- Key People:
- Alexander Agassiz
Asexual reproduction in echinoderms usually involves the division of the body into two or more parts (fragmentation) and the regeneration of missing body parts. Fragmentation is a common method of reproduction used by some species of asteroids, ophiuroids, and holothurians, and in some of these species sexual reproduction is not known to occur. Successful fragmentation and regeneration require a body wall that can be torn and an ability to seal resultant wounds. In some asteroids fragmentation occurs when two groups of arms pull in opposite directions, thereby tearing the animal into two pieces. Successful regeneration requires that certain body parts be present in the lost pieces; for example, many asteroids and ophiuroids can regenerate a lost portion only if some part of the disk is present. In sea cucumbers, which divide transversely, considerable reorganization of tissues occurs in both regenerating parts.
The ability to regenerate, or regrow, lost or destroyed parts is well developed in echinoderms, especially sea lilies, starfishes, and brittle stars, all of which can regenerate new arms if existing ones are broken off. Echinoderm regeneration frustrated early attempts to keep starfishes from destroying oyster beds; when captured starfishes were chopped into pieces and thrown back into the sea, they actually increased in numbers. So long as a portion of a body, or disk, remained associated with an arm, new starfishes regenerated. Some sea cucumbers can expel their internal organs (autoeviscerate) under certain conditions (i.e., if attacked, if the environment is unfavourable, or on a seasonal basis), and a new set of internal organs regenerates within several weeks. Sea urchins (Echinoidea) readily regenerate lost spines, pincerlike organs called pedicellariae, and small areas of the internal skeleton, or test.
Sexual reproduction
In sexual reproduction, eggs (up to several million) from females and spermatozoa from males are shed into the water (spawning), where the eggs are fertilized. Most echinoderms spawn on an annual cycle, with the spawning period normally lasting one or two months during spring or summer; several species, however, are capable of spawning throughout the year. Spawn-inducing factors are complex and may include external influences such as temperature, light, or salinity of the water. In the case of one Japanese feather star (Crinoidea), spawning is correlated with phases of the Moon and takes place during early October when the Moon is in the first or last quarter. Many echinoderms aggregate before spawning, thus increasing the probability of fertilization of eggs. Some also display a characteristic behaviour during the spawning process; some asteroids and ophiuroids raise the centre of the body off the seafloor; holothurians may raise the front end of the body and wave it about. These movements are presumably intended to prevent eggs and sperm from becoming entrapped in the sediment.
Development
After an egg is fertilized, the development of the resulting embryo into a juvenile echinoderm may proceed in a variety of ways. Small eggs without much yolk develop into free-swimming larvae that become part of the plankton, actively feeding on small organisms until they transform, or metamorphose, into juvenile echinoderms and begin life on the seafloor. Larger eggs with greater amounts of yolk may develop into a larval form that is planktonic but subsists upon its own yolk material, rather than feeding upon small organisms, before eventually transforming into a juvenile echinoderm. Development involving an egg, planktonic larval stages, and a juvenile form is termed indirect development. Echinoderm development in which large eggs with abundant yolk transform into juvenile echinoderms without passing through a larval stage is termed direct development.
In direct development the young usually are reared by the female parent. Parental care or brood protection ranges from actual retention of young inside the body of the female until they are born as juveniles to retention of the young on the outer surface of the body. Brood protection is best developed among Antarctic, Arctic, and deep-sea echinoderms, in which young may be held around the mouth or on the underside of the parent’s body, as in some starfishes and sea cucumbers, or in special pouches on the upper surface of the body, as in some sea urchins, sea cucumbers, and asteroids.
During indirect development, the fertilized egg divides many times to produce a hollow ciliated ball of cells (blastula); cleavage is total, indeterminate, and radical. The blastula invaginates at one end to form a primitive gut, and the cells continue to divide to form a double-layered embryo called the gastrula. Echinoderms resemble vertebrates and some invertebrate groups (chaetognaths and hemichordates) in being deuterostomes; the hole through which the gut opens to the outside (blastopore) marks the position of the future anus; the mouth arises anew at the opposite end of the body from the blastopore. A pair of subdivided hollow pouches arise from the gut and develop into the body cavity (coelom) and water-vascular system.
The gastrula develops into a basic larval type called a dipleurula larva, characterized by bilateral symmetry; hence the name, which means “little two sides.” A single band of hairlike projections, or cilia, is found on each side of the body and in front of the mouth and anus. The characteristic larvae found among the living classes of echinoderms are modifications of the basic dipleurula pattern.
Because the ciliated band of the dipleurula larva of holothurians becomes sinuous and lobed, thus resembling a human ear, the larva is known as an auricularia larva. The dipleurula larva of asteroids develops into a bipinnaria larva with two ciliated bands, which also may become sinuous and form lobes or arms; one band lies in front of the mouth, the other behind it and around the edge of the body. In most asteroids the larval form in the next stage of development is called a brachiolaria, which has three additional arms used for attaching the larva to the seafloor. Echinoids and ophiuroids have complex advanced larvae closely similar in type. The larva, named pluteus, resembles an artist’s easel turned upside down. It has fragile arms formed by lobes of ciliated bands and is supported by fragile rods of calcite, the skeletal material. The echinoid larva (echinopluteus) and the ophiuroid larva (ophiopluteus) usually have four pairs of arms but may have fewer or more. An extra unpaired arm on the plutei of sand dollars and cake urchins extends downward, presumably to help keep the larva upright. The crinoids, which apparently lack a dipleurula larval stage, have a barrel-shaped larva called a doliolaria larva. The doliolaria larva also occurs in other groups; in holothurians, for example, it is the developmental stage after the auricularia larva, which may not occur in some species. A doliolaria larva usually contains large quantities of yolk material and moves with the aid of several ciliated bands arranged in hoops around the body.
Although most larval stages are small, often less than 1 mm (0.04 inch) in length, some holothurians are known to be 15 mm (0.59 inch) long in the larval stage, and the length of bipinnaria larvae of some starfishes may exceed 25 mm (0.98 inch).
After a few days to several weeks in a free-swimming form (plankton), echinoderm larvae undergo a complex transformation, or metamorphosis, that results in the juvenile echinoderm. During metamorphosis, the fundamental bilateral symmetry is overshadowed by a radial symmetry dominated by formation of five water-vascular canals (see below Form and function of external features). Among holothurians, echinoids, and ophiuroids, the larvae may metamorphose as they float, and the young then sink to the seafloor; among crinoids and asteroids, however, the larvae firmly attach to the seafloor prior to metamorphosis. The average life span of echinoderms is about four years, and some species may live as long as eight or 10.
Life activities
Food and feeding habits
Echinoderms feed in a variety of ways. A distinct feeding rhythm frequently occurs, with many forms feeding only at night, others feeding continuously. Feeding habits range from active, selective predation to omnivorous scavenging or nonselective mud swallowing.
Crinoids are suspension feeders, capturing planktonic organisms in a network of mucus produced by soft appendages, called tube feet, contained in grooves on the tentacles, or arms. The arms are spread into a characteristic “fan” at right angles to the prevailing current, and small prey animals are passed to the mouth along the grooves by activity of the cilia and the tube feet.
Many asteroids are active predators on shellfishes and even upon other starfishes; other asteroids are mud swallowers. When feeding, some asteroid species extrude their stomach through the mouth onto the prey, which then is partially digested externally, after which the stomach is retracted and digestion is completed inside the body. Most ophiuroids feed on small organisms floating in the water or lying on the bottom, which are captured by the arms and tube feet and passed toward the mouth. Ophiuroids with arms branched in a complex manner may feed in a way similar to that of the crinoids. Feeding methods of concentricycloids are not yet known.
The more primitive, so-called regular, sea urchins are omnivorous or vegetarian browsers, either scraping algae and other small organisms from rocks with their hard teeth or eating seaweed. Several deep-sea regular echinoids feed exclusively on plants carried into the sea from the land. The more advanced irregular echinoids, which usually lack teeth, are burrowers and pass small organisms to the mouth with the aid of spines and tube feet. Several species of sand dollars sometimes feed on suspended organisms carried to them by ocean currents as they lie on the seafloor. Some sea cucumbers remain attached to a surface for indefinite periods of time, capturing plankton in a network of branching, sticky tentacles; others select food from the seafloor and push it into their mouths with their tentacles. A large number of holothurians feed by actively swallowing mud and sand, digesting the organic material, and egesting the waste in the form of characteristic castings, in a manner similar to that of earthworms.
Under artificial conditions, as in aquariums, echinoderms can survive apparent starvation for several weeks at a time. After a holothurian has autoeviscerated, it is unable to feed during the several weeks required for gut regeneration. Echinoderms may derive a significant amount of nourishment, at least for the outer cell layers of the body, from organic material dissolved in seawater.