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See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/227982250 Cnidaria (Coelenterates) Chapter · September 2005 DOI: 10.1038/npg.els.0004117 CITATIONS READS 0 6,314 1 author: Stanley Shostak University of Pittsburgh 88 PUBLICATIONS 459 CITATIONS SEE PROFILE All content following this page was uploaded by Stanley Shostak on 31 March 2015. The user has requested enhancement of the downloaded file. Cnidaria (Coelenterates) Introductory article Stanley Shostak, University of Pittsburgh, Pennsylvania, USA Article Contents . Basic Design Cnidaria (Gr., cnidae, nettle) is a phylum of mostly marine Metazoa distinguished by . Diversity cnidocysts, subcellular capsules containing an inverted tubule capable of everting and, in . Habitats and Abundance some cases, discharging venom. Sexual reproduction results in solid planula embryos that . Habits and Lifestyles develop into columnar polyps. Asexual reproduction results in clones of polyps, polypoidal . Life Histories colonies or disk- to bell-shaped sexual medusas (jellyfish). Oddities with the Phylum . Fossil History Basic Design . Phylogeny . Future Developments Cnidarians exist in either of the two adult forms (pheno- morphs) known as polyps and medusas and an embryonic doi: 10.1038/npg.els.0004117 or larval form known as a planula. Polyps are columnar and rest on a base, although the polyps of colonial varieties may protrude from a stalk or stolon. Medusas are cup- shaped jellyfish, varying from disk-like to bulbous. The body of polyps and medusas consists of a double-layered cellular wall enclosing a cavity known as the coelenteron (or gastrovascular cavity), opening by a mouth or stomodeum. Extracellular breakdown of food occurs in the coelenteron, and food particles are distributed through the coelenteron to the body, where they undergo intracel- lular digestion. The coelenteron serves as a hydrostatic skeleton when pressure is applied by muscle in the body wall and may provide a brooding chamber for planulas. Polyps and medusas have tentacles, which are hollow or solid projections of the body wall. Sets or multiples of four, five, six or eight tentacles, sometimes arranged in rings, surround the apical end of polyps and margin of medusas. Fertilized eggs develop into hollow (coelo-) or solid (ster- Figure 1 Cnidarian life cycles. Anthozoa (dashed circle on left) consists eo-) gastrulas that rapidly become solid, pear-shaped and exclusively of polyps, while Medusozoa (dashed circle on right) consists of both medusas and polyps. The sexual phase, in which eggs and flattened or rod-like planulas that develop into polyps. spermatozoa are produced, occurs in anthozoan polyps and medusozoan Cnidarians are divided into two subphyla, Anthozoa medusas or reduced medusoids attached to polyps. Polyps alone are and Medusozoa, on the basis of their polyp’s structure and produced by sexual reproduction (single arrow). Medusas are produced by the existence of a medusa stage (Figure 1). Anthozoans exist the asexual reproduction of polyps (double arrow). exclusively as polyps, while medusozoans alternate be- tween polyps and medusas. Medusas are made by polyps Scyphozoa, represented by massive, ocean-going medusas; through asexual reproduction, and the passage from polyp and Cubozoa, represented by small, box-like medusas with to medusa is known as the ‘alternation of generation’ (also a dangerous sting. called metagenesis and sometimes heterogenesis). Polyps Polyps and medusas appear radially symmetrical, but are the sexual phase of Anthozoa, while medusas are the anthozoans are biradially or bilaterally symmetrical. They sexual phase of Medusozoa, although medusas are fre- are biradial when the slit-like stomodeum and pharynx quently reduced to sexual medusoids attached to the body have ciliated grooves (siphonoglyphs or sulci) at both ends, wall of polyps and sometimes to mere gonads on polyps. and bilaterally symmetrical when a siphonoglyph appears In medusozoans, lips surrounding the mouth project at one end. Tentacles and pairs of mesenteries, or septa, beyond the body wall as a manubrium or hypostome. In linking the body wall to the pharynx and partitioning the anthozoans, lips are folded deeply into the coelenteron, coelenteron, are also biradially or bilaterally arranged. forming a pharynx that opens at the surface as a slit-like The cnidarian body wall consists of an outer epithelium stomodeum within an oral plate. The Medusozoa is further (ectoderm or epidermis), and an inner epithelium (end- divided into three classes: Hydrozoa, represented by en- oderm or gastrodermis) lining the coelenteron. A noncel- crusting or arborizing polyps and small, delicate medusas; lular layer, the mesoglea, is located between the two ENCYCLOPEDIA OF LIFE SCIENCES & 2005, John Wiley & Sons, Ltd. www.els.net 1 Cnidaria (Coelenterates) Figure 2 Light micrograph of section through wall of young sea anemone Figure 3 Phase-contrast micrograph of macerated Hydra viridis cells Bunodactis verrucosa at the origin of a mesentery. Two cellular layers are spread on a haemocytometer grid. Large endodermal cells containing separated by a dense mesoglea. refractive endosymbiotic algae have basal muscular extensions. Smaller cnidoblasts in nests of 2–8 cells contain differentiating cnidocysts. epithelial layers. In some large medusas and polyps, the mesoglea is infiltrated by cells that may even congregate in ectoderm (Hydrozoa) or endoderm (Anthozoa, Cubozoa, massive, muscular sphincters. The mesoglea in medusas is Scyphozoa) and in different body regions. distended by a buoyant gelatinous filler, 99% water. In Cnidocysts (or cnidae) exist in 10 common varieties and Anthozoa, the mesoglea is condensed into a leathery 20 or more rare varieties. Ever since the pioneering work of framework (Figure 2). Robert Weill in the 1930 s, cnidocysts have been classified The epithelia are simple, consisting of squamous to co- as adhesive, ensnaring and penetrating nematocysts and lumnar epitheliocytes or epitheliomuscular cells with an entangling spirocysts. Another cnidocyst, the ptychocyst, apical cilium or flagellum and a basal contractile muscle- builds the wall of tube anemones. Cnidarian species typ- like fibre organized into layers. In polyps, the ectoderm ically have 1–3 different kinds of cnidocysts (range 1–7) fibres may be longitudinally orientated, while endoderm comprising their cnidome or census of cnidocysts with fibres may be circularly or both longitudinally and circu- more complex cnidomes occurring among the med- larly orientated. Contraction of circular fibres applies usozoans. Cnidomes may be species-specific or, more of- pressure on the coelenteron, creating a hydrostatic skele- ten, subclass- or family-specific. ton. Coordinated contractions change the curvature of the Cnidocysts are produced in cells known as cnidoblasts. body and produce directional movement. In medusas, only These cells divide incompletely, forming small nests of ectodermal muscle is present and restricted to the concave, interconnected cells that differentiate synchronously oral surface where it is oriented circularly and radially. (Figure 3). The cells are known as cnidocytes following the In tentacles, the ectoderm differentiates as pavement and differentiation of their cnidocysts. Cnidocytes migrate to battery cells. Elsewhere, ectoderm forms adhesive and se- sites in tentacles and elsewhere, where they are able to cretory cells capable of secreting an extracellular, mucilag- discharge their cnidocyst (Figure 4). inous or skelatogenic matrix. Endodermal epitheliocytes are frequently populated by endosymbiotic algae (Figure 3). ‘Specialized cells’ are derived from stem or blast cells Diversity originating from amoeboid or interstitial cells insinuated among epitheliocytes. Pigment cells, mucous and other Cnidaria consist of about 9000 species. Subphylum Ant- gland cells, nerve and sensory cells, sex cells and cnidocytes hozoa has one class, also called Anthozoa, with 6100 spe- probably differentiate from unique stem cells. cies, while Subphylum Medusozoa has three classes: Nerve and sensory cells form the diffuse cnidarian Hydrozoa with 2700 species, Scyphozoa with 150 species, nervous system or nerve net. Its cells exhibit gap junctions and Cubozoa with 50 species. Classification within these similar to those in other animals, and nerves effect neuro- taxonomic categories relies on morphological characteris- muscular facilitation and spontaneous rhythmic discharge. tics, especially of sexual structures, and on cnidomes. In addition, cnidarians exhibit nonnervous conduction of See also: Classification excitatory impulses. Sensory cells in the vicinity of battery Anthozoans are polypoidal. Anemones are solitary; hard cells probably play a role in cnidocyst discharge. corals are colonial (Montastrea, star corals) and solitary Germ cells are diffusely distributed, since gonads are not (Fungia, mushroom corals); octocorallians are nearly all co- present continuously. Sex cells develop in specific layers, in lonial. The class is divided into two subclasses: Hexacorallia 2 Cnidaria (Coelenterates) running through an internal skeleton containing bony spicules. The Hexacorallia are usually clustered into two major orders and several minor ones. The solitary sea anemones comprise Actiniaria (1000 species), and true or stony corals constitute Scleractinia or Madreporaria (2500 species). Other orders include the coral-like Corallimorpharia and anemone-like Ptychodactiaria and Zoanthidea (300 spe-
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