第28章無脊椎動物

The phyla Echinodermata and Chordata (the phylum that includes humans) both belong to the superphylum Deuterostomia. Recall that protostomes and deuterostomes differ in certain aspects of their embryonic development, and they are named based on which opening of the archenteron (primitive gut tube) develops first. The word deuterostome comes from the Greek word meaning “mouth second,” indicating that the mouth develops as a secondary structure opposite the location of the blastopore, which becomes the anus. In protostomes (“mouth first”), the first embryonic opening becomes the mouth, and the second opening becomes the anus.

There are a series of other developmental characteristics that differ between protostomes and deuterostomes, including the type of early cleavage (embryonic cell division) and the mode of formation of the coelom of the embryo: Protosomes typically exhibit spiral mosaic cleavage whereas deuterostomes exhibit radial regulative cleavage. In deuterostomes, the endodermal lining of the archenteron usually forms buds called coelomic pouches that expand and ultimately obliterate the embryonic blastocoel (the cavity within the blastula and early gastrula) to become the embryonic mesoderm, the third germ layer. This happens when the mesodermal pouches become separated from the invaginating endodermal layer forming the archenteron, then expand and fuse to form the coelomic cavity. The resulting coelom is termed an enterocoelom. The archenteron develops into the alimentary canal, and a mouth opening is formed by invagination of ectoderm at the pole opposite the blastopore of the gastrula. The blastopore forms the anus of the alimentary system in the juvenile and adult forms. Cleavage in most deuterostomes is also indeterminant, meaning that the developmental fates of early embryonic cells are not decided at that point of embryonic development (this is why we could potentially clone most deuterostomes, including ourselves).

The deuterostomes consist of two major clades—the Chordata and the Ambulacraria. The Chordata include the vertebrates and two invertebrate subphyla, the urochordates and the cephalochordates. The Ambulacraria include the echinoderms and the hemichordates, which were once considered to be a chordate subphylum (Figure 28.46). The two clades, in addition to being deuterostomes, have some other interesting features in common. As we have seen, the vast majority of invertebrate animals do not possess a defined bony vertebral endoskeleton, or a bony cranium. However, one of the most ancestral groups of deuterostome invertebrates, the Echinodermata, do produce tiny skeletal “bones” called ossicles that make up a true endoskeleton, or internal skeleton, covered by an epidermis. The Hemichordata (acorn worms and pterobranchs) will not be covered here, but share with the echinoderms a three-part (tripartite) coelom, similar larval forms, and a derived metanephridium that rids the animals of nitrogenous wastes. They also share pharyngeal slits with the chordates (Figure 28.46). In addition, hemichordates have a dorsal nerve cord in the midline of the epidermis, but lack a neural tube, a true notochord and the endostyle and post-anal tail characteristic of chordates.

Echinodermata are named after their “prickly skin” (from the Greek “echinos” meaning “prickly” and “dermos” meaning “skin”). This phylum is a collection of about 7,000 described living species of exclusively marine, bottom-dwelling organisms. Sea stars (Figure 28.47), sea cucumbers, sea urchins, sand dollars, and brittle stars are all examples of echinoderms.

Despite the adaptive value of bilaterality for most free-living cephalized animals, adult echinoderms exhibit pentaradial symmetry (with “arms” typically arrayed in multiples of five around a central axis). Echinoderms have an endoskeleton made of calcareous ossicles (small bony plates), covered by the epidermis. For this reason, it is an endoskeleton like our own, not an exoskeleton like that of arthropods. The ossicles may be fused together, embedded separately in the connective tissue of the dermis, or be reduced to minute spicules of bone as in sea cucumbers. The spines for which the echinoderms are named are connected to some of the plates. The spines may be moved by small muscles, but they can also be locked into place for defense. In some species, the spines are surrounded by tiny stalked claws called pedicellaria, which help keep the animal's surface clean of debris, protect papulae used in respiration, and sometimes aid in food capture.

The endoskeleton is produced by dermal cells, which also produce several kinds of pigments, imparting vivid colors to these animals. In sea stars, fingerlike projections (papillae) of dermal tissue extend through the endoskeleton and function as gills. Some cells are glandular, and may produce toxins. Each arm or section of the animal contains several different structures: for example, digestive glands, gonads, and the tube feet that are unique to the echinoderms. In echinoderms like sea stars, every arm bears two rows of tube feet on the oral side, running along an external ambulacral groove. These tube feet assist in locomotion, feeding, and chemical sensations, as well as serve to attach some species to the substratum.