- Key People:
- Alexander Agassiz
Water-vascular system
The water-vascular system, which functions in the movement of tube feet, is a characteristic feature of echinoderms, and evidence of its existence has been found in even the oldest fossil forms. It comprises an internal hydraulic system of canals and reservoirs containing a watery fluid, the system consisting of a sieve plate, or madreporite, and a ring vessel, or water-vascular ring, that are connected by a frequently calcified vessel called the stone canal. Five radial water canals extend outward from the ring vessel and give rise to branches that end in the tube feet, which are in contact with the sea. The ring vessel in ophiuroids, asteroids, concentricycloids, and holothurians has bulbous cavities called Polian vesicles, which apparently maintain pressure in the system and hold reserve supplies of fluid; ophiuroids have four or more vesicles, asteroids five, holothurians from one to 50. Crinoids lack Polian vesicles, and echinoids have five structures known as either Polian vesicles or spongy bodies.
The madreporite, which is usually located externally, takes in water from outside the body; if internally located, as is the case in many holothurians, fluid is taken from the body cavity. The water or fluid passes from the madreporite to the ring vessel and along the radial canals to the tube feet. The tube feet are extended by contractions of localized muscle areas in the radial canals (ophiuroids) or by contractions of offshoots of the radial canals called ampullae (asteroids, concentricycloids, echinoids, and holothurians); the contractions force fluid into the tube feet, which then extend.
The structure of the system varies from group to group; asteroids frequently have more than one madreporite, and in holothurians, the madreporite is usually internal, hanging in the coelom. Radial canals may lie inward or outward from the skeleton. The tube feet may have well-developed suckers with great holding power, may taper to a point, or may be adapted for respiration, feeding, burrow building, mucus production, or sensory perception. Attachment of tube feet to hard substrates is achieved through a combination of suction and mucus production. The mucus contains adhesive and de-adhesive mucopolysaccharides. Respiratory tube feet have high oxygen uptake; they are usually located on parts of the body where water flow is unimpeded. Tube feet have been implicated in photoreception and chemoreception; the eyespots in the terminal tentacles of asteroids are the most conspicuous photoreceptors.
The tube feet of crinoids are arranged in clumps of three on the arms and on the pinnules. They secrete and spread a net of sticky mucus that traps small organisms. In ophiuroids the tube feet are used to gain a hold on a surface and to pass food to the mouth. The numerous tube feet of asteroids are used in locomotion; asteroids with suckered feet may use them to exert a continuous pull on the valves of shellfish (e.g., oysters, mussels) until muscles holding the valves tire and open slightly, allowing the asteroid to insert its stomach. In sea daisies the ring of tube feet is probably used for attachment to substrates. Holothurians use tube feet for the same purpose. Tentacles around the mouth of holothurians are modified tube feet used to capture food; tentacles used to capture plankton are branched and sticky, while those used to scoop mud and shovel it into the mouth have a simpler structure.
The tube feet of echinoids serve a variety of functions. The mouth of regular echinoids is surrounded by sensory tube feet, and tube feet farther from the mouth are used in locomotion. On the upper side of the body near the anus, the tube feet have respiratory and sensory functions. The tube feet of irregular echinoids, which burrow, are modified in various ways for feeding, burrow construction, and sensory and respiratory functions.
Body wall and body cavity
The outer body wall (epidermis) contains hairlike projections (cilia) in most echinoderms except ophiuroids; the body wall of crinoids has relatively few. The cilia produce a waving motion that carries food particles toward the mouth or removes unwanted particles from the body. The epidermis also contains glandular and sensory cells. The epidermis of skeletal elements such as spines and pedicellariae, which project from the body surface, often is worn away. The next layer, the dermis, includes the calcareous skeleton and connective tissues. Internal to the dermis are circular and longitudinal muscle layers. The extensive body cavity (coelom) is modified to form several specialized regions. Two subdivisions of the coelom are the perivisceral coelom and the water-vascular system. The perivisceral coelom is a large, fluid-filled cavity in which the major organs, particularly the digestive tube and sex organs, are suspended. Other regions of the coelom include the axial sinus (absent from adult holothurians and all echinoids), the madreporic vesicle, and the hyponeural sinus (often called the perihemal system).
Alimentary and blood systems
The digestive canal consists of a tube, which is almost straight (asteroids and ophiuroids), coiled in a clockwise direction (crinoids and holothurians), or coiled first clockwise, then counterclockwise (echinoids). The tube may be divided into esophagus, stomach, intestine, and rectum. Specialized branches of the digestive tube enlarge the digestive surface and may serve other functions; e.g., digestive glands of asteroids, diverticula of echinoids and crinoids, siphons in echinoids, and respiratory trees in holothurians. The anus, absent in ophiuroids and a few asteroids, is present in most groups. The mouth is near the centre of the oral surface, at the point of convergence of the areas containing the tube feet.
The blood system is a complex system of spaces that are neither part of the coelom nor true vessels. A hemal ring and five radial hemal canals surround the esophagus and radial canals of the water-vascular system. A sixth hemal space arises from the hemal ring and enters the axial organ. In addition, a complex network of hemal spaces is associated with the alimentary canal and gonads.
Axial organ
The axial organ, a complex and elongated mass of tissue found in all echinoderms except holothurians, represents the common junction of the perivisceral coelom, the water-vascular system, and the hemal system. Although its functions are not yet well understood, the axial organ plays a part in defense against invading organisms, can contract, is responsible for a circulation of fluids, and may have excretory and secretory activity.
Nervous system and sense organs
The echinoderm nervous system is complex. In all groups, a nerve plexus lies within and below the skin. In addition, the esophagus is surrounded by one to several nerve rings, from which run radial nerves often in parallel with branches of the water-vascular system. Ring and radial nerves coordinate righting activity.
Although echinoderms have few well-defined sense organs, they are sensitive to touch and to changes in light intensity, temperature, orientation, and the surrounding water. The tube feet, spines, pedicellariae, and skin respond to touch, and light-sensitive organs have been found in echinoids, holothurians, and asteroids.
Reproductive system
The masses of sex cells that compose the gonads of crinoids fill special cavities in the arms or pinnules. Crinoids are the only echinoderms with gonads outside the main body cavity, probably because its volume is reduced. Asteroids typically have 10 gonads, two in each arm, which are located near the arm base, appearing as a feathery tuft or a mass of tubules resembling a bunch of grapes. The gonads of some species are arranged in rows along each arm. Concentricycloids have five pairs of saclike gonads. Ophiuroids have gonads attached to sacs that hang into the body cavity; the sacs, which open outside the body at the bases of the arms, may have one gonad or as many as 1,000.
Regular echinoids typically have five gonads attached to the interambulacra. A duct from each gonad opens to the exterior near the anus. Most irregular echinoids have four gonads, some have three or five, a few have two; the ducts are on the upper surface of the body. Holothurians differ from all other living echinoderms in having a single gonad, which consists of branching or unbranched tubules; the tubules open into a single duct, which opens to the exterior near the front end of the body. Since many early fossil echinoderms have a single genital opening, or gonopore, it is assumed that these forms also had only one gonad; the condition in holothurians thus is regarded as primitive.