PHYLUM ECHINODERMATA Introduction The echinoderms have a long evolutionary history and were once some of the most abundant animals in the ancient oceans; only 7,000 species are living today compared to 13,000 fossil species. This makes the Echinodermata another one of those animal groups that have a body plan that was more successful in the past than now. The ancestral echinoderms were sessile organisms attached to the substrate with their mouth pointed up and feeding on the organic material that fell through the water of the ancient oceans. Like other early deuterostomes, the feeding appendages surrounding the mouth were hollow extensions of the tripartite coelom. To trap even more food the ancestor of the group had arms that extended out from the central disc, and on the arms were tube feet that passed the captured food to a central disc where the mouth was located. Echinoderms are marine animals; the phylum has no freshwater or terrestrial representatives. They come in a variety of different shapes and sizes including: sea stars, sand dollars, sea urchins, brittle stars, sea cucumbers, and basket stars to name a few. They're usually found in shallow waters but sea lilies and sea cucumbers are also found in the abyssal depths of the oceans. One of the distinctive characteristics of the phylum is their pentaradiate symmetry. At one time biologists combined them with the radially symmetric cnidarians and named them the Radiata. But, once the bilaterally symmetric planktonic larval stage of echinoderms was identified, it became clear that radial symmetry in the adult was a derived trait associated with sessile suspension feeding. The first echinoderms lived attached to the ocean bottom with their mouths facing up. The tube feet on their arms used mucus to collect food floating to the ocean bottom or suspended in the ocean currents. Asteroidea Commonly called sea stars, Asteroidea display the typical pentaradiate symmetry of the phylum and have arms that gradually increase in diameter as they fuse with the central disc. There are approximately 1,600 species living in the shallower coastal waters of the world's oceans; like all echinoderms, they're only found in the marine environment. The size and number of arms that a sea star has varies. Some have the usual five, others more. If there are more, it's usually based on some multiple of five. Sea stars are usually between 12 and 24 centimeters in diameter, although some may become more than a meter across. Fig. 1. External anatomy of the aboral and oral surface of the sea star. © BIODIDAC Asteris External Anatomy Phylum Echinodermata -1- Digital Zoology LabManual © Houseman Examine the external features of your starfish identifying the aboral and oral surfaces with the mouth located in the centre of the peristomial membrane. Does a starfish have a dorsal and ventral surface? Five arms radiate from a central disc. The groove on the oral surface of each arm is the ambulacral groove and contains four rows of tube feet with sucker like discs located at their ends. The delicate tube feet are protected by the ambulacral groove and the spines found along its edge. In your specimen, the water vascular system has been injected with a blue or red dye and tube feet are easy to see in the ambulacral groove. At the end of each ambulacral groove is a minute eye spot formed from a pigmented epidermal invagination. This may or may not be visible depending on how the preservative and the injected dye have affected the pigments. Fig. 2. Aboral surface of the sea star. © BIODIDAC The aboral surface is composed of a central disc with the madreporite on one side of the disc, rather in the centre. Examine the madreporite under the dissecting microscope. How is its appearance related to its role in controlling the movement of water into the water vascular system? The anal opening is located in the approximate middle of the central disc. Why is the anal opening so small and how is this related to the way that starfish feed? Fig. 3. Structure of a sea star pedicellarium. © BIODIDAC The position of the madreporite is used to name and number the starfish regions and arms. The arms on each side of the madreporite are the bivium and the remaining three the trivium. Arm A is immediately opposite the madreporite and in the middle of the trivium and when viewed from above, each arm is lettered consecutively, counterclockwise, from A to E. Using this nomenclature the madreporite is located between what arms? Phylum Echinodermata -2- Digital Zoology Lab Manual © Houseman The whole aboral surface is covered with spines. Take a close look under the dissecting microscope to see the finger like sacs at the base of the spines. These are dermal branchia and are outpockets of the underlying coelom involved in excretion and respiration. They are lined internally with cilia that insure that the coelomic fluids are continually mixed so that the exchange can occur. What kind of material is being exchanged across this surface? The pedicellaria are small pincer-like structures located around the spines in a ring of tissue. Scrape away a portion of the epidermis to expose the calcareous ossicles that form the skeleton. Place your scrapings on a microscope slide and observe the pedicellaria. Compare yours to the prepared microscope slide of pedicellaria. Internal Anatomy If there is time you are more than welcome to take a preserved starfish and observe the internal anatomy. It’s really straightforward and you can just as easily see the main internal features on the preserved museum mount. The protocol that follows is for if you have the time. Fig. 4. Internal antomy of the sea star. © BIODIDAC Starting at the tip of each ray carefully lift the aboral body wall off the underlying tissue and as you do this gently free the tissue from the body wall. Be particularly careful when you remove the body wall over the central disk, especially near the madreporite, so that connections with the underlying water vascular system are not broken. Examine the internal surface of a piece of the body wall that you have removed so that you can see the fused calcareous ossicles that form the endoskeleton of the starfish. The part of the coelom that you have exposed is the pervisceral cavity and one of the three parts of the tripartite coelom. Which part is it? The pervisceral cavity is filled with fluid that circulates throughout the cavity by the cilia that line it. It was extensions of this cavity that formed the dermal branchia that you saw on the external surface. The movement of fluid in the pervisceral cavity, combined with the same in the water vascular system essentially defines the circulatory system of the animal. The water vascular system is also one of the three parts of the coelom; which is it? Digestive system Starfish feed on just about any organic material that they can find. The mouth, and an inverted stomach, are used to cover the surface being fed on. This creates an enclosed space into which the digestive enzymes are released. The organic material on the covered surface is broken down before being swept into the alimentary tract for final digestion by the cilia that line the system. The major structures of the digestive system are best seen by submersing your specimen under water. That way the different regions can float free of each other. A short esophagus leads from the mouth, and opens into a stomach divided into two regions, the pyloric and cardiac stomachs. A large cardiac portion everts during feeding and subsequently retracts back into the body cavity. The cardiac region connects to the smaller pyloric region that is in turn connected to the five paired pyloric ceca (digestive glands) contained in each arm. The pyloric ceca are the sites of final digestion and their position throughout the body insures that there is an Phylum Echinodermata -3- Digital Zoology LabManual © Houseman adequate supply of nutrients to the whole organism by nutrient diffusion in the coelomic fluid circulated by cilia. A rectal caecum is connected to the tube leading from the pyloric part of the stomach and the anus on the aboral surface. Why is the anal opening so small? Carefully remove the pyloric ceca to expose the paired gonads found in each of the arms. Their size will depend on the reproductive state of the animal when it was collected. You will not be able to see the external openings for the gonads in the interambulacral region. The sexes are difficult to distinguish using physical features. Can you suggest a way to determine the sex of your specimen? Fig. 5. Organization of the water ascular system inn the sea star. © BIODIDAC Water vascular system Remove the gonads and the remainder of the digestive system to expose the underlying water vascular system. Be careful when you remove this in the area near the madreporite. You don’t want to damage the madreporite’s connection to the rest of the water vascular system. The principle elements of the system include the madreporite, stone canal, and ring canal which are connected to the five radial canals and their associated short transverse canals. The tube feet are connected to the transverse canals. The ring canal has nine Tiedemann bodies and the position of the tenth is occupied by the connection of the stone canal. Polian vesicles that are present on some species of starfish are absent in the one that you are looking at today. The bulb-like ampulla of each tube foot is visible from the inner surface of the ambulacral groove as it passes between the ambulacral plates. How many ampullae are there on each transverse canal? The remaining parts of the tube feet are only visible when viewed from the oral surface. You will notice how the ambulacral spines protect them. Cut a section through the arm to see the relationship between the podia, ampulla of the tube feet and the ambulacral skeleton. Nervous and Circulatory systems The nervous system is composed primarily of a nerve net over the body surface. If you remove the tube feet from one of the ambulacral grooves a minute yellowish brown ridge running along the entire length of the groove may be apparent if you fold the ambulacral groove back. This is the radial nerve cord that connects with a circumoral nerve ring. The circulatory system is all but absent and its remnants form a hemal system closely associated with the first part of the tripartite coelom. You will not be able to see the system. Echinoidea There are approximately 900 species of echinoids, which come in two shapes. They are either spherical, regular echinoids, such as the sea urchins, or disk-shaped, irregular echinoids, such as the sand dollars. Like all echinoderms, the echinoids are exclusively marine animals and can be found anywhere from the intertidal zone to five kilometers deep. The pentaradiate symmetry of the phylum is still visible externally if you look closely at the surface of both. Sea urchins have spherical symmetry and the ambulacral zones alternate around the surface of their circular test, and the aboral region has been reduced to a set of plates that surround the anus. Sand dollars Phylum Echinodermata -4- Digital Zoology Lab Manual © Houseman have become secondarily bilaterally symmetric. The ambulacral zones are on the aboral side, and the mouth and anus are on what was the oral side. As you would expect with the solid shell, the body is not part of a hydrostatic skeleton, and there is no musculature associated with the body wall. Sea Urchin Sea urchins are either herbivores or detritivores grazing on large marine plants or the substrate, using the unique feeding structure of Aristotle’s lantern. External anatomy The outer body surface of a sea urchin is covered with a large variety of different spines, some of which articulate through a ball and socket type of joint. Submerse your specimen under water and use the dissecting microscope to examine the spines, pedicellaria, and the long tube feet that extend beyond the spines. Fig. 6. External anatomy of the aboral surface of a sea urchin. © BIODIDAC Use one of the dried sea urchin tests to see the interlocking hexagonal plates that make up the test. The plates are arranged into rows organized into alternating double columns of which five are ambulacral and five are interambulacral. The ambulacral region is easily identified by the perforations through which the tube feet extend. On the aboral surface a small distinct plate surrounds the anal opening on the aboral surface and the opening of the gonopore perforates Phylum Echinodermata -5- Digital Zoology LabManual © Houseman the five genital plates. The madreporite is found on the largest of the five genital plates. Use the dissecting microscope and look at one of the plates that make up the test. You should be able to see the tubercles; and the ball and socket joint that articulates the large surface spines. Fig. 7. External anatomy of the oral surface of the sea urchin © BIODIDAC Return to your preserved specimen in the dissecting dish and examine the oral surface. In the centre you can see the five white protruding teeth of Aristotle’s lantern. The membranous peristome surrounds the lantern. The feeding structure in a sea urchin is Aristotle’s lantern and is an intricate series of interlocking plates. Observe the specimen available on demonstration. We will remove Aristotle’s lantern for a closer look after we have looked at the internal anatomy. Fig. 8. Detail of the anal plates (left) and the spines and pedicellaria on the surface of the sea urchin. © BIODIDAC Internal Anatomy Scrape the surface of the your urchin to remove the spines from the sea urchin test and submerse your specimen under water in a finger bowl. With scissors carefully cut through the equatorial region of the sea urchin and separate the two halves in much the same way that you would crack an egg. As you separate the two halves be sure to keep them submersed so that none of the internal tissues are damaged. Orient the two halves as shown in the museum mount taking care Phylum Echinodermata -6- Digital Zoology Lab Manual © Houseman to not rip the organs that extend between the two hemispheres of the animal. The complex framework supporting the teeth is Aristotle’s lantern and is composed of muscles and calcareous plates. Fig. 9. Internal anatomy of the sea urchin. © BIODIDAC The esophagus extends from the centre of Aristotle’s lantern and connects with the flat stomach which runs around the edge of the body cavity followed by the intestine. The digestive system winds back on itself before moving into the aboral hemisphere containing the rectum and anus. Because of the way that they feed, sea urchins consume a large volume of water along with the food and a second part of the alimentary tract, the siphon running along the inner margin of the looping stomach. It bypasses the stomach and allows water to be removed from the ingested food so that it becomes concentrated. There are no digestive ceca and the looping passage of the intestine through the test insures the distribution of nutrients. Phylum Echinodermata -7- Digital Zoology LabManual © Houseman You may not be able to see it, but the water vascular ring extends around the oral surface and spongy Polian vesicles are attached to it. Identify the ampullae of the tube feet. Gonads are also apparent in the upper hemisphere. Fig. 10. Ossicles and muscles of Aristotle’s lantern in the sea urchin. © BIODIDAC Carefully remove Aristotle’s lantern and examine the arrangement of the musculature and the plates and teeth. How does this work? Sand dollar The sand dollar is an example of another echinoid type and compared to the sea urchin the modifications are dramatic. The echinoid body plan has been compressed in the oral aboral plane and these animals are specialized for burrowing. Fig. 11. External anatomy of the aboral surface of the sand dollar. © BIODIDAC The surface is covered with short spines that give a velvety appearance to the body. Look at the oral and aboral surface through the dissecting microscope. On the aboral surface the ambulacral area are visible as petaloids and in these animals the tube feet that extend through here are involved in gas exchange. The five openings to the gonopore surround the madreporite. The mouth and the anus are both on the oral surface and the feeding mechanism is not fully Phylum Echinodermata -8- Digital Zoology Lab Manual © Houseman understood. The openings in the sand dollar shell are lunules and it is not known whether they are involved in feeding allowing food to pass to the aboral surface or whether they help stabilize the animals where there are strong currents. Holothuroidea The sea cucumber is an example of the Holothuroidea and is an interesting class because many of the ancestral echinoderm characters have been modified with a return to a bilateral type of symmetry. External Anatomy The endoskeleton has been reduced to spicules buried in the leathery body of the animal. The tube feet and the ambulacral grooves are still visible on the outside and you can still tell the ambulacral from the interambulacral parts of the body. But, because these animals always orient one side of their body against the substrate the tube feet on this side are much more visible as they are still involved in locomotion, just like other Echinoderms. The tube feet on the opposite side, which are not in contact with the substrate are much smaller or may have disappeared completely depending on the species. The tube feet aren’t the only way that these animals move and contractions of the muscles embedded in the body wall allows sea cucumbers to wriggle across the substrate. The next most obvious external feature are the tentacles surrounding the mouth and used for deposit feeding typical of the class. Food is gathered on the tentacles and the whole tentacle is pushed into the mouth and wiped clean of food before being extended back outside the animal. Look at the opposite end of the animal and you’ll see another opening commonly referred to as a vent. That’s because water is pumped in and out of the opening to aerate the unique respiratory trees that these animals use for gas exchange. Fig. 12. Internal anatomy of the sea cucumber. © BIODIDAC Internal Anatomy Make a cut from the base of the tentacles to the anal opening (vent or cloaca) and pin back the body wall. Inside is a mess of organs and structures and to see them you’ll need to flood the specimen and gently tease the material apart. The cavity that you opened is the enterocoelom of the animal and the walls are lined with cilia that constantly mix the fluid content. The mouth leads to a pharyngeal area that can be seen as an enlargement of the digestive tract at the base of the mouth. There are a number of structures that can be identified in this area. The five retractor muscles are attached to a calciferous ring. Take a good look at this structure under your dissecting microscope and you’ll see the ring canal behind the calciferous ring. This is part of the water vascular system and the stone canal, and the madreporite attached to it. The polian vesicles are also connected to the ring canal and they vary in size depending on the species. In ours they are large. The rest of the water vascular system may be hard to see, but if Phylum Echinodermata -9- Digital Zoology LabManual © Houseman you look closely, five radial canals are connected to the ring and follow the large longitudinal muscles embedded in the body wall. Don’t confuse these muscles with the tentacle retractor muscles that you looked at earlier. The ring canal branches as it passes to the posterior of the animal and ultimately connecting to the tube feet. As we mentioned earlier, tube feet aren’t the only means of locomotion that these animals have and in addition to the longitudinal muscles in the body there are circular muscles as well. Fig. 13. Detail of the main internal systems of the sea cucumber. © BIODIDAC These animals are particle or suspension feeders and food collected by the tentacles is passed into the mouth down a short esophagus and into the stomach. From here the food is passed to the intestine that loops backwards and forwards through the body cavity. The intestine ends at the vent. Two large respiratory trees are connected at the vent and water is pumped into the respiratory tree by the musculature associated with the vent. At the base of the respiratory tree are the Cuvierian organs which are filled with sticky mucous threads. These are used in defence and when sea cucumbers are disturbed they release the threads. If a severe defensive reaction is required then sea cucumbers eviscerate expelling part, or all of the digestive tract and the respiratory trees. A single gonad is present and gametes are released through a genital pore positioned at the base of the tentacle. Internally the gonad looks like an old mop and the bigger the gonad is the more mature the specimen is. Just like all echinoderms its hard to tell the sex of your specimen. Echinoderm Diversity Identify the major features of animals representing the other classes in Echinodermata. The feather stars and sea lilies, Class Crinoidea, are the most primitive of the Echinoderms. Distinguish between the aboral and oral surfaces. The oral surface has both the mouth and anal openings, the latter projects away from the mouth by being positioned at the tip of an anal papillae. Cilia in the ambulacral grooves are used to move food towards the mouth. Dermal vesicles which may appear as yellowish brown dots along the sides of the ambulacral grooves Phylum Echinodermata - 10 - Digital Zoology Lab Manual © Houseman have an unknown function. The opening to the water vascular system is through a single central aboral ossicle. Members of this class often possess a stem composed of a stack of disc shaped ossicles containing a central axial canal which terminates in a holdfast. Fig. 14. External anatomy of a crinoid. © BIODIDAC The class Ophiuroidea includes the brittle stars that are superficially similar to the Asteroidea, and get their name from the way they drop their arms off when disturbed. They differ from the starfish because of the distinct separation of the central disc and the radial arms. These animals lack ambulacral grooves and the arms are completely flexible along their length. Calcareous vertebrae like structures running the length of the arm facilitate this movement. Examine an arm that has been broken off and attempt to locate vertebrae and four sets of muscles contained inside. Fig. 15. External anatomy of the oral surface of a brittle star. © BIODIDAC Some interesting things that you might want to make note of as you work with your specimen is the location of the madreporite on the oral rather than aboral surface of the animal. While you’re looking at this surface make a note of the bursal slits that open into an enclosed space called the bursa. Seawater moves in and out for gas exchange and can also be used to protect developing larval young. Phylum Echinodermata - 11 - Digital Zoology LabManual © Houseman