radial canal velum feeding polyps
Hydroid anatomy. A. Colony; B. Medusa. (Illustration by Patricia Ferrer)
tentacle exumbrella manubrium endodermal lamella ring canal velum
Hydroid anatomy. A. Colony; B. Medusa. (Illustration by Patricia Ferrer)
and are inappropriately called "actinulae," like the juvenile polyps of some Anthomedusae. The Trachymedusae have a bell-shaped umbrella, with circular and radial canals. Gametes ripen on the radial canals. The manubrium is often on a peduncle.
The Hydroidomedusa class is represented by a succession of three stages during indirect development. The planula is a ciliated motile gastrula; it typically develops into a benthic, modular, larval stage, polyp or hydroid (except in the Porpiti-dae, Margelopsis and Pelagohydra, where the hydroid is floating). Hydroids can be solitary, but generally form modular colonies by simple budding. Polyps can be specialized for different functions (defensive dactylozooids, reproductive gonozooids, nutritive gastrozooids, etc.). Polyps give rise, by asexual budding, to planktonic, free-swimming, and solitary hydromedusae, representing the sexual adult. The sense organs of pelagic hy-droidomedusae, when present, are ocelli (Anthomedusae, some Leptomedusae), or statocysts (Leptomedusae, Limnomedusae); sometimes cordyli of unknown function are also present (Lep tomedusae). The Siphonophores have no visible sense organs. Medusae are often reduced to sporosacs (fixed gonophores), so that hydroids, by paedomorphosis, secondarily become the sexual stages. The Hydroidomedusa may also form pelagic, highly polymorphic colonies (Siphonophores). Medusa budding occurs via a medusary nodule or entocodon, forming a coelom-like cavity, the subumbrellar cavity, lined by striated muscle cells; primary marginal tentacles always develop after the sub-umbrellar cavity and the gastro vascular system. Both embryonic and larval stages, the planula and the polyp, are typically diploblastic; the adult sexual stages, the hydromedusae, acquire a triploblastic kind of organization during embryonic development (medusary nodule formation). Hydroidomedusae are frequently seasonal; the hydroid stage may develop several types of resting stages (frustules, propagules, cysts, stolon system) to overcome unfavorable ecological conditions.
The hydroids of the subclass Anthomedusae do not have a protective perisarc sheath around the polyps and are said athecates; they are usually colonial (but the most famous hy-
droid, Hydra, is a solitary paedomorphic anthomedusa). The colonies can be either monomorphic or polymorphic, while the structure of the tentacles is in two forms: filiform tentacles do not present particular aggregations of cnidocysts, whereas capitate tentacles have cnidocyst knobs. The medusae are typically bell shaped; their gonads (aggregates of gametes commonly referred to as gonads) are confined on manubrium, sometimes extending on the most proximal parts of the radial canals. Their marginal sense organs, if present, are ocelli; the marginal tentacles are peripheral, hollow, or solid, usually with tentacular bulbs; sexual reproduction occurs through a complex planula. The hydroids of the subclass Laingiome-dusae are unknown. The medusae have an almost hemispherical umbrella, with lobed margin, divided by peronial grooves or similar structures. The radial canals are four; there is no typical circular canal but a solid core of endodermal cells around the umbrella margin. The tentacles are solid, inserted above the exumbrellar margin, on the exumbrella. The manubrium is simple, quadrangular, tubular, or conical; the mouth opening is simple, quadrangular to circular; gametes are in four masses on the manubrium or as epidermal lining of interradial pockets of the manubrium. The hydroids of the subclass Leptomedusae are thecate: all parts of the colonies are typically protected by a chitinous perisarcal structure. The hydranth is protected by a hydrotheca, the nematophore by a nematotheca, and the gonophore by a gonotheca. Rarely, hydranths are naked. The medusae are typically with hemispherical or flattened umbrella; the masses of gametes are confined to radial canals, exceptionally extending onto the proximal part of manubrium; when present, the marginal sense organs are ectodermal velar statocysts, rarely cordyli, occasionally adaxial ocelli. The marginal tentacles are peripheral and hollow (except in Obelia), with tentacular bulbs. Sexual reproduction occurs through a complex planula. The hydroids of the subclass Limnomedusae are very simple; solitary or colonial; small, sessile; with or without tentacles; often close to planula structure and budding planula-like structures or frustules; there are no perisarcal thecae, but cysts and stolons are covered by chitin. The medusae usually have gamete masses along the radial canals or, exceptionally, on the manubrium. The marginal tentacles are peripheral, hollow, without a true basal bulb; their base is usually with a parenchymatic endodermal core embedded in umbrellar mesoglea. The marginal sense organs are internal, enclosed ecto-endodermal statocysts that are embedded in the mesoglea near the ring canal or in the velum. Exceptionally, medusae can be reduced medusoids. Sexual reproduction leads to simple planulae, without embryonic glandular cells.
The subclass Siphonophorae comprises generally pelagic, free-swimming, or floating species, forming highly polymor-
Reduced Medusa, Non-feeding and Reproducing Sexually
Non-feeding, Abolished -Medusa
Complete Medusa, Feeding and Reproducing Sexually
Direct Development without hydroid stage Indirect Development with hydroid stage
Hydroid life cycles vary by species. Here is shown an example of a "typical" life cycle. (Illustration by Emily Damstra)
phic modular colonies of polypoid and medusoid zooids attached to a stem or stolon supported by a floating and swimming system or nectosome (pneumatophores and nectophores).
The Polypodiozoa class is represented by a single species, Polypodium hydriforme Ussow, 1885, the only known metazoan adapted to intracellular parasitism. Polypodium has a unique lifecycle, having a succession of a free-living stage and of an intracellular parasitic stage of some Acipenseridae and Poly-odontidae eggs. The earliest parasitic stage known is a binu-cleate cell observed in previtellogenetic fish oocytes. Further parasitic development leads to a didermic stolonal structure, with inverted germ layers, growing at the expense of the egg's yolk and forming numerous inverted buds. Before becoming free at fish spawning, eversion takes place and the germ layers take their normal position (ectoderm out, endoderm inside). Once liberated, the stolon becomes fragmented into individual buds, each originating a free-creeping globular stage that can multiply by longitudinal fission. These stages can move and feed, having an oral mouth-cone and tentacles. Germ cells are endodermal; the females have two kinds of go-nads, each with a gonoduct opening in the gastral cavity; the gonads of the males form gametophores carrying cnidocysts. The free-living stage presumably represents the sexual medusae, the parasitic stages being considered as polypoid.
They differ, however, from all other Hydrozoa by unique features: bilateral symmetry; presence of gonoducts; aberrant ga-metogenesis; unique structure of cnidocil apparatus; inversion of germ layers during parasitic life; and the complete separation of epidermal and muscle cells.
The umbrella of Hydroido- and Automedusae generally measures between 0.02 in (0.5 mm) and 2 in (50 mm), but in numerous species the size may be greater, reaching 3.9-7.8 in (100-200 mm) (Aequorea) and even, exceptionally, 15.7 in (400 mm) of diameter (Rhacostoma atlanticum). The colonies of the Hydroidomedusae usually have a reduced size. Most of them do not exceed a few inches (centimeters to a few decimeters) (i.e., Cladocarpus lignosus is 27.5 in [70 cm]); the hydranths are usually very tenuous, not exceeding a few inches (millimeters), but there are exceptions (i.e., Hydrocoryne miuren-sis: 2.3 in [6 cm]; Corymorpha nutans: 4.7 in [12 cm]; Monocoryne gigantea: 15.7 in [40 cm]; Candelabrumpenola: 33.4 in [85 cm]; and Branchiocerianthus imperator: more than 6.5 ft [2 m]). The smallest polyps are those of the Microhydrulidae (Limnome-dusae): they are reduced to a spherical or irregular body, ranging from 20 to 480 pm. The longest siphonophore is Apolemia uvaria, with colonies reaching 98.4 ft (30 m). The medusae of the hydrozoa are usually diaphanous, as are their polyps. Colored species, however, are frequent. Pigments derive from the diet or are produced directly. The most common color is reddish, deriving from crustaceans; other colors for medusae can be green, white, or orange; whereas polyps can be, according to the species, reddish, pink, white, or blue.
The medusae have typical and easily recognizable body architecture; the main feature that distinguishes them from other medusae is the velum. For this reason, they have also been called Craspedotae (with velum), as opposed to the medusae of the Scypozoa and Cubozoa, which were called Acraspedae (without velum). The polyps are quite varied in architecture, ranging from coral-like colonies (Millepora and the stylasterids) to gigantic polyps resembling those of the Anthozoa (Branchiocerianthus) to microscopic polyps reduced to a simple ball of tissue (Microhydrula).
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