| Symbiodinium Temporal range: Inferred as Eocene–Recent[1] |
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| Scientific classification | |
| Domain: | Eukaryota |
| Kingdom: | Chromalveolata |
| Superphylum: | Alveolata |
| Phylum: | Dinoflagellata |
| Class: | Dinophyceae |
| Order: | Suessiales |
| Family: | Blastodiniaceae |
| Genus: | Symbiodinium |
Zooxanthellae ( /ˌzoʊ.əzænˈθɛliː/) are flagellate protozoa that are golden-brown intracellular endosymbionts of various marine animals and protozoa, especially anthozoans such as the scleractinian corals and the tropical sea anemone, Aiptasia.
Zooxanthellae live in other protozoa (foraminiferans and radiolarians) and in some invertebrates. Most are autotrophs and provide the host with energy in the form of translocated reduced carbon compounds, such as glucose, glycerol, and amino acids, which are the products of photosynthesis.[2] Zooxanthellae can provide up to 90% of a coral’s energy requirements.[3] In return, the coral provides the zooxanthellae with protection, shelter, nutrients (mostly waste material containing nitrogen and phosphorus) and a constant supply of carbon dioxide required for photosynthesis. Available nutrients, incident light, and expulsion of excess cells limits their population.
Hermatypic (reef-building) corals largely depend on zooxanthellae, which limits that coral's growth to the photic zone. The symbiotic relationship enables corals' success as reef-building organisms in tropical waters. However, under high environmental stress, corals die after losing their zooxanthellae either by expulsion or digestion resulting in coral bleaching.
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The coral and zooxanthella relationship has traditionally been considered mutualistic -- that is, both partners benefit from the arrangement. However, whilst the benefit for the coral is clear in terms of its enhanced growth and calcification rate, the benefit for the algae has been called into question.[1]
Benefits cited for the algae include protection from predation and enhanced provision with chemicals such as carbon dioxide and ammonium.[1] However, a number of conditions are thought to be necessary in order to maintain a symbiotic relationship; on a very simple level, the symbiont's optimal reproductive strategy must be to remain in the host. It is not clear that these conditions are met in the coral-zooxanthella instance; reproduction of the zooxanthella is retarded by almost two orders of magnitude when it dwells inside a coral.[1]
The relationship may be better thought of as a parasitic relationship, with the coral parasitic upon its enslaved algae. The coral ensnares the algae by secreting a chemical attractant, before ingesting the algae and incorporating it into its cells. They are then surrounded by a 'symbiosome' membrane and confined within the host cell, separated from its cytoplasm. The host cell then emits chemical signals that prevent the zooxanthella from reproducing.[1]
The presence of the alga results in the production of excess oxygen, which must be removed from the cell quickly in order to avoid destructive oxidation. The corals have mechanisms whereby they can kill over-oxygenated cells if necessary.[1]
Coral bleaching occurs when zooxanthellae densities within coral tissue become low or the concentration of photosynthetic pigments within each zooxanthella decline. Color loss also comes from reduced concentrations of pigments produced by the cnidarian itself. The result is a ghostly white calcareous skeleton. The coral will then die unless conditions improve enough to allow the zooxanthellae to return.
Zooxanthellae directly or indirectly experience the stress that their containing corals undergo. Exposure to air during low tides and damage from solar radiation in shallow water environments are two of the ecological stressors coral and zooxanthellae face. Temperature changes now provide the most stress to the zooxanthellae-coral relationship. A 1-2 degree Celsius temperature rise for 5–10 weeks and a 3-5 degree decline for 5–10 days have produced a coral bleaching event. Such temperature changes induce cell adhesion dysfunction which detaches zooxanthellae from their cnidarian endodermal cells.
Whilst there are genetic differences between the zooxanthellae of different species of coral, there is not agreement as to whether these are significant enough to denote different species.[1]
Some see several different species of zooxanthellae within the Symbiodinium monophylum.[4] Symbiodinium is related to Gymnodinium simplex, Gymnodinium beii, and Polarella glacialis.[4]
The scleractinian-zooxanthella relationship evolved multiple times independently in the mid-Triassic, 245 million years ago, and probably enabled the corals to build carbonate skeletons in the adverse Triassic seas[5] (which had caused the tabulate and rugose corals to become extinct). However, genetic evidence suggests that Symbiodinium did not begin to diversify until the early-to-middle Eocene, just 50 million years ago.[1]
The genus, Symbiodinium, was defined by Hugo Freudenthal in 1970, after his identification of the life cycle of zooxanthella from Cassiopea. At that time he proved that symbodinia had a motile stage which resembled a "gymnodinioid" dinoflagellate. Being both symbiotic and a dinoflagellate, he named the genus Symbiodinium, and the species epithet microadriaticium, after its resemblance to a similar free-living species.
Dr. Freudenthal demonstrated that zooxanthellae go through a vegetative stage, a cyst stage, and a motile stage as part of their life cycle.
Polyps can acquire Zooxanthellae by direct ingestion. However, their hosts do not digest them. In other cases, zooxanthellae may be transmitted by coral eggs and planulae.
Zooxanthellae reproduce asexually by budding (one individual splitting into multiple descendants).
Other organisms which can host zooxanthellae include jellyfish, clams, foraminifera, sea slugs i.e. nudibranchs such as Pteraeolidia ianthina, and also ciliates and radiolaria.
Rudman, W.B., 2000 (October 10) What are Zooxanthellae?. [In] Sea Slug Forum. Australian Museum, Sydney. Available from http://www.seaslugforum.net/factsheet.cfm?base=zoox1
Riddle, D., 2006 (January) Lighting by numbers: ”types” of Zooxanthellae and what they tell us. Advanced Aquarist’s Online Magazine. Available from http://www.advancedaquarist.com/2006/1/aafeature1
Buchheim, J., 1998 Coral Reef Bleaching. Odyssey Expeditions – Marine Biology Learning Center Publications. Available from http://www.marinebiology.org/coralbleaching.htm
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