Symbiodinium

These unicellular microalgae commonly reside in the endoderm of tropical cnidarians such as corals, sea anemones, and jellyfish, where the products of their photosynthetic processing are exchanged in the host for inorganic molecules.

Cnidarians that are associated with Symbiodinium occur mostly in warm oligotrophic (nutrient-poor), marine environments where they are often the dominant constituents of benthic communities.

Under normal conditions, symbiont and host cells exchange organic and inorganic molecules that enable the growth and proliferation of both partners.

Coral reefs have economic benefits – valued at hundreds of billions of dollars each year – in the form of ornamental, subsistence and commercial fisheries, tourism and recreation, coastal protection from storms, a source of bioactive compounds for pharmaceutical development, and more.

A chief mechanism for widespread reef degradation has been stress-induced coral bleaching caused by unusually high seawater temperature.

[27][28][29] The symbiosis Symbodinium-coral could provide higher resistance to multiple stress (desiccation, high UVR) to the coral holobiont through its mycosporine-like amino acids (MAAs).

It is present in small numbers in coral globally and is common in the Andaman Sea, where the water is about 4 °C (7 °F) warmer than in other parts of the Indian Ocean.

In the Caribbean Sea in late 2005, water temperature was elevated for several months and it was found that S. trenchi, a symbiont not normally abundant, took up residence in many corals in which it had not previously been observed.

The application of this methodology helped overturn the long-held belief that (traditional understood) Symbiodinium comprised a single genus, a process which began in earnest with the morphological, physiological, and biochemical comparisons of cultured isolates.

Currently, genetic markers are exclusively used to describe ecological patterns and deduce evolutionary relationships among morphologically cryptic members of this group.

The earliest ribosomal gene sequence data indicated that Symbiodinium had lineages whose genetic divergence was similar to that seen in other dinoflagellates from different genera, families, and even orders.

The high concordance found among nuclear, mitochondrial and chloroplast DNA argues that a hierarchical phylogenetic scheme, combined with ecological and population genetic data, can unambiguously recognize and assign nomenclature to reproductively isolated lineages, i.e.

[46] Through the use of microsatellite markers, multilocus genotypes identifying a single clonal line of Symbiodinium can be resolved from samples of host tissue.

[citation needed] The earliest genetic methods for assessing Symbiodinium diversity relied on low-resolution molecular markers that separated the genus into a few evolutionarily divergent lineages, referred to as "clades".

Previous characterizations of geographic distribution and dominance have focused on the clade-level of genetic resolution, but more detailed assessments of diversity at the species level are needed.

[citation needed] The large diversity of Symbiodinium revealed by genetic analyses is distributed non-randomly and appears to comprise several guilds with distinct ecological habits.

[54] Finally, there appears to be another group of Symbiodinium that are incapable of establishing endosymbiosis yet exist in environments around the animal or associate closely with other substrates (i.e. macro-algal surfaces, sediment surface)[47][55] Symbiodinium from functional groups 2, 3, and 4 are known to exist because they culture easily, however species with these life histories are difficult to study because of their low abundance in the environment.

The use of aposymbiotic host polyps deployed as "capture vessels" and the application of molecular techniques has allowed for the detection of environmental sources of Symbiodinium.

[53][56] With these methods employed, investigators may resolve the distribution of different species on various benthic surfaces[55] and cell densities suspended in the water column.

[50] Learning more about the "private lives" of these environmental populations and their ecological function will further our knowledge about the diversity, dispersal success, and evolution among members within this large genus.

The comparison of cultured isolates under identical conditions show clear differences in morphology, size, biochemistry, gene expression, swimming behavior, growth rates, etc.

Culturing is a selective process, and many Symbiodinium isolates growing on artificial media are not typical of the species normally associated with a particular host.

Samples for genetic analysis should be acquired from the source colony in order to match the resulting culture with the identity of the dominant and ecologically relevant symbiont originally harbored by the animal.

For isolates that are in log phase growth, division rates occur every 1–3 days, with Symbiodinium cells alternating between a spherical, or coccoid, morphology and a smaller flagellated motile mastigote stage.

While several similar schemes are published that describe how each morphological state transitions to other, the most compelling life history reconstruction was deduced from light and electron microscopy and nuclear staining evidence.

[64] During asexual propagation (sometimes referred to as mitotic or vegetative growth), cells undergo a diel cycle of karyokinesis (chromosome/nuclear division) in darkness.

[64] Approaching or at the end of the photoperiod the mastigotes cease swimming, release their flagella, and undergo a rapid metamorphosis into the coccoid form.

Mucocysts (an ejectile organelle[69]) located beneath the plasmalemma are found in S. pilosum and their function is unknown, but may be involved in heterotrophic feeding.

[67] The term cyst usually refers to a dormant, metabolically quiescent stage in the life history of other dinoflagellates, initiated by several factors, including nutrient availability, temperature, and day length.

All cultured isolates (i.e. strains) are capable of phenotypic adjustment in their capacity for light harvesting (i.e. photoacclimation), such as by altering the cellular Chl.

Light and confocal images of Symbiodinium cells in hospite (living in a host cell) within scyphistomae of the jellyfish Cassiopea xamachana . This animal requires infection by these algae to complete its life cycle . The chloroplast imaged in 3-D is highly reticulated and distributed around the cell's periphery
Symbiodinium reach high cell densities through prolific mitotic division in the endodermal tissues of many shallow tropical and sub-tropical cnidarians . This is a SEM of a freeze-fractured internal mesentery from a reef coral polyp ( Porites porites ) that shows the distribution and density of symbiont cells.
Genetic disparity between clades in the legacy genus Symbiodinium sensu lato compared to other dinoflagellates. Analysis of conserved mitochondrial sequences ( CO1 ) and rDNA ( SSU ) suggest that a taxonomic revision of this group was required. See clades A—F , Polarella , Scrippsiella , Pfiesteria , Peridinium , Lingulodinium , Alexandrium , Karlodinium , Karenia , Gymnodinium , Gyrodinium , Aka­shiwo , and Prorocentrum .
The investigation of Symbiodinium diversity, ecology, and evolution is enhanced by analysis of ribosomal and single copy nuclear , plastid , and mitochondrial DNA. The use of multiple markers , along with a hierarchical phylogenetic classification provides the genetic resolution necessary for investigating species diversity, biogeography, dispersal, natural selection, and adaptive radiations.
Global distribution and preliminary diversity estimate of common Symbiodinium species associated with cnidarians (rare “species” excluded). Internal transcribed spacer region 2 ( ITS2 ) data ( sensu LaJeunesse 2002) are used here as a proxy for species diversity.
The distributions of Symbiodinium species comprising different ecological guilds in a coral reef ecosystem . A . Host cnidarians can expel millions of symbiont cells (viable and necrotic) per day into the surrounding environment ( a ). In turn these animals pass large volumes of water through their gastrovascular system for respiration and cellular waste removal , a process that introduces numerous small particles including food and various other Symbiodinium spp. (indicated by differences in cell color) ( b ). B . The ecological niche , from the viewpoint of functional groups, differs among species of Symbiodinium . Several ecological guilds exist, including abundant host-specific and host-generalist species ( 1 ), low background and potentially opportunistic species ( 2 ), and non-symbiotic species closely associated with the coral's biome ( 3 ) and/or occupying unrelated habitats ( 4 )
Symbiodinium life cycle
The life stages of dinoflagellates in the genus Symbiodinium . ( A ) Electron micrographs of a Symbiodinium mastigote (motile cell) with characteristic gymnodinioid morphology ( S. natans ) and ( B ) the coccoid cell in hospite. As free-living cells the mastigote allows for short-range dispersal and can exhibit chemotaxis toward sources of nitrogen . Once within the host, these symbionts rapidly proliferate in and in many cases dominate the cytoplasm of the host cell.
Fine scale features and organelles found among Symbiodinium species. ( A ) Close-up of divided cells of a tetrad from a culture of S. corculorum that is genetically similar to S. pilosum (both are type/subclade A2). This particular lineage is unusual among described Symbiodinium by possessing mucocysts , and chloroplasts with parallel and peripheral thylakoid arrangements. Flagella are seen in cross section in the space between daughter cells. ( B ) A thick cellulosic cell wall is observed for many cultured isolates. ( C ) TEM of a cross-section through a chloroplast lobe at the periphery of the cell with parallel and peripheral thylakoids grouped in sets of three.
TEM of cross-section through a dividing cell (the doublet). The light micrograph inset depicts a doublet in culture. Flagella are present from the daughter mastigote cells (white arrowheads) prior to emerging from the mother cell wall. n = nucleus ; acc = accumulation body ; pyr = pyrenoid . (Robert K. Trench et al . 1981)
The reticulated chloroplast structures of ( A ) Cladocopium goreaui (formerly S. goreaui type C1), ( B ) Symbiodinium fitti (type A3), and ( C ) Effrenium voratum (formerly S. californium type E1) [ 3 ] imaged in 3-D using confocal microscopy of chlorophyll autofluorescence .
Symbiodinium kawagutii