateral symmetry. In these urchins, the upper surface of the body is slightly domed, but the underside is flat, while the sides are devoid of tube feet. This "irregular" body form has evolved to allow the animals to burrow through sand or other soft materials.

Sea urchins

Sea urchins do not have arms, but have rows of tube feet that can be extended out of pores of the internal shell.

Sea lilies and feather stars are examples of Crinoidea. Both of these species are suspension feeders. They live both in shallow water and in depths as great as 6,000 meters. Sea lilies refer to the crinoids which, in their adult form, are attached to the sea bottom by a stalk . Feather stars or comatulids refer to the unstalked forms. Crinoids are characterized by a mouth on the top surface that is surrounded by feeding arms. They have a U-shaped gut; their anus is located next to the mouth. Although the basic echinoderm pattern of fivefold symmetry can be recognized, most crinoids have many more than five arms. Crinoids usually have a stem used to attach themselves to a substrate, but many live attached only as juveniles and become free-swimming as adults.

Sea lilies

Sea lilies, like feather stars, have a mouth on their upper surface that is surrounded by arms used for feeding.

Sea cucumbers of class Holothuroidea are extended in the oral-aboral axis and have five rows of tube feet . These are the only echinoderms that demonstrate "functional" bilateral symmetry as adults because the uniquely-extended oral-aboral axis compels the animal to lie horizontally rather than stand vertically. Like all echinoderms, sea cucumbers have an endoskeleton just below the skin: calcified structures that are usually reduced to isolated microscopic ossicles joined by connective tissue. In some species these can sometimes be enlarged to flattened plates, forming armor. In pelagic species, such as Pelagothuria natatrix, the skeleton and a calcareous ring are absent.

Sea cucumbers

Sea cucumbers are the only echinoderms that demonstrate "functional" bilateral symmetry as adults, as they lie horizontally as opposed to the vertical axis of other echinoderms.

28.5.3: Phylum Chordata

Learning Objective

Name the features that distinguish the members of the phylum chordata

Key Points

• The phylum chordata is named for the notochord, a longitudinal, flexible rod between the digestive tube and the nerve cord; in vertebrates, this is the spinal column.
• The chordates are also characterized by a dorsal nerve cord, which splits into the brain and spinal cord.
• Chordata contains two clades of invertebrates: Urochordata (tunicates) and Cephalochordata (lancelets), both of which are suspension feeders.
• The phylum chordata includes all animals that share four characteristics, although they might each possess some of them at different stages of their development: a notochord, a dorsal nerve cord, pharyngeal slits, and a postanal tail.
• Chordata contains five classes of animals: fish, amphibians, reptiles, birds, and mammals; these classes are separated by whether or not they can regulate their body temperature, the manner by which they consume oxygen, and their method of reproduction.

Key Terms

pharyngeal slit

filter-feeding organs found in non-vertebrate chordates (lancelets and tunicates) and hemichordates living in aquatic environments

notochord

a flexible rodlike structure that forms the main support of the body in the lowest chordates; a primitive spine

dorsal nerve cord

a hollow cord dorsal to the notochord, formed from a part of the ectoderm that rolls, forming a hollow tube.

Phylum Chordata

Animals in the phylum Chordata share four key features that appear at some stage of their development:

• A notochord, or a longitudinal, flexible rod between the digestive tube and the nerve cord . In most vertebrates, it is replaced developmentally by the vertebral column. This is the structure for which the phylum is named.
• A dorsal nerve cord which develops from a plate of ectoderm that rolls into a tube located dorsal to the notochord. Other animal phyla have solid nerve cords ventrally located. A chordate nerve cord splits into the central nervous system: the brain and spinal cord.
• Pharyngeal slits, which allow water that enters through the mouth to exit without continuing through the entire digestive tract. In many of the invertebrate chordates, these function as suspension feeding devices; in vertebrates, they have been modified for gas exchange, jaw support, hearing, and other functions.
• A muscular, postanal tail which extends posterior to the anus. The digestive tract of most nonchordates extends the length of the body. In chordates, the tail has skeletal elements and musculature, and can provide most of the propulsion in aquatic species.

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) and Cephalochordata (lancelets). However, even though they are invertebrates, they share characteristics with other chordates that places them in this phylum. For example, tunicate larvae have both a notochord and a nerve cord which are lost in adulthood . Most tunicates live on the ocean floor and are suspension feeders. Cephalochordates, or lancelets, have a notochord and a nerve cord (but no brain or specialist sensory organs) and a very simple circulatory system. Lancelets are suspension feeders that feed on phytoplankton and other microorganisms.

Structures present in a tunicate larva

While tunicates are invertebrates and may seem very different from the more familiar members of Chordata, the tunicate larva possesses both a notochord and a dorsal nerve cord, although both are lost in adulthood.

The phylum Chordata contains all of the animals that have a rod-like structure used to give them support. In most cases this is the spine or backbone. Within Chordata there are five classes of animals: fish, amphibians, reptiles, birds, and mammals. Three dividing factors separate these classes:

• Regulation of body temperature: animals are either homeothermic (can regulate their internal temperature so that it is kept at an optimum level) or poikilothermic (cannot regulate their internal temperature, the environment affects how hot or cold they are)
• Oxygen Absorption: the way in which oxygen is taken in from the air, which can be through gills, the skin (amphibians), or lungs
• Reproduction: this factor is particularly varied. Animals can be oviparous (lay eggs) or viviparous (birth live young). Fertilization can occur externally or internally. In mammals, the mother produces milk for the young.

Notochord

All chordates possess a notochord, or a type of flexible support rod, at some point in their development. In this dissected lungfish, which is a member of the chordates, tissues have been dissected away to display the notochord.

Attributions

• Phylum Echinodermata
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• Phylum Chordata
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