第28章無脊椎動物

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.

We will start our investigation with the simplest of all the invertebrates—animals sometimes classified within the clade Parazoa (“beside the animals”). This clade currently includes only the phylum Placozoa (containing a single species, Trichoplax adhaerens), and the phylum Porifera, containing the more familiar sponges (Figure 28.2). The split between the Parazoa and the Eumetazoa (all animal clades above Parazoa) likely took place over a billion years ago.

We should reiterate here that the Porifera do not possess “true” tissues that are embryologically homologous to those of all other derived animal groups such as the insects and mammals. This is because they do not create a true gastrula during embryogenesis, and as a result do not produce a true endoderm or ectoderm. But even though they are not considered to have true tissues, they do have specialized cells that perform specific functions like tissues (for example, the external “pinacoderm” of a sponge acts like our epidermis). Thus, functionally, the poriferans can be said to have tissues; however, these tissues are likely not embryologically homologous to our own.

Sponge larvae (e.g., parenchymula and amphiblastula) are flagellated and able to swim; however, adults are non-motile and spend their life attached to a substratum. Since water is vital to sponges for feeding, excretion, and gas exchange, their body structure facilitates the movement of water through the sponge. Various canals, chambers, and cavities enable water to move through the sponge to allow the exchange of food and waste as well as the exchange of gases to nearly all body cells.

There are at least 5,000 named species of sponges, likely with thousands more yet to be classified. The morphology of the simplest sponges takes the shape of an irregular cylinder with a large central cavity, the spongocoel, occupying the inside of the cylinder (Figure 28.3). Water enters into the spongocoel through numerous pores, or ostia, that create openings in the body wall. Water entering the spongocoel is expelled via a large common opening called the osculum. However, we should note that sponges exhibit a range of diversity in body forms, including variations in the size and shape of the spongocoel, as well as the number and arrangement of feeding chambers within the body wall. In some sponges, multiple feeding chambers open off of a central spongocoel and in others, several feeding chambers connecting to one another may lie between the entry pores and the spongocoel.

While sponges do not exhibit true tissue-layer organization, they do have a number of functional “tissues” composed of different cell types specialized for distinct functions. For example, epithelial-like cells called pinacocytes form the outermost body, called a pinacoderm, that serves a protective function similar that of our epidermis. Scattered among the pinacoderm are the ostia that allow entry of water into the body of the sponge. These pores have given the sponges their phylum name Porifera—pore-bearers. In some sponges, ostia are formed by porocytes, single tube-shaped cells that act as valves to regulate the flow of water into the spongocoel. In other sponges, ostia are formed by folds in the body wall of the sponge. Between the outer layer and the feeding chambers of the sponge is a jelly-like substance called the mesohyl, which contains collagenous fibers. Various cell types reside within the mesohyl, including amoebocytes, the “stem cells” of sponges, and sclerocytes, which produce skeletal materials. The gel-like consistency of mesohyl acts like an endoskeleton and maintains the tubular morphology of sponges.

The feeding chambers inside the sponge are lined by choanocytes ("collar cells"). The structure of a choanocyte is critical to its function, which is to generate a directed water current through the sponge and to trap and ingest microscopic food particles by phagocytosis. These feeding cells are similar in appearance to unicellular choanoflagellates (Protista). This similarity suggests that sponges and choanoflagellates are closely related and likely share common ancestry. The body of the choanocyte is embedded in mesohyl and contains all the organelles required for normal cell function. Protruding into the “open space” inside the feeding chamber is a mesh-like collar composed of microvilli with a single flagellum in the center of the column. The beating of the flagella from all choanocytes draws water into the sponge through the numerous ostia, into the spaces lined by choanocytes, and eventually out through the osculum (or osculi, if the sponge consists of a colony of attached sponges). Food particles, including waterborne bacteria and unicellular organisms such as algae and various animal-like protists, are trapped by the sieve-like collar of the choanocytes, slide down toward the body of the cell, and are ingested by phagocytosis. Choanocytes also serve another surprising function: They can differentiate into sperm for sexual reproduction, at which time they become dislodged from the mesohyl and leave the sponge with expelled water through the osculum.

The amoebocytes (derived from stem-cell-like archaeocytes), are so named because they move throughout the mesohyl in an amoeba-like fashion. They have a variety of functions: In addition to delivering nutrients from choanocytes to other cells within the sponge, they also give rise to eggs for sexual reproduction. (The eggs remain in the mesohyl, whereas the sperm cells are released into the water.) The amoebocytes can differentiate into other cell types of the sponge, such as collenocytes and lophocytes, which produce the collagen-like protein that support the mesohyl. Amoebocytes can also give rise to sclerocytes, which produce spicules (skeletal spikes of silica or calcium carbonate) in some sponges, and spongocytes, which produce the protein spongin in the majority of sponges. These different cell types in sponges are shown in Figure 28.3.

As we’ve seen, most sponges are supported by small bone-like spicules (usually tiny pointed structures made of calcium carbonate or silica) in the mesohyl. Spicules provide support for the body of the sponge, and may also deter predation. The presence and composition of spicules form the basis for differentiating three of the four classes of sponges (Figure 28.4). Sponges in class Calcarea produce calcium carbonate spicules and no spongin; those in class Hexactinellida produce six-rayed siliceous (glassy) spicules and no spongin; and those in class Demospongia contain spongin and may or may not have spicules; if present, those spicules are siliceous. Sponges in this last class have been used as bath sponges. Spicules are most conspicuously present in the glass sponges, class Hexactinellida. Some of the spicules may attain gigantic proportions. For example, relative to typical glass sponge spicules, whose size generally ranges from 3 to 10 mm, some of the basal spicules of the hexactinellid Monorhaphis chuni are enormous and grow up to 3 meters long! The glass sponges are also unusual in that most of their body cells are fused together to form a multinucleate syncytium. Because their cells are interconnected in this way, the hexactinellid sponges have no mesohyl. A fourth class of sponges, the Sclerospongiae, was described from species discovered in underwater tunnels. These are also called coralline sponges after their multilayered calcium carbonate skeletons. Dating based on the rate of deposition of the skeletal layers suggests that some of these sponges are hundreds of years old.