第28章無脊椎動物

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.

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.

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.

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!

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.

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 subtropical 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.