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   » » Wiki: Photosymbiosis
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Photosymbiosis is a type of where one of the is capable of .

Examples of photosymbiotic relationships include those in , , , and many organisms including , , , and .

Photosymbiosis is important in the development, maintenance, and of terrestrial and aquatic , for example in biological soil crusts, , supporting highly diverse microbial populations in and , and growth and maintenance.

When one organism lives within another symbiotically it's called . Photosymbiotic relationships where and/or live within a host organism, are believed to have led to acquiring photosynthesis and to the evolution of .


Occurrence

Lichens
represent an association between one or more mycobionts and one or more photosynthetic or cyanobacterial photobionts. The mycobiont provides protection from predation and , while the photobiont provides energy in the form of fixed carbon. Cyanobacterial partners are also capable of fixing nitrogen for the fungal partner. Recent work suggests that non-photosynthetic bacterial associated with lichens may also have functional significance to lichens.

Most mycobiont partners derive from the , and the largest class of lichenized fungi is . The vast majority of lichens derive photobionts from (green algae). The co-evolutionary dynamics between mycobionts and photobionts are still unclear, as many photobionts are capable of free-living, and many lichenized fungi display traits adaptive to lichenization such as the capacity to withstand higher levels of reactive oxygen species (ROS), the conversion of sugars to that help withstand dedication, and the downregulation of fungal . However, it is still unclear whether these are derived or ancestral traits.

Currently described photobiont species number about 100, far less than the 19,000 described species of fungal mycobionts, and factors such as geography can predominate over mycobiont preference. Phylogenetic analyses in lichenized fungi have suggested that, throughout evolutionary history, there has been repeated loss of photosymbionts, switching of photosymbionts, and independent lichenization events in previously unrelated fungal taxa. Loss of lichenization has likely led to the coexistence of non-lichenized fungi and lichenized fungi in lichens.


Sponges
(phylum Porifera) have a large diversity of photosymbiote associations. Photosymbiosis is found in four classes of Porifera (, , Homoscleromorpha, and Calcarea), and known photosynthetic partners are cyanobacteria, , , and red () and green (Chlorophyta) algae. Relatively little is known about the evolutionary history of sponge photosymbiois due to a lack of data. However, it has been shown that photosymbiotes are acquired vertically (transmission from parent to offspring) and/or horizontally (acquired from the environment). Photosymbiotes can supply up to half of the host sponge's respiratory demands and can support sponges during times of nutrient stress.


Cnidaria
Members of certain classes in phylum are known for photosymbiotic partnerships. Members of corals (Class ) in the orders and form well-characterized partnerships with the dinoflagellate genus . Some jellyfish (class ) in the genus (upside-down jellyfish) also possess Symbiodinium. Certain species in the genus Hydra (class ) also harbor green algae and form a stable photosymbiosis.

The evolution of photosymbiosis in corals was likely critical for the global establishment of . Corals are likewise adapted to eject damaged photosymbionts that generate high levels of toxic reactive oxygen species, a process known as . The identity of the Symbiodinium photosymbiont can change in corals, although this depends largely on the mode of transmission: some species vertically transmit their algal partners through their eggs, while other species acquire environmental dinoflagellates as newly-released eggs. Since algae are not preserved in the coral fossil record, understanding the evolutionary history of the symbiosis is difficult.


Bilaterians
In basal , photosymbiosis in marine or systems is present only in the family . In the group there is limited knowledge on the symbionts present, and they have been vaguely identified as or . Some species have a symbiotic relationship with the chlorophyte convolutae while others have a symbiotic relationship with the dinoflagellates , klebsii, or in the genus Licomorpha.

In freshwater systems, photosymbiosis is present in belonging to the group. In this group, members of the , , and families are symbiotic. Members of Provorticidae likely feed on diatoms and retain their symbionts. Typhloplanidae have symbiotic relationships with the chlorophytes in the genus .


Molluscs
Photosymbiosis is taxonomically restricted in . Tropical marine in the Cardiidae family form a symbiotic relationship with the dinoflagellate . This family boasts large organisms often referred to as and their large size is attributed to the establishment of these symbiotic relationships. Additionally, the Symbiodinium are hosted extracellularly, which is relatively rare. The only known freshwater bivalve with a symbiotic relationship are in the genus which hosts the chlorophyte Chlorella in the gills and mantle of the host. In bivalves, photosymbiosis is thought to have evolved twice, in the genus Anodonta and in the family Cardiidae. However, how it has evolved in Cardiidae could have occurred through different gains or losses in the family.


Gastropods
In , photosymbiosis can be found in several genera.

The species Strombus gigas hosts which is acquired during the larval stage, at which point it is a mutualistic relationship. However, during the adult stage, Symbiodinium becomes as the shell prevents photosynthesis.

Another group of gastropods, sea slugs, have two different systems for symbiosis. The first, , acquire their symbionts through feeding on prey that are in symbiotic relationships. In Nudibranchs, photosymbiosis has evolved twice, in and . In Aeolidida it is likely there have been several gains and losses of photosymbiosis as most genera include both photosymbiotic and non-photosymbiotic species. The second, , removes from macroalgae when feeding and sequesters them into their digestive tract at which point they are called . Whether these kleptoplasts maintain their photosynthetic capabilities depends on the host species ability to digest them properly. In this group, functional kleptoplasty has been acquired twice, in and .


Chordates
Photosymbiosis is relatively uncommon in species. One such example of photosymbiosis is in , the sea squirts. In the genus , 30 species establish symbiotic relationships. The photosynthetic ascidians are associated with in the genus of as well as, in some cases, the species trididemni. The 30 species with a symbiotic relationship span four genera where the congeners (species within the same genus) are primarily non-symbiotic, suggesting multiple origins of photosymbiosis in ascidians.

In addition to sea squirts, embryos of some species ( Ambystoma maculatum, Ambystoma gracile, Ambystoma jeffersonium, , Hynobius nigrescens, , and Lithobates aurora) form symbiotic relationships with the in the genus of Oophila. This algae is present in the egg masses of the species, causing them to appear green and providing oxygen and carbohydrates to the embryos. Similarly, little is known about the evolution of symbiosis in amphibians, but there appear to be multiple origins.


Protists
Photosymbiosis has evolved multiple times in the protist taxa , , , , and . Foraminifera and Radiolaria are taxa that serve as in open ocean communities. Photosynthetic plankton species associate with the symbiotes of dinoflagellates, diatoms, , , and that can be transferred both vertically and horizontally. In Foraminifera, species will either have a symbiotic relationship with Symbiodinium or retain the chloroplasts present in algal prey species. The planktonic species of Foraminifera associate primarily with Pelagodinium. These species are often considered indicator species due to their bleaching in response to environmental stressors. In the Radiolarian group , photosynthetic species inhabit surface waters whereas non-photosynthetic species inhabit deeper waters. Photosynthetic Acantharia are associated with similar microalgae as the Foraminifera groups, but were also found to be associated with , , Scrippsiella, and which were not previously known to be involved in photosynthetic relationships. In addition, several of the species present in symbiotic relationships with Acantharia were oftentimes identical to the free-living species, suggesting horizontal transfer of symbiotes. This provides insight into the evolutionary patterns responsible for these symbiotic relationships, suggesting that the selection for symbiosis is relatively weak and symbiosis is likely a result of the adaptive capacity of the host plankton species.


See also

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