第28章無脊椎動物

Phylum Cnidaria includes animals that exhibit radial or biradial symmetry and are diploblastic, meaning that they develop from two embryonic layers, ectoderm and endoderm. Nearly all (about 99 percent) cnidarians are marine species.

Whereas the defining cell type for the sponges is the choanocyte, the defining cell type for the cnidarians is the cnidocyte, or stinging cell. These cells are located around the mouth and on the tentacles, and serve to capture prey or repel predators. Cnidocytes have large stinging organelles called nematocysts, which usually contain barbs at the base of a long coiled thread. The outer wall of the cell has a hairlike projection called a cnidocil, which is sensitive to tactile stimulation. If the cnidocils are touched, the hollow threads evert with enormous acceleration, approaching 40,000 times that of gravity. The microscopic threads then either entangle the prey or instantly penetrate the flesh of the prey or predator, releasing toxins (including neurotoxins and pore-forming toxins that can lead to cell lysis) into the target, thereby immobilizing it or paralyzing it (see Figure 28.5).

Two distinct body plans are found in Cnidarians: the polyp or tuliplike "stalk" form and the medusa or "bell" form. (Figure 28.6). An example of the polyp form is found in the genus Hydra, whereas the most typical form of medusa is found in the group called the “sea jellies” (jellyfish). Polyp forms are sessile as adults, with a single opening (the mouth/anus) to the digestive cavity facing up with tentacles surrounding it. Medusa forms are motile, with the mouth and tentacles hanging down from an umbrella-shaped bell.

Some cnidarians are dimorphic, that is, they exhibit both body plans during their life cycle. In these species, the polyp serves as the asexual phase, while the medusa serves as the sexual stage and produces gametes. However, both body forms are diploid.

An example of cnidarian dimorphism can be seen in the colonial hydroid Obelia. The sessile asexual colony has two types of polyps, shown in Figure 28.7. The first is the gastrozooid, which is adapted for capturing prey and feeding. In Obelia, all polyps are connected through a common digestive cavity called a coenosarc. The other type of polyp is the gonozooid, adapted for the asexual budding and the production of sexual medusae. The reproductive buds from the gonozooid break off and mature into free-swimming medusae, which are either male or female (dioecious). Each medusa has either several testes or several ovaries in which meiosis occurs to produce sperm or egg cells. Interestingly, the gamete-producing cells do not arise within the gonad itself, but migrate into it from the tissues in the gonozooid. This separate origin of gonad and gametes is common throughout the eumetazoa. The gametes are released into the surrounding water, and after fertilization, the zygote develops into a blastula, which soon develops into a ciliated, bilaterally symmetrical planula larva. The planula swims freely for a while, but eventually attaches to a substrate and becomes a single polyp, from which a new colony of polyps is formed by budding.

All cnidarians are diploblastic and thus have two “epithelial” layers in the body that are derived from the endoderm and ectoderm of the embryo. The outer layer (from ectoderm) is called the epidermis and lines the outside of the animal, whereas the inner layer (from endoderm) is called the gastrodermis and lines the digestive cavity. In the planula larva, a layer of ectoderm surrounds a solid mass of endoderm, but as the polyp develops, the digestive or gastrovascular cavity opens within the endoderm. A non-living, jelly-like mesoglea lies between these two epithelial layers. In terms of cellular complexity, cnidarians show the presence of differentiated cell types in each tissue layer, such as nerve cells, contractile epithelial cells, enzyme-secreting cells, and nutrient-absorbing cells, as well as the presence of intercellular connections. However, with a few notable exceptions such as statocysts and rhopalia (see below), the development of organs or organ systems is not advanced in this phylum.

The nervous system is rudimentary, with nerve cells organized in a network scattered across the body. This nerve net may show the presence of groups of cells that form nerve plexi (singular: plexus) or nerve cords. Organization of the nervous system in the motile medusa is more complex than that of the sessile polyp, with a nerve ring around the edge of the medusa bell that controls the action of the tentacles. Cnidarian nerve cells show mixed characteristics of motor and sensory neurons. The predominant signaling molecules in these primitive nervous systems are peptides, which perform both excitatory and inhibitory functions. Despite the simplicity of the nervous system, it is remarkable that it coordinates the complicated movement of the tentacles, the drawing of captured prey to the mouth, the digestion of food, and the expulsion of waste.

The gastrovascular cavity has only one opening that serves as both a mouth and an anus; this arrangement is called an incomplete digestive system. In the gastrovascular cavity, extracellular digestion occurs as food is taken into the gastrovascular cavity, enzymes are secreted into the cavity, and the cells lining the cavity absorb nutrients. However, some intracellular digestion also occurs. The gastrovascular cavity distributes nutrients throughout the body of the animal, with nutrients passing from the digestive cavity across the mesoglea to the epidermal cells. Thus, this cavity serves both digestive and circulatory functions.

Cnidarian cells exchange oxygen and carbon dioxide by diffusion between cells in the epidermis and water in the environment, and between cells in the gastrodermis and water in the gastrovascular cavity. The lack of a circulatory system to move dissolved gases limits the thickness of the body wall and necessitates a non-living mesoglea between the layers. In the cnidarians with a thicker mesoglea, a number of canals help to distribute both nutrients and gases. There is neither an excretory system nor organs, and nitrogenous wastes simply diffuse from the cells into the water outside the animal or into the gastrovascular cavity.

The phylum Cnidaria contains about 10,000 described species divided into two monophyletic clades: the Anthozoa and the Medusozoa. The Anthozoa include the corals, sea fans, sea whips, and the sea anemones. The Medusozoa include several classes of Cnidaria in two clades: The Hydrozoa include sessile forms, some medusoid forms, and swimming colonial forms like the Portuguese man-of-war. The other clade contains various types of jellies including both Scyphozoa and Cubozoa. The Anthozoa contain only sessile polyp forms, while the Medusozoa include species with both polyp and medusa forms in their life cycle.