Sea anemone
Sea anemones are a group of predatory marine invertebrate animals constituting the order Actiniaria. Because of their colourful appearance, they are named after the Anemone, a terrestrial flowering plant. Sea anemones are classified in the phylum Cnidaria, class Anthozoa, subclass Hexacorallia.
As cnidarians, sea anemones are related to corals, jellyfish, tube-dwelling anemones, and Hydra. Unlike jellyfish, sea anemones do not have a medusa stage in their life cycle.
A typical sea anemone is a single polyp attached to a hard surface by its base, but some species live in soft sediment, and a few float near the surface of the water. The polyp has a columnar trunk topped by an oral disc with a ring of tentacles and a central mouth. The tentacles can be retracted inside the body cavity or expanded to catch passing prey. They are armed with cnidocytes. In many species, additional nourishment comes from a symbiotic relationship with single-celled dinoflagellates, with zooxanthellae, or with green algae, zoochlorellae, that live within the cells. Some species of sea anemone live in association with clownfish, hermit crabs, small fish, or other animals to their mutual benefit.
Sea anemones breed by liberating sperm and eggs through the mouth into the sea. The resulting fertilized eggs develop into planula larvae which, after being planktonic for a while, settle on the seabed and develop directly into juvenile polyps. Sea anemones also breed asexually, by breaking in half or into smaller pieces which regenerate into polyps. Sea anemones are sometimes kept in reef aquariums; the global trade in marine ornamentals for this purpose is expanding and threatens sea anemone populations in some localities, as the trade depends on collection from the wild.
Anatomy
A typical sea anemone is a sessile polyp attached at the base to the surface beneath it by an adhesive foot, called a basal or pedal disc, with a column-shaped body topped by an oral disc. Most are from in diameter and in length, but they are inflatable and vary greatly in dimensions. Some are very large; Urticina columbiana and Stichodactyla mertensii can both exceed in diameter and Metridium farcimen a metre in length. Some species burrow in soft sediment and lack a basal disc, having instead a bulbous lower end, the physa, which anchors them in place.The column or trunk is generally more or less cylindrical and may be plain and smooth or may bear specialised structures; these include solid papillae, adhesive papillae, cinclides, and small protruding vesicles. In some species the part immediately below the oral disc is constricted and is known as the capitulum. When the animal contracts, the oral disc, tentacles and capitulum fold inside the pharynx and are held in place by a strong bone all the way up the column. There may be a fold in the body wall, known as a parapet, at this point, and this parapet covers and protects the anemone when it is retracted.
The oral disc has a central mouth, usually slit-shaped, surrounded by one or more whorls of tentacles. The ends of the slit lead to grooves in the wall of the pharynx known as siphonoglyphs; there are usually two of these grooves, but some groups have a single one. The tentacles are generally tapered and often tipped by a pore, but in some species they are branched, club-tipped, or reduced to low knobs. The tentacles are armed with many cnidocytes, cells that are both defensive and used to capture prey. Cnidocytes contain stinging nematocysts, capsule-like organelles capable of everting suddenly, giving the phylum Cnidaria its name. Each nematocyst contains a small venom vesicle filled with actinotoxins, an inner filament, and an external sensory hair. A touch to the hair mechanically triggers a cell explosion, which launches a harpoon-like structure that attaches to the organism that triggered it, and injects a dose of venom in the flesh of the aggressor or prey.
At the base of the tentacles in some species, primarily aggregating anemones, lie acrorhagi, elongated inflatable tentacle-like organs armed with cnidocytes, that can flail around and fend off other encroaching anemones; one or both anemones can be driven off or suffer injury in such battles.
Many sea anemones also have acontia, thin filaments covered in cnidae that can be ejected and retracted for defence.
The venom is a mix of toxins, including neurotoxins, that paralyzes the prey so the anemone can move it to the mouth for digestion inside the gastrovascular cavity. Actinotoxins are highly toxic to prey species of fish and crustaceans. However, Amphiprioninae, small banded fish in various colours, are not affected by their host anemone's sting and shelter themselves from predators among its tentacles. Several other species have similar adaptions and are also unaffected. Most sea anemones are harmless to humans, but a few highly toxic species have caused severe injuries and are potentially lethal.
Digestive system
Sea anemones have what can be described as an incomplete gut: the gastrovascular cavity functions as a stomach and possesses a single opening to the outside, which operates as both a mouth and anus. Waste and undigested matter are excreted through this opening. The mouth is typically slit-like in shape and bears a groove at one or both ends. The groove, termed a siphonoglyph, is ciliated, and helps to move food particles inwards and circulate water through the gastrovascular cavity.The mouth opens into a flattened pharynx. This consists of an infolding of the body wall, and is therefore lined by the animal's epidermis. The pharynx typically runs for about one-third the length of the body before opening into the gastrovascular cavity that occupies the remainder of the body.
The gastrovascular cavity itself is divided into several chambers by mesenteries radiating inwards from the body wall. Some of the mesenteries form complete partitions with a free edge at the base of the pharynx, where they connect, but others reach only partway across. The mesenteries are usually found in multiples of twelve and are symmetrically arranged around the central lumen. They have a stomach lining on both sides, separated by a thin layer of mesoglea, and include filaments of tissue specialised for secreting digestive enzymes. In some species, these filaments extend below the lower margin of the mesentery, hanging free in the gastrovascular cavity as thread-like acontial filaments. These acontia are armed with nematocysts and can be extruded through cinclides, blister-like holes in the wall of the column, for use in defence.
Musculature and nervous system
A primitive nervous system, without centralization, coordinates the processes involved in maintaining homeostasis, as well as biochemical and physical responses to various stimuli. There are two nerve nets, one in the epidermis and one in the gastrodermis; these unite at the pharynx, the junctions of the septa with the oral disc and the pedal disc, and across the mesogloea. No specialized sense organs are present, but sensory cells include nematocytes and chemoreceptors.The muscles and nerves are much simpler than those of most other animals, although more specialised than in other cnidarians, such as corals. Cells in the outer layer and the inner layer have microfilaments that group into contractile fibers. These fibers are not true muscles because they are not freely suspended in the body cavity as they are in more developed animals. Longitudinal fibres are found in the tentacles and oral disc, and also within the mesenteries, where they can contract the whole length of the body. Circular fibers are found in the body wall and, in some species, around the oral disc, allowing the animal to retract its tentacles into a protective sphincter.
Since the anemone lacks a rigid skeleton, the contractile cells pull against the fluid in the gastrovascular cavity, forming a hydrostatic skeleton. The anemone stabilizes itself by flattening its pharynx, which acts as a valve, keeping the gastrovascular cavity at a constant volume and making it rigid. When the longitudinal muscles relax, the pharynx opens and the cilia lining the siphonoglyphs beat, wafting water inwards and refilling the gastrovascular cavity. In general, the sea anemone inflates its body to extend its tentacles and feed, and deflates it when resting or disturbed. The inflated body is also used to anchor the animal inside a crevice, burrow or tube.
Life cycle
Unlike other cnidarians, anemones entirely lack the free-swimming medusal stage of their life cycle; the polyp produces eggs and sperm, and the fertilized egg develops into a planula larva, which develops directly into another polyp. Both sexual and asexual reproduction can occur.The sexes in sea anemones are separate in some species, while other species are sequential hermaphrodites, changing sex at some stage in their life. The gonads are strips of tissue within the mesenteries. In sexual reproduction, males may release sperm to stimulate females to release eggs, and fertilization occurs, either internally in the gastrovascular cavity or in the water column. The eggs and sperm, or the larvae, usually emerge through the mouth, but in some species, such as Metridium dianthus, may be swept out from the body cavity through the cinclides. In many species the eggs and sperm rise to the surface where fertilisation occurs. The fertilized egg develops into a planula larva, which drifts for a while before sinking to the seabed and undergoing metamorphosis into a juvenile sea anemone. Some larvae preferentially settle onto certain suitable substrates; the mottled anemone for example, settles onto green algae, perhaps attracted by a biofilm on the surface.
The brooding anemone is gynodioecious, starting life as a female and later becoming hermaphroditic, so that populations consist of females and hermaphrodites. As a female, the eggs can develop parthenogenetically into female offspring without fertilisation, and as a hermaphrodite, the eggs are routinely self-fertilised. The larvae emerge from the anemone's mouth and tumble down the column, lodging in a fold near the pedal disc. Here they develop and grow, remaining for about three months before crawling off to start independent lives.
Sea anemones have great powers of regeneration and can reproduce asexually, by budding, fragmentation, or longitudinal or transverse binary fission. Some species such as certain Anthopleura divide longitudinally, pulling themselves apart, resulting in groups of individuals with identical colouring and markings. Transverse fission is less common, but occurs in Anthopleura stellula and Gonactinia prolifera, with a rudimentary band of tentacles appearing halfway up the column before it splits horizontally. Some species can also reproduce by pedal laceration. In this process, a ring of material may break off from the pedal disc at the base of the column, which then fragments, the pieces regenerating into new clonal individuals. Alternatively, fragments detach separately as the animal creeps across a surface. In Metridium dianthus, fragmentation rates were higher in individuals living among live mussels than among dead shells, and all the new individuals had tentacles within three weeks.
The sea anemone Aiptasia diaphana displays sexual plasticity. Thus asexually produced clones derived from a single founder individual can contain both male and female individuals. When eggs and sperm are formed, they can produce zygotes derived from "selfing" or out-crossing, which then develop into swimming planula larvae. Anemones tend to grow and reproduce relatively slowly. The magnificent sea anemone, for example, may live for decades, with one individual surviving in captivity for eighty years.