Lichens represent an association between one or more fungal mycobionts and one or more photosynthetic algal or cyanobacterial photobionts. The mycobiont provides protection from predation and desiccation, while the photobiont provides energy in the form of fixed carbon. Cyanobacterial partners are also capable of fixing nitrogen for the fungal partner.[9] Recent work suggests that non-photosynthetic bacterial microbiomes associated with lichens may also have functional significance to lichens.[10]
Most mycobiont partners derive from the ascomycetes, and the largest class of lichenized fungi is Lecanoromycetes.[11] The vast majority of lichens derive photobionts from Chlorophyta (green algae).[9] 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 polypols that help withstand dedication, and the downregulation of fungal virulence. However, it is still unclear whether these are derived or ancestral traits.[9]
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.[12][13] 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.[11][14] Loss of lichenization has likely led to the coexistence of non-lichenized fungi and lichenized fungi in lichens.[14]
Sponges
Sponges (phylum Porifera) have a large diversity of photosymbiote associations. Photosymbiosis is found in four classes of Porifera (Demospongiae, Hexactinellida, Homoscleromorpha, and Calcarea), and known photosynthetic partners are cyanobacteria, chloroflexi, dinoflagellates, and red (Rhodophyta) and green (Chlorophyta) algae. Relatively little is known about the evolutionary history of sponge photosymbiois due to a lack of genomic data.[15] However, it has been shown that photosymbiotes are acquired vertically (transmission from parent to offspring) and/or horizontally (acquired from the environment).[16] Photosymbiotes can supply up to half of the host sponge’s respiratory demands and can support sponges during times of nutrient stress.[17]
Cnidaria
Members of certain classes in phylum Cnidaria are known for photosymbiotic partnerships. Members of corals (Class Anthozoa) in the orders Hexacorallia and Octocorallia form well-characterized partnerships with the dinoflagellate genus Symbiodinium. Some jellyfish (class Scyphozoa) in the genus Cassiopea (upside-down jellyfish) also possess Symbiodinium. Certain species in the genus Hydra (class Hydrozoa) also harbor green algae and form a stable photosymbiosis.[15]
The evolution of photosymbiosis in corals was likely critical for the global establishment of coral reefs.[18] Corals are likewise adapted to eject damaged photosymbionts that generate high levels of toxic reactive oxygen species, a process known as bleaching.[19] 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,[20] while other species acquire environmental dinoflagellates as newly-released eggs.[21] Since algae are not preserved in the coral fossil record, understanding the evolutionary history of the symbiosis is difficult.[22]
In freshwater systems, photosymbiosis is present in platyhelminths belonging to the Rhabdocoela group.[34] In this group, members of the Provorticidae, Dalyeliidae, and Typhloplanidae families are symbiotic.[35] Members of Provorticidae likely feed on diatoms and retain their symbionts.[36] Typhloplanidae have symbiotic relationships with the chlorophytes in the genus Chlorella.[37]
Molluscs
Photosymbiosis is taxonomically restricted in Mollusca.[38] Tropical marine bivalves in the Cardiidae family form a symbiotic relationship with the dinoflagellate Symbiodinium.[39] This family boasts large organisms often referred to as giant clams and their large size is attributed to the establishment of these symbiotic relationships. Additionally, the Symbiodinium are hosted extracellularly, which is relatively rare.[40] The only known freshwater bivalve with a symbiotic relationship are in the genus Anodonta which hosts the chlorophyte Chlorella in the gills and mantle of the host.[41] In bivalves, photosymbiosis is thought to have evolved twice, in the genus Anodonta and in the family Cardiidae.[42] However, how it has evolved in Cardiidae could have occurred through different gains or losses in the family.[43]
Gastropods
In gastropods, photosymbiosis can be found in several genera.
The species Strombus gigas hosts Symbiodinium which is acquired during the larval stage, at which point it is a mutualistic relationship.[44] However, during the adult stage, Symbiodinium becomes parasitic as the shell prevents photosynthesis.[45]
Another group of gastropods, heterobranch sea slugs, have two different systems for symbiosis. The first, Nudibranchia, acquire their symbionts through feeding on cnidarian prey that are in symbiotic relationships.[46] In Nudibranchs, photosymbiosis has evolved twice, in Melibe and Aeolidida.[47] In Aeolidida it is likely there have been several gains and losses of photosymbiosis as most genera include both photosymbiotic and non-photosymbiotic species.[48] The second, Sacoglossa, removes chloroplasts from macroalgae when feeding and sequesters them into their digestive tract at which point they are called kleptoplasts.[49] Whether these kleptoplasts maintain their photosynthetic capabilities depends on the host species ability to digest them properly.[50] In this group, functional kleptoplasy has been acquired twice, in Costasiellidae and Plakobranchacea.[51]
Chordates
Photosymbiosis is relatively uncommon in chordate species.[52] One such example of photosymbiosis is in ascidians, the sea squirts. In the genus Didemnidae, 30 species establish symbiotic relationships.[53] The photosynthetic ascidians are associated with cyanobacteria in the genus of Prochloron as well as, in some cases, the species Synechocystis trididemni.[54] The 30 species with a symbiotic relationship span four genera where the congeners are primarily non-symbiotic, suggesting multiple origins of photosymbiosis in ascidians.[55]
In addition to sea squirts, embryos of some amphibian species (Ambystoma maculatum, Ambystoma gracile, Ambystoma jeffersonium, Ambystoma trigrinum, Hynobius nigrescens, Lithobates sylvaticus, and Lithobates aurora) form symbiotic relationships with the green alga in the genus of Oophila.[56][57][58] This algae is present in the egg masses of the species, causing them to appear green and providing oxygen and carbohydrates to the embryos.[59] Similarly, little is known about the evolution of symbiosis in amphibians, but there appears to be multiple origins.
Protists
Photosymbiosis has evolved multiple times in the protist taxa Ciliophora, Foraminifera, Radiolaria, Dinoflagellata, and diatoms.[60] Foraminifera and Radiolaria are planktonic taxa that serve as primary producers in open ocean communities.[61] Photosynthetic plankton species associate with the symbiotes of dinoflagellates, diatoms, rhodophytes, chlorophytes, and cyanophytes that can be transferred both vertically and horizontally.[62] In Foraminifera, benthic species will either have a symbiotic relationship with Symbiodinium or retain the chloroplasts present in algal prey species.[63] The planktonic species of Foraminifera associate primarily with Pelagodinium.[64] These species are often considered indicator species due to their bleaching in response to environmental stressors.[65] In the Radiolarian group Acantharia, 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 Phaeocystis, Heterocapsa, Scrippsiella, and Azadinium which were not previously known to be involved in photosynthetic relationships.[66] 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.[67] 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.
2010 novel by John G. Hemry This article consists almost entirely of a plot summary. Please help improve the article by adding more real-world context. (June 2012) (Learn how and when to remove this template message) The Lost Fleet: Victorious First Edition Book CoverAuthorJack CampbellCover artistPeter BollingerCountryUnited StatesLanguageEnglish languageSeriesThe Lost FleetGenreMilitary science fictionPublisherAce BooksPublication dateApril 27, 2010Pages352 (Paperback)ISBN978-0-441-018...
Map all coordinates using OpenStreetMap Download coordinates as: KML GPX (all coordinates) GPX (primary coordinates) GPX (secondary coordinates) The State Register of Heritage Places is maintained by the Heritage Council of Western Australia. As of 2023[update], 141 places are heritage-listed in the Shire of Denmark,[1] of which three are on the State Register of Heritage Places.[2] List State Register of Heritage Places The Western Australian State Register of Herita...
Artikel atau sebagian dari artikel ini mungkin diterjemahkan dari Rd Mochtar di en.wikipedia.org. Isinya masih belum akurat, karena bagian yang diterjemahkan masih perlu diperhalus dan disempurnakan. Jika Anda menguasai bahasa aslinya, harap pertimbangkan untuk menelusuri referensinya dan menyempurnakan terjemahan ini. Anda juga dapat ikut bergotong royong pada ProyekWiki Perbaikan Terjemahan. (Pesan ini dapat dihapus jika terjemahan dirasa sudah cukup tepat. Lihat pula: panduan penerjemahan ...
Cinema of Malaysia List of Malaysian filmsList of Malaysian cinemasList of Malaysian animated films pre-1960 1960s 1970s 1980s 1990s 2000s 2000 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010s 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019 2020s 2020 2021 2022 2023 2024 2025 2026 2027 2028 2029 vte This is an index for the list of films produced in Malaysia ordered by decade on separate pages. For an alphabetical listing of Malaysian films see Category:Malaysian films. Before 1960 List ...
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Turkish novelist and columnist This biography of a living person needs additional citations for verification. Please help by adding reliable sources. Contentious material about living persons that is unsourced or poorly sourced must be removed immediately from the article and its talk page, especially if potentially libelous.Find sources: Ayşe Kulin – news · newspapers · books · scholar · JSTOR (February 2009) (Learn how and when to remove this templa...
Relative representation of a species in anr ecosystem Effects of herbivore abundance in seagrass meadows. In ecology, local abundance is the relative representation of a species in a particular ecosystem.[1] It is usually measured as the number of individuals found per sample. The ratio of abundance of one species to one or multiple other species living in an ecosystem is referred to as relative species abundances.[1] Both indicators are relevant for computing biodiversity. A ...
John SaxbyBiographieNaissance 17 août 1821BrightonDécès 22 avril 1913 (à 91 ans)HassocksNationalité britanniqueActivité Ingénieurmodifier - modifier le code - modifier Wikidata John Saxby, né le 17 août 1821 à Brighton et mort le 22 avril 1913 à Hassocks[1],[2], est un inventeur et industriel anglais, spécialiste dans les enclenchements et la signalisation ferroviaire lors du développement des réseaux de chemins de fer dans le monde[3]. Biographie France - Gare de Nîmes - ...
American businessman (born 1957) Christopher MellonMellon in October 2017BornChristopher Karl Mellon (1957-10-02) October 2, 1957 (age 66)Topeka, Kansas, United StatesEducationColby College, Yale UniversityOccupation(s)Deputy Assistant Secretary of Defense, Staff Director of the Senate Intelligence CommitteeYears active1985-2004Organization(s)Department of Defense, United States SenateRelativesJudge Thomas Mellon, Andrew Mellon, Matthew Mellon, Richard Mellon Scaife, Paul MellonFami...
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Alphabet used for writing the Mandaic language MandaicScript type Alphabet Time period2nd century AD — presentDirectionRight-to-left script LanguagesClassical MandaicNeo-MandaicRelated scriptsParent systemsPhoenicianAramaicMandaicISO 15924ISO 15924Mand (140), Mandaic, MandaeanUnicodeUnicode aliasMandaicUnicode rangeU+0840–U+085F This article contains phonetic transcriptions in the International Phonetic Alphabet (IPA). For an introductory guide on IPA symbols, s...
Channel access method used by various radio communication technologies This article is about a channel access method. For the mobile phone technology referred to as CDMA, see cdmaOne and CDMA2000. Multiplexing Analog modulation AM FM PM QAM SM SSB Circuit mode(constant bandwidth) TDM FDM / WDM SDMA Polarization Spatial OAM Statistical multiplexing(variable bandwidth) Packet switching Dynamic TDMA FHSS DSSS OFDMA SC-FDM MC-SS Related topics Channel access methods Medium access control...
City in California, United States Not to be confused with Scott Valley. City in California, United StatesCity of Scotts ValleyCityBack view of the Scotts Valley Civic Center/City Hall and Police Department FlagSealLocation in Santa Cruz County and the state of CaliforniaCity of Scotts ValleyLocation in the United StatesCoordinates: 37°3′5″N 122°0′48″W / 37.05139°N 122.01333°W / 37.05139; -122.01333CountryUnited StatesStateCaliforniaCountySanta CruzIncorpora...
Gastone BeanBean al MilanNazionalità Italia Altezza170 cm Peso70 kg Calcio RuoloAllenatore (ex attaccante) Termine carriera1972 - giocatore CarrieraGiovanili 1946-1952 San Canzian1953-1955 Milan Squadre di club1 1952-1953 San Canzian30 (26)1955 Milan0 (0)1955-1956→ Piacenza21 (23)1956-1960 Milan87 (39)1960-1964 Genoa123 (46)1964-1967 Napoli45 (10)1967-1969 SPAL22 (2)1969-1972 Bellaria74 (18) Nazionale 1957-1958 Italia4 (0) Carriera da...
Newspaper for Edwardsville, Illinois This article needs additional citations for verification. Please help improve this article by adding citations to reliable sources. Unsourced material may be challenged and removed.Find sources: Edwardsville Intelligencer – news · newspapers · books · scholar · JSTOR (February 2015) (Learn how and when to remove this message) Edwardsville IntelligencerTypeDaily newspaperFormatBroadsheetOwner(s)Hearst CorporationPubl...
يفتقر محتوى هذه المقالة إلى الاستشهاد بمصادر. فضلاً، ساهم في تطوير هذه المقالة من خلال إضافة مصادر موثوق بها. أي معلومات غير موثقة يمكن التشكيك بها وإزالتها. (ديسمبر 2018) هذه المقالة عن قسنطينة. لمعانٍ أخرى، طالع قسنطينة (توضيح). 36°17′00″N 6°37′00″E / 36.283333°N 6.616667°E&...