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Dlya termina Arhej sm takzhe drugie znacheniya Arhe i v edinstvennom chisle arhe ya ot lat Archaea ot dr grech ἀrxaῖos izvechnyj drevnij pervozdannyj staryj domen zhivyh organizmov po tryohdomennoj sisteme Karla Vyoze naryadu s bakteriyami i eukariotami Arhei predstavlyayut soboj odnokletochnye mikroorganizmy ne imeyushie yadra a takzhe kakih libo membrannyh organell ArheiHalobacteria shtamm NRC 1 kazhdaya kletka dlinoj okolo 5 mkmNauchnaya klassifikaciyaDomen ArheiMezhdunarodnoe nauchnoe nazvanieArchaea Woese angl and angl 1990Sinonimypo dannym NCBI Archaebacteria Woeseand Fox 1977 Mendosicutes MetabacteriaTipyCrenarchaeota Krenarheoty Euryarchaeota Evriarheoty Thaumarchaeota Candidatus Aigarchaeota Candidatus Candidatus Korarchaeota Korarheoty Candidatus Lokiarchaeota Candidatus Nanoarchaeota Nanoarheoty Candidatus Candidatus Candidatus Candidatusi drugie vklyuchayaCandidatus ThorarchaeotaSistematika v VikividahIzobrazheniya na VikiskladeITIS 935939NCBI 2157EOL 7920Proslushat vvedenie v statyu source source noiconAudiozapis sozdana na osnove versii stati ot 4 oktyabrya 2012 goda Ranee arhei obedinyali s bakteriyami v obshuyu gruppu nazyvaemuyu prokarioty ili carstvo Drobyanki lat Monera i oni nazyvalis arhebakterii odnako sejchas takaya klassifikaciya schitaetsya ustarevshej ustanovleno chto arhei imeyut svoyu nezavisimuyu evolyucionnuyu istoriyu i harakterizuyutsya mnogimi biohimicheskimi osobennostyami otlichayushimi ih ot drugih form zhizni Sejchas arhei podrazdelyayut na bolee chem 7 tipov Iz nih naibolee izucheny krenarheoty Crenarchaeota i evriarheoty Euryarchaeota Klassificirovat arhei po prezhnemu slozhno tak kak podavlyayushee bolshinstvo iz nih nikogda ne vyrashivalis v laboratornyh usloviyah i identificirovalis tolko po analizu nukleinovyh kislot iz prob poluchennyh iz mest ih obitaniya Arhei i bakterii ochen pohozhi po razmeru i forme kletok hotya nekotorye arhei imeyut dovolno neobychnuyu formu naprimer kletki Haloquadratum walsbyi ploskie i kvadratnye Nesmotrya na vneshnee shodstvo s bakteriyami nekotorye geny i metabolicheskie puti arhej sblizhayut ih s eukariotami v chastnosti fermenty kataliziruyushie processy transkripcii i translyacii Drugie aspekty biohimii arhej unikalny k primeru prisutstvie v kletochnyh membranah lipidov soderzhashih prostuyu efirnuyu svyaz Bolshaya chast arhej hemoavtotrofy Oni ispolzuyut znachitelno bolshe istochnikov energii chem eukarioty nachinaya ot obyknovennyh organicheskih soedinenij takih kak sahara i zakanchivaya ammiakom ionami metallov i dazhe vodorodom Soleustojchivye arhei galoarhei Haloarchaea ispolzuyut v kachestve istochnika energii solnechnyj svet drugie vidy arhej fiksiruyut uglerod odnako v otlichie ot rastenij i cianobakterij sinezelyonyh vodoroslej ni odin vid arhej ne delaet i to i drugoe odnovremenno Razmnozhenie u arhej bespoloe binarnoe delenie fragmentaciya i pochkovanie V otlichie ot bakterij i eukariot ni odin izvestnyj vid arhej ne formiruet spor Iznachalno arhej schitali ekstremofilami zhivushimi v surovyh usloviyah goryachih istochnikah solyonyh ozyorah odnako potom ih nashli i v bolee privychnyh mestah vklyuchaya pochvu okeany bolota i tolstuyu kishku cheloveka Arhej osobenno mnogo v okeanah i vozmozhno planktonnye arhei samaya mnogochislennaya gruppa nyne zhivushih organizmov Arhei priznany vazhnoj sostavlyayushej zhizni na Zemle Oni igrayut rol v krugovorotah ugleroda i azota Ni odin iz izvestnyh predstavitelej arhej ne yavlyaetsya ni parazitom za isklyucheniem nanoarheot yavlyayushihsya parazitami drugih arhej ni patogennym organizmom odnako oni chasto byvayut mutualistami i kommensalami Nekotorye predstaviteli yavlyayutsya metanogenami i obitayut v pishevaritelnom trakte cheloveka i zhvachnyh gde oni pomogayut osushestvlyat pishevarenie Metanogeny ispolzuyutsya v proizvodstve biogaza i pri ochistke kanalizacionnyh stochnyh vod a fermenty ekstremofilnyh mikroorganizmov sohranyayushie aktivnost pri vysokih temperaturah i v kontakte s organicheskimi rastvoritelyami nahodyat svoyo primenenie v biotehnologii Istoriya otkrytiyaVpervye arhei byli obnaruzheny v ekstremalnyh mestah obitaniya goryachih vulkanicheskih istochnikah Pervye predstaviteli gruppy obnaruzheny v razlichnyh ekstremalnyh sredah obitaniya naprimer geotermalnyh istochnikah Na protyazhenii bolshej chasti XX veka prokarioty schitalis edinoj gruppoj i klassificirovalis po biohimicheskim morfologicheskim i metabolicheskim osobennostyam K primeru mikrobiologi pytalis klassificirovat mikroorganizmy v zavisimosti ot formy kletok detalej stroeniya kletochnoj stenki i potreblyaemyh mikroorganizmami veshestv V 1965 godu bylo predlozheno ustanavlivat stepen rodstva raznyh prokariot na osnovanii shodstva stroeniya ih genov Etot podhod filogenetika v nashi dni yavlyaetsya osnovnym Vpervye arhei byli vydeleny v kachestve otdelnoj gruppy prokariot na filogeneticheskom dreve v 1977 godu Karlom Vyoze i Dzhordzhem Edvardom Foksom pri sravnitelnom analize 16S rRNK Iznachalno eti dve gruppy oboznachalis kak arhebakterii lat Archaebacteria i eubakterii lat Eubacteria i rassmatrivalis kak carstva ili podcarstva kotorye Vyoze i Foks nazyvali terminom Urkingdoms Vyoze nastaival chto eta gruppa prokariot est fundamentalno otlichnyj tip zhizni Chtoby podcherknut eto otlichie vposledstvii dve gruppy prokariot byli nazvany arheyami i bakteriyami V tryohdomennoj sisteme Karla Vyoze obe eti gruppy i eukarioty byli vozvedeny v rang domena Etot termin byl predlozhen Vyoze v 1990 godu dlya oboznacheniya samogo verhnego ranga v klassifikacii organizmov vklyuchayushej odno ili neskolko carstv V pervoe vremya k novomu domenu prichislyali tolko metanogennye mikroorganizmy Schitalos chto arhei naselyayut tolko mesta s ekstremalnymi usloviyami goryachie istochniki solyonye ozyora Odnako k koncu XX veka mikrobiologi prishli k vyvodu chto arhei bolshaya i raznoobraznaya gruppa organizmov shiroko rasprostranyonnaya v prirode Mnogie vidy arhej naselyayut vpolne obychnye sredy obitaniya naprimer pochvy ili vody okeana Takaya pereocenka byla vyzvana primeneniem metoda polimeraznoj cepnoj reakcii dlya identifikacii prokariot v obrazcah vody i pochvy po ih nukleinovym kislotam Dannyj metod pozvolyaet vyyavlyat i identificirovat organizmy kotorye po tem ili inym prichinam ne kultiviruyutsya v laboratornyh usloviyah Proishozhdenie i evolyuciyaHotya vozmozhnye okamenelosti prokarioticheskih kletok datirovany vozrastom v 3 5 mlrd let bolshinstvo prokariot ne imeet harakternyh morfologicheskih osobennostej i poetomu okamenelye formy nelzya opredelyonno identificirovat imenno kak ostanki arhej V to zhe vremya himicheskie ostatki unikalnyh dlya arhej lipidov bolee informativny tak kak eti soedineniya u drugih organizmov ne vstrechayutsya V nekotoryh publikaciyah ukazyvaetsya chto ostanki lipidov arhej ili eukariot prisutstvuyut v porodah vozrastom 2 7 mlrd let odnako dostovernost etih dannyh ostayotsya pod somneniem Eti lipidy obnaruzheny v dokembrijskih formaciyah Drevnejshie iz podobnyh ostatkov najdeny v na zapade Grenlandii gde nahodyatsya samye starye na Zemle osadochnye porody sformirovavshiesya 3 8 mlrd let nazad Arhei mogut byt drevnejshimi zhivymi sushestvami naselyayushimi Zemlyu Vyoze utverzhdal chto arhei bakterii i eukarioty predstavlyayut soboj tri razdelnye linii rano otdelivshiesya ot obshej predkovoj gruppy organizmov Vozmozhno eto proizoshlo eshyo do kletochnoj evolyucii kogda otsutstvie tipichnoj kletochnoj membrany davalo vozmozhnosti k neogranichennomu gorizontalnomu perenosu genov i predki tryoh domenov razlichalis mezhdu soboj po fiksiruemym komplektam genov Ne isklyucheno chto poslednij obshij predok arhej i bakterij byl termofilom eto dayot osnovaniya predpolozhit chto nizkie temperatury byli ekstremalnoj sredoj dlya arhej i organizmy prisposobivshiesya k nim poyavilis tolko pozzhe Sejchas arhei i bakterii svyazany mezhdu soboj ne bolshe chem s eukariotami i termin prokarioty oboznachaet lish ne eukarioty chto ogranichivaet ego primenimost Sravnitelnaya harakteristika arhej i drugih domenov V privedyonnoj tablice pokazany nekotorye cherty arhej svojstvennye i ne svojstvennye drugim domenam Mnogie iz etih svojstv takzhe obsuzhdayutsya nizhe Svojstvenno arheyam i bakteriyam Svojstvenno arheyam i eukariotam Svojstvenno tolko arheyamNet oformlennogo yadra i membrannyh organell Net peptidoglikana mureina Struktura kletochnoj stenki k primeru kletochnye stenki nekotoryh arhej soderzhat Kolcevaya hromosoma DNK svyazana s gistonami V kletochnoj membrane prisutstvuyut lipidy soderzhashie prostuyu efirnuyu svyazGeny obedineny v operony Translyaciya nachinaetsya s metionina Struktura flagellinovShozhie RNK polimeraza promotory i drugie komponenty transkripcionnogo kompleksa est introny i processing RNK Struktura ribosom nekotorye priznaki sblizhayut s bakteriyami nekotorye s eukariotami Policistronnaya mRNK Shozhie replikaciya i reparaciya DNK Struktura i metabolizm tRNKRazmer kletok na neskolko poryadkov menshe chem u eukariot Shozhaya ATFaza tip V Rodstvo s drugimi prokariotami Ustanovlenie stepeni rodstva mezhdu tremya domenami imeet klyuchevoe znachenie dlya ponimaniya vozniknoveniya zhizni Bolshinstvo metabolicheskih putej v kotoryh zadejstvovana bolshaya chast genov organizma shozhi u bakterij i arhej v to vremya kak geny otvechayushie za ekspressiyu drugih genov ochen pohozhi u arhej i eukariot Po stroeniyu kletok arhei naibolee blizki k grampolozhitelnym bakteriyam kletka pokryta edinstvennoj plazmaticheskoj membranoj dopolnitelnaya vneshnyaya membrana harakternaya dlya gramotricatelnyh bakterij otsutstvuet kletochnye stenki razlichnogo himicheskogo sostava kak pravilo tolstye V filogeneticheskom dreve osnovannom na sravnitelnom analize struktur gomologichnyh genov belkov prokariot gomologi arhej naibolee blizki k takovym grampolozhitelnyh bakterij V nekotoryh vazhnejshih belkah arhej i grampolozhitelnyh bakterij takih kak Hsp70 i glutaminovaya sintetaza I obnaruzhivayutsya odinakovye evolyucionno konservativnye vstavki i delecii Gupta angl Gupta predpolozhil chto arhei otdelilis ot grampolozhitelnyh bakterij v rezultate otbora po priznaku ustojchivosti k dejstviyu antibiotikov Eto osnovyvaetsya na nablyudenii chto arhei ustojchivy k ogromnomu chislu antibiotikov v osnovnom proizvodimyh grampolozhitelnymi bakteriyami i chto eti antibiotiki dejstvuyut glavnym obrazom na geny kotorye otlichayut bakterij ot arhej Soglasno gipoteze Gupty davlenie otbora v napravlenii formirovaniya ustojchivosti k antibiotikam grampolozhitelnyh bakterij v konce koncov privelo k sushestvennym izmeneniyam v strukture genov mishenej antibiotikov u nekotoryh mikroorganizmov kotorye stali obshimi predkami sovremennyh arhej Predpolagaemaya evolyuciya arhej pod dejstviem antibiotikov i drugih neblagopriyatnyh faktorov takzhe mozhet obyasnit ih adaptaciyu k ekstremalnym usloviyam takim kak povyshennye temperatura i kislotnost kak rezultat poiska nish svobodnyh ot produciruyushih antibiotiki organizmov T Kavalir Smit vydvinul shozhee predpolozhenie Versiya Gupty takzhe podtverzhdaetsya drugimi rabotami izuchayushimi rodstvennye cherty v strukturah belkov i issledovaniyami pokazavshimi chto grampolozhitelnye bakterii mogli byt pervoj vetvyu otdelivshejsya ot obshego dreva prokariot Rodstvo s eukariotami Evolyucionnoe rodstvo mezhdu arheyami i eukariotami ostayotsya neyasnym Pomimo shodstva v strukture i funkciyah kletok mezhdu nimi sushestvuet shodstvo na geneticheskom urovne Ustanovleno chto gruppa arhej krenarheoty stoyat blizhe k eukariotam chem k drugomu tipu arhej evriarheotam Krome togo u nekotoryh bakterij kak obnaruzheny arheepodobnye geny peredannye putyom gorizontalnogo perenosa Naibolee rasprostranena gipoteza soglasno kotoroj predok eukariot rano otdelilsya ot arhej a eukarioty voznikli v rezultate sliyaniya arhei i eubakterii stavshih citoplazmoj i yadrom novoj kletki Eta gipoteza obyasnyaet razlichnye geneticheskie shodstva no stalkivaetsya s trudnostyami v obyasnenii kletochnoj struktury StroenieForma kletok i kolonij Otdelnye kletki arhej dostigayut ot 0 1 do 15 mkm v diametre i mogut imet razlichnuyu formu shara palochki spirali ili diska Nekotorye krenarheoty imeyut druguyu formu naprimer nepravilnoj dolchatoj formy tonkoj nitevidnoj formy i menshe 1 mkm v diametre a i pochti idealno pryamougolnye Haloquadratum walsbyi ploskie kvadratnye arhei zhivushie v sverhsolyonyh vodoyomah Takie neobychnye formy kletok veroyatno obespechivayutsya kletochnoj stenkoj i prokarioticheskim citoskeletom U arhej obnaruzheny belki rodstvennye komponentam citoskeleta drugih organizmov a takzhe pokazano prisutstvie filamentov v ih kletkah odnako u arhej v otlichie ot drugih organizmov eti struktury ploho izucheny U Thermoplasma i Ferroplasma kletochnaya stenka otsutstvuet poetomu ih kletki imeyut nepravilnuyu formu i pohozhi na amyob Kletki nekotoryh vidov arhej mogut obedinyatsya v agregaty i filamenty dlinoj do 200 mkm Eti organizmy mogut formirovat bioplyonki V kulturah kletki slivayutsya drug s drugom formiruya odnu krupnuyu kletku Arhei roda obrazuyut slozhnye mnogokletochnye kolonii v kotoryh kletki obedineny s pomoshyu dlinnyh tonkih polyh trubok nazyvaemyh cannulae kotorye vystupayut nad poverhnostyami kletok i sobirayut ih v gustoe kustovidnoe skoplenie Funkcii etih trubok ne yasny no vozmozhno oni osushestvlyayut kommunikaciyu i obmen pitatelnymi veshestvami mezhdu sosednimi kletkami Sushestvuyut i mnogovidovye kolonii kak naprimer nit zhemchuga obnaruzhennaya v 2001 godu v bolote v Germanii Kruglye belovatye kolonii nekotoryh neobychnyh evriarheot peremezhayutsya tonkimi nityami kotorye mogut dostigat do 15 sm v dlinu i sostoyat iz osobyh vidov bakterij Arhei i bakterii imeyut ochen pohozhuyu strukturu kletok odnako ih sostav i organizaciya otdelyayut arhej ot bakterij Kak u bakterij u nih otsutstvuyut vnutrennie membrany i organelly kletochnye membrany kak pravilo ogranicheny kletochnoj stenkoj a plavanie osushestvlyaetsya za schyot odnogo ili bolee zhgutikov Strukturno arhei naibolee shozhi s grampolozhitelnymi bakteriyami Bolshinstvo imeet odnu plazmaticheskuyu membranu i kletochnuyu stenku periplazmaticheskoe prostranstvo otsutstvuet Isklyucheniem iz etogo glavnogo pravila yavlyaetsya u kotorogo krupnoe periplazmaticheskoe prostranstvo ogranichennoe naruzhnoj membranoj soderzhit okruzhyonnye membranoj vezikuly Membrany Struktura membran Vverhu fosfolipidy arhej 1 izoprenovye cepochki 2 prostye efirnye svyazi 3 ostatok L glicerina 4 fosfatnaya gruppa Poseredine bakterialnye ili eukarioticheskie fosfolipidy 5 cepochki zhirnyh kislot 6 slozhnoefirnye svyazi 7 ostatok D glicerina 8 fosfatnaya gruppa Snizu 9 lipidnyj bisloj bakterij i eukariot 10 lipidnyj monosloj nekotoryh arhej Molekuly iz kotoryh postroeny membrany arhej silno otlichayutsya ot teh kotorye ispolzuyutsya v membranah drugih organizmov Eto ukazyvaet na to chto arhei sostoyat lish v otdalyonnom rodstve s bakteriyami i eukariotami U vseh zhivyh organizmov kletochnye membrany postroeny iz fosfolipidov Molekuly fosfolipidov sostoyat iz dvuh chastej gidrofilnoj polyarnoj sostoyashej iz fosfatov i gidrofobnoj nepolyarnoj sostoyashej iz lipidov Eti komponenty obedineny cherez ostatok glicerina V vode molekuly fosfolipidov klasterizuyutsya pri etom fosfatnye golovki okazyvayutsya obrashyonnymi k vode a lipidnye hvosty obrashyonnymi ot neyo i spryatannymi vnutr klastera Glavnaya sostavlyayushaya membrany dva sloya takih fosfolipidov nazyvaemye lipidnym bisloem Eti fosfolipidy u arhej obladayut chetyrmya neobychnymi chertami U bakterij i eukariot membrany sostoyat glavnym obrazom iz glicerin slozhnoefirnyh lipidov togda kak u arhej oni slozheny iz glicerin efirnyh lipidov Razlichaetsya tip svyazi mezhdu ostatkami lipidov i glicerina Svyazi dvuh tipov oboznacheny zhyoltym na sheme sprava V slozhnoefirnyh lipidah svyaz slozhnoefirnaya a v efirnyh efirnaya Efirnye svyazi himicheski bolee stojkie chem slozhnoefirnye Eta stabilnost pomogaet arheyam vyzhivat pri vysokih temperaturah a takzhe v silnokislyh i silnoshelochnyh sredah Bakterii i eukarioty soderzhat nekotoroe kolichestvo efirnyh lipidov no po sravneniyu s arheyami oni ne yavlyayutsya glavnoj sostavlyayushej membran Imeetsya otlichie v stereohimii u arhej asimmetricheskij centr glicerinovoj sostavlyayushej imeet L konfiguraciyu a ne D kak u drugih organizmov Poetomu dlya sinteza fosfolipidov arhei ispolzuyut sovershenno drugie fermenty chem bakterii i eukarioty Takie fermenty poyavilis ochen rano v istorii zhizni chto ukazyvaet na to chto arhei rano otdelilis ot dvuh drugih domenov Lipidnye hvosty arhej himicheski otlichny ot takovyh u drugih organizmov Osnovu lipidov arhej sostavlyaet izoprenoidnaya bokovaya cep i ih lipidy predstavlyayut soboj dlinnye cepi s mnozhestvom pobochnyh vetvej inogda dazhe s ciklopropanovymi i ciklogeksanovymi kolcami Hotya izoprenoidy igrayut vazhnuyu rol v biohimii mnogih organizmov tolko arhei ispolzuyut ih dlya sozdaniya fosfolipidov Predpolagayut chto eti razvetvlyonnye cepi kak i efirnye svyazi sluzhat dlya prisposobleniya k obitaniyu pri vysokih temperaturah Ustanovleno chto izoprenoidnye membrany sohranyayut v shirokom diapazone temperatur 0 100 C zhidkokristallicheskoe sostoyanie chto neobhodimo dlya ih normalnogo biologicheskogo funkcionirovaniya Pronicaemost takih membran dlya ionov i nizkomolekulyarnyh organicheskih veshestv takzhe malo izmenyaetsya s povysheniem temperatury v otlichie ot membran iz obychnyh lipidov u kotoryh ona rezko vozrastaet U nekotoryh arhej lipidnyj bisloj zamenyaetsya monosloem Fakticheski pri etom lipidnye hvosty dvuh raznyh fosfolipidnyh molekul slivayutsya s obrazovaniem odnoj molekuly s dvumya polyarnymi golovkami Eti sliyaniya delayut membranu bolee stojkoj i luchshe prisposoblennoj dlya surovyh uslovij K primeru ferroplazma imeet lipidy etogo tipa i oni pomogayut ej vyzhivat v silnokislyh usloviyah Kletochnaya stenka Bolshinstvo arhej no ne Thermoplasma i Ferroplasma obladayut kletochnoj stenkoj U bolshej chasti iz nih ona sformirovana molekulami poverhnostnyh belkov obrazuyushih naruzhnyj S sloj S sloj predstavlyaet soboj zhyostkuyu setku iz belkovyh molekul pokryvayushih kletku snaruzhi podobno kolchuge Etot sloj zashishaet kletku ot fizicheskih i himicheskih vozdejstvij a takzhe predotvrashaet kontakt makromolekul s kletochnoj membranoj V otlichie ot bakterij kletochnaya stenka arhej ne soderzhit peptidoglikan Metanobakterii lat Methanobacteriales imeyut kletochnye stenki soderzhashie kotoryj napominaet peptidoglikan eubakterij po morfologii funkcii i fizicheskoj strukture no otlichen po himicheskoj v nyom net ostatkov D aminokislot i N acetilmuramovoj kisloty Zhgutiki Osnovnaya statya Arhellum Zhgutik arhej inogda nazyvayut arhellum Zhgutiki arhej rabotayut tak zhe kak i u bakterij ih dlinnye niti privodyatsya v dvizhenie vrashatelnym mehanizmom v osnovanii zhgutika Etot mehanizm rabotaet za schyot transmembrannogo protonnogo gradienta Tem ne menee zhgutiki arhej znachitelno otlichayutsya ot bakterialnyh po stroeniyu i sposobu sborki Dva tipa zhgutikov razvilis iz raznyh predkovyh struktur Bakterialnyj zhgutik i sistema sekrecii III tipa imeli obshuyu predkovuyu strukturu a arhejnyj zhgutik proizoshyol ot bakterialnyh pilej IV tipa Zhgutik bakterij polyj i sobiraetsya iz subedinic kotorye prohodyat vverh po centralnoj pore k koncu zhgutika Zhgutiki zhe arhej stroyatsya putyom dobavleniya subedinic v ih osnovanie Krome togo v otlichie ot bakterialnyh zhgutikov v zhgutiki arhej vhodit neskolko vidov flagellinov MetabolizmArhei demonstriruyut ogromnoe raznoobrazie himicheskih reakcij protekayushih v ih kletkah v processe metabolizma a takzhe istochnikov energii Eti reakcii klassificiruyutsya po gruppam pitaniya v zavisimosti ot istochnikov energii i ugleroda Nekotorye arhei poluchayut energiyu iz neorganicheskih soedinenij takih kak sera ili ammiak oni yavlyayutsya litotrofami K nim otnosyatsya nitrificiruyushie arhei metanogeny i anaerobnye metanookisliteli V etih reakciyah odno soedinenie otdayot elektrony drugomu okislitelno vosstanovitelnye reakcii a vydelyayushayasya pri etom energiya sluzhit toplivom dlya osushestvleniya razlichnyh kletochnyh processov Soedinenie otdayushee elektrony nazyvaetsya donorom a prinimayushee akceptorom Vydelyayushayasya energiya idyot na obrazovanie ATF putyom hemiosmosa V sushnosti eto osnovnoj process protekayushij v mitohondriyah eukarioticheskih kletok Drugie gruppy arhej ispolzuyut v kachestve istochnika energii solnechnyj svet ih nazyvayut fototrofami Odnako ni odin iz etih organizmov ne obrazuet kislorod v processe fotosinteza Mnogie bazovye metabolicheskie processy yavlyayutsya obshimi dlya vseh form zhizni naprimer arhei ispolzuyut modificirovannyj variant glikoliza put Entnera Dudorova a takzhe polnyj ili chastichnyj cikl Krebsa trikarbonovyh kislot Eto veroyatno otrazhaet rannee vozniknovenie etih putej v istorii zhizni i ih vysokuyu effektivnost Tipy pitaniya arhej Tip pitaniya Istochnik energii Istochnik ugleroda PrimeryFototrofy Solnechnyj svet Organicheskie soedineniya HalobacteriaLitotrofy Neorganicheskie soedineniya Organicheskie soedineniya ili fiksaciya ugleroda Methanobacteria Organotrofy Organicheskie soedineniya Organicheskie soedineniya ili fiksaciya ugleroda Methanosarcinales Nekotorye evriarheoty yavlyayutsya metanogenami i obitayut v anaerobnyh sredah takih kak bolota Takoj tip metabolizma poyavilsya rano i vozmozhno dazhe chto pervyj svobodnozhivushij organizm byl metanogenom Obychnaya dlya etih organizmov biohimicheskaya reakciya predstavlyaet soboj okislenie vodoroda s ispolzovaniem uglekislogo gaza v kachestve akceptora elektronov Dlya osushestvleniya metanogeneza neobhodimo mnozhestvo razlichnyh kofermentov unikalnyh dlya etih arhej takih kak koferment M i metanofuran Nekotorye organicheskie soedineniya takie kak spirty uksusnaya i muravinaya kisloty mogut ispolzovatsya metanogenami v kachestve alternativnyh akceptorov elektronov Podobnye reakcii protekayut u arhej zhivushih v pishevaritelnom trakte U acidotrofnyh arhej uksusnaya kislota raspadaetsya neposredstvenno na metan i uglekislyj gaz Takie acidotrofnye arhei otnosyatsya k otryadu Methanosarcinales Oni yavlyayutsya vazhnoj sostavlyayushej soobshestv mikroorganizmov produciruyushih biogaz Drugie arhei ispolzuyut atmosfernyj uglekislyj gaz kak istochnik ugleroda blagodarya processu fiksacii ugleroda to est yavlyayutsya avtotrofami Etot process vklyuchaet v sebya libo silno izmenyonnyj cikl Kalvina libo nedavno otkrytyj metabolicheskij put izvestnyj kak 3 gidroksilpropionat 4 gidroksibutiratnyj cikl Krenarheoty takzhe ispolzuyut obratnyj cikl Krebsa a evriarheoty vosstanovitelnyj acetil SoA process Fiksaciya ugleroda osushestvlyaetsya za schyot energii poluchaemoj iz neorganicheskih soedinenij Ni odin izvestnyj vid arhej ne fotosinteziruet Istochniki energii kotorye ispolzuyut arhei chrezvychajno raznoobrazny nachinaya ot okisleniya ammiaka do okisleniya serovodoroda ili elementarnoj sery provodimogo pri etom v kachestve akceptorov elektronov mogut ispolzovatsya kislorod ili iony metallov Bakteriorodopsin Retinolovyj kofaktor i ostatki osushestvlyayushie perenos protonov predstavleny v vide sharo sterzhnevoj modeli Fototrofnye arhei ispolzuyut solnechnyj svet dlya polucheniya himicheskoj energii v vide ATF U Halobacteria aktiviruemye svetom ionnye nasosy kak bakteriorodopsin i sozdayut ionnyj gradient putyom vykachivaniya ionov iz kletki cherez plazmaticheskuyu membranu Zapasyonnaya v etom elektrohimicheskom gradiente energiya preobrazuetsya v ATF s pomoshyu ATF sintazy Etot process predstavlyaet soboj formu fotofosforilirovaniya Sposobnost etih nasosov perenosit iony cherez membrany pri osveshenii obuslovlena izmeneniyami kotorye proishodyat v strukture retinolovogo kofaktora skrytogo v centre belka pod dejstviem sveta GenetikaKak pravilo arhei imeyut odinochnuyu kolcevuyu hromosomu razmer kotoroj mozhet dostigat 5 751 492 par nukleotidov u Methanosarcina acetivorans obladayushej samym bolshim izvestnym genomom sredi arhej Odnu desyatuyu razmera etogo genoma sostavlyaet genom s 490 885 parami nukleotidov u Nanoarchaeum equitans imeyushego samyj malenkij izvestnyj genom sredi arhej on soderzhit lish 537 genov kodiruyushih belki Takzhe u arhej obnaruzheny bolee melkie nezavisimye molekuly DNK tak nazyvaemye plazmidy Vozmozhno plazmidy mogut peredavatsya mezhdu kletkami pri fizicheskom kontakte v hode processa shodnogo s konyugaciej bakterij porazhyonnyj DNK virusom STSV1 Dlina otrezka 1 mkm Arhei mogut porazhatsya virusami soderzhashimi dvuhcepochechnuyu DNK Virusy arhej chasto nerodstvenny drugim gruppam virusov i imeyut razlichnye neobychnye formy vklyuchaya butylki kryuchki i kapli Eti virusy byli tshatelno izucheny na termofilah v osnovnom otryadov Sulfolobales i Thermoproteales V 2009 godu byl otkryt virus soderzhashij odnocepochechnuyu DNK i porazhayushij galofilnye arhei Zashitnye reakcii arhej protiv virusov mogut vklyuchat mehanizm blizkij k RNK interferencii eukariot Arhei geneticheski otlichny ot eukariot i bakterij prichyom do 15 belkov kodiruemyh odnim genomom arhei unikalny dlya etogo domena hotya funkcii bolshinstva etih belkov neizvestny Bolshaya chast unikalnyh belkov funkciya kotoryh izvestna prinadlezhit evriarheotam i zadejstvovana v metanogeneze Belki obshie dlya arhej bakterij i eukariot uchastvuyut v osnovnyh kletochnyh funkciyah i kasayutsya v osnovnom transkripcii translyacii i metabolizma nukleotidov K drugim osobennostyam arhej mozhno otnesti organizaciyu genov vypolnyayushih svyazannye funkcii k primeru geny otvetstvennye za raznye etapy odnogo i togo zhe metabolicheskogo processa v operony i bolshie otlichiya v stroenii genov tRNK i ih aminoacil tRNK sintetaz Transkripciya i translyaciya arhej bolshe napominayut eti processy v kletkah eukariot chem bakterij prichyom RNK polimeraza i ribosomy arhej ochen blizki k analogichnym strukturam u eukariot Hotya u arhej est lish odin tip RNK polimerazy po stroeniyu i funkcii v transkripcii ona blizka k RNK polimeraze II eukariot pri etom shozhie gruppy belkov glavnye faktory transkripcii obespechivayut svyazyvanie RNK polimerazy s promotorom gena V to zhe vremya drugie faktory transkripcii arhej bolee blizki k takovym u bakterij Processing RNK u arhej proshe chem u eukariot tak kak bolshinstvo genov arhej ne soderzhit intronov hotya v genah ih tRNK i rRNK ih dostatochno mnogo takzhe oni prisutstvuyut v nebolshom kolichestve genov kodiruyushih belki RazmnozhenieArhei razmnozhayutsya bespolym putyom binarnym ili mnozhestvennym deleniem fragmentaciej ili pochkovaniem Mejoza ne proishodit poetomu dazhe esli predstaviteli konkretnogo vida arhej sushestvuyut bolee chem v odnoj forme vse oni imeyut odinakovyj geneticheskij material Kletochnoe delenie opredelyaetsya kletochnym ciklom posle togo kak hromosoma replicirovalas i dve dochernie hromosomy razoshlis kletka delitsya Detali izucheny lish u roda no osobennosti ego cikla ochen shozhi s takovymi i u eukariot i u bakterij Replikaciya hromosom nachinaetsya s mnozhestvennyh tochek nachala replikacii s pomoshyu DNK polimerazy pohozhej na analogichnye fermenty eukariot Odnako belki upravlyayushie kletochnym deleniem takie kak FtsZ kotorye formiruyut szhimayushee kolco vokrug kletki i komponenty septy prohodyashej cherez centr kletki shozhi s ih bakterialnymi ekvivalentami Arhei ne obrazuyut spory Nekotorye vidy mogut preterpevat smenu fenotipa i sushestvovat kak kletki neskolkih razlichnyh tipov vklyuchaya tolstostennye kletki ustojchivye k osmoticheskomu shoku i pozvolyayushie arheyam vyzhivat v vode s nizkoj koncentraciej soli Odnako eti struktury ne sluzhat dlya razmnozheniya a skoree pomogayut arheyam osvaivat novye sredy obitaniya EkologiyaPyatno cvetnyh termofilnyh arhej Jelloustonskij nacionalnyj park SShA Sredy obitaniya Arhei zhivut v shirokom diapazone sred obitaniya i yavlyayutsya vazhnoj chastyu globalnoj ekosistemy mogut sostavlyat do 20 obshej biomassy Pervye otkrytye arhei byli ekstremofilami Dejstvitelno mnogie arhei vyzhivayut pri vysokih temperaturah chasto svyshe 100 C i obnaruzheny v gejzerah chyornyh kurilshikah i maslosbornikah Drugie prisposobilis k zhizni v ochen holodnyh usloviyah v silnosolyonyh silnokislyh i silnoshelochnyh sredah a takzhe pri vysokom davlenii do 700 atmosfer barofily Odnako sredi arhej est i mezofily obitayushie v myagkih usloviyah v bolotistyh mestnostyah stochnyh vodah okeanah i pochve Ekstremofilnye arhei otnosyatsya k chetyryom glavnym fiziologicheskim gruppam galofilam termofilam acidofilam kislotoustojchivye i alkalifilam shelocheustojchivye Eti gruppy nelzya rassmatrivat v range tipa ili kak drugie samostoyatelnye taksony Oni ne vzaimoisklyuchayut drug druga i nekotorye arhei otnosyat odnovremenno k neskolkim gruppam Tem ne menee oni yavlyayutsya udobnoj startovoj tochkoj dlya klassifikacii Galofily vklyuchaya rod zhivut v ekstremalno solyonyh sredah takih kak solyonye ozyora i pri mineralizacii bolshe 20 25 prevoshodyat po chislennosti svoih sosedej bakterij Termofily luchshe vsego rastut na temperaturah svyshe 45 C v takih mestah kak goryachie istochniki dlya gipertermofilov optimalnaya temperatura 80 C i vyshe Methanopyrus kandleri shtamm 116 rastyot pri 122 C rekordno vysokoj temperature dlya vseh organizmov Drugie arhei obitayut v ochen kislyh ili shelochnyh sredah Naprimer naibolee ustojchivyj acidofil rastyot pri pH 0 chto ekvivalentno 1 2 molyarnoj sernoj kislote Ustojchivost k ekstremalnym usloviyam vneshnej sredy sdelala arhej centralnoj temoj v obsuzhdeniyah vozmozhnyh svojstv zhizni na drugih planetah Nekotorye sredy v kotoryh obitayut ekstremofily ne silno otlichayutsya ot takovyh na Marse chto navodit na mysl o vozmozhnom perenose takih ustojchivyh mikroorganizmov mezhdu planetami na meteoritah Nedavno neskolko rabot pokazali chto arhei obitayut ne tolko v termofilnyh i mezofilnyh usloviyah no takzhe vstrechayutsya inogda v bolshom kolichestve i v mestah s nizkimi temperaturami Naprimer arhei vstrechayutsya v holodnyh vodah takih kak polyarnye morya Eshyo bolee vazhno chto ogromnoe kolichestvo arhej obnaruzheno povsemestno v okeanah v neekstremalnyh usloviyah v sostave planktona kak chast pikoplanktona Hotya eti arhei mogut prisutstvovat v poistine kolossalnom kolichestve do 40 ot obshej biomassy mikrobov pochti ni odin iz etih vidov ne byl izolirovan vyrashen i izuchen v chistoj kulture Poetomu nashe ponimanie roli arhej v ekologii okeana ih vliyaniya na globalnyj biogeohimicheskij krugovorot ostayotsya v znachitelnoj mere nepolnym Nekotorye morskie krenarheoty sposobny k nitrifikacii poetomu veroyatno chto oni okazyvayut vliyanie na okeanicheskij krugovorot azota hotya eti okeanicheskie krenarheoty mogut ispolzovat i drugie istochniki energii Bolshoe chislo arhej takzhe obnaruzheno v osadke pokryvayushem okeanicheskoe dno prichyom oni sostavlyayut bolshinstvo zhivyh kletok na glubine bolshe 1 m ot urovnya okeanicheskogo dna Rol v krugovorote veshestv Arhei vtorichno ispolzuyut takie elementy kak uglerod azot i seru v svoih razlichnyh sredah obitaniya Hotya takie prevrasheniya neobhodimy dlya normalnogo funkcionirovaniya ekosistemy arhei mogut takzhe sodejstvovat vrednym izmeneniyam vyzvannym deyatelnostyu cheloveka i dazhe vyzvat zagryaznenie Arhei osushestvlyayut mnogie etapy krugovorota azota Eto vklyuchaet v sebya kak reakcii udalyayushie azot iz ekosistemy k primeru azotnoe dyhanie i denitrifikaciya tak i processy v hode kotoryh pogloshaetsya azot takie kak usvoenie nitratov i fiksaciya azota Nedavno byla otkryta prichastnost arhej k okisleniyu ammiaka Eti reakcii osobenno vazhny v okeanah Arhei takzhe igrayut vazhnuyu rol v pochvennom okislenii ammiaka Oni obrazuyut nitrity kotorye zatem okislyayutsya drugimi mikrobami v nitraty Poslednie potreblyayutsya rasteniyami i drugimi organizmami V krugovorote sery arhei zhivushie za schyot okisleniya soedinenij sery poluchayut ih iz kamenistyh porod i delayut ih dostupnymi dlya drugih organizmov Odnako vidy osushestvlyayushie eto takie kak obrazuyut sernuyu kislotu kak pobochnyj produkt i sushestvovanie takih organizmov v zabroshennyh shahtah mozhet sovmestno s kislotnymi shahtnymi vodami prichinit vred okruzhayushej srede V krugovorote ugleroda metanogeny udalyayut vodorod i igrayut vazhnuyu rol v razlozhenii organicheskoj materii populyaciyami mikroorganizmov vystupayushih kak razlagateli v anaerobnyh ekosistemah takih kak ily bolota i vodoochistnye sooruzheniya Odnako metan odin iz samyh rasprostranyonnyh gazov v zemnoj atmosfere vyzyvayushih parnikovyj effekt dostigaya 18 ot obshego obyoma parnikovyh gazov On v 25 raz bolee effektiven po sposobnosti vyzyvat parnikovyj effekt chem uglekislyj gaz Metanogeny glavnyj istochnik atmosfernogo metana vydelyayut bolshuyu chast ezhegodnogo vybrosa metana Poetomu eti arhei prichastny k sozdaniyu parnikovogo effekta na Zemle i globalnomu potepleniyu Vzaimodejstvie s drugimi organizmami Metanoobrazuyushie arhei vstupayut v simbioz s termitami Horosho izuchennye otnosheniya mezhdu arheyami i drugimi organizmami mutualizm i kommensalizm Poka ne sushestvuet chyotkih dokazatelstv sushestvovaniya patogennyh ili paraziticheskih vidov arhej Odnako byla predpolozhena svyaz mezhdu nekotorymi vidami metanogenov i infekciyami polosti rta Krome togo vid Nanoarchaeum equitans vozmozhno yavlyaetsya parazitom drugogo vida arhej poskolku on vyzhivaet i razmnozhaetsya tolko na kletkah krenarheota i ne prinosit nikakoj ochevidnoj vygody svoemu hozyainu S drugoj storony arheepodobnye acidofilnye nanoorganizmy Richmondskih rudnikov ARMAN inogda prikreplyayutsya k kletkam drugih arhej v bioplyonkah kislyh stochnyh vod rudnikov Priroda etogo vzaimodejstviya ne yasna no v otlichie ot sluchaya Nanorchaeaum Ignicoccus sverhmelkie kletki ARMAN vsegda ostayutsya nezavisimymi ot kletok Thermoplasmatales Mutualizm Odin iz horosho ponyatnyh primerov mutualizma vzaimodejstvie prostejshih i metanoobrazuyushih arhej obitayushih v pishevaritelnom trakte zhivotnyh sposobnyh perevarivat cellyulozu takih kak zhvachnye i termity V etih anaerobnyh usloviyah prostejshie razlagayut cellyulozu dlya polucheniya energii V etom processe v kachestve pobochnogo produkta osvobozhdaetsya vodorod odnako vysokij ego uroven sokrashaet poluchenie energii Metanogeny prevrashayut vodorod v metan i prostejshie mogut dalshe normalno poluchat energiyu V sluchae anaerobnyh prostejshih vrode arhei zhivut vnutri kletki prostejshego i potreblyayut vodorod obrazuemyj v ego gidrogenosomah Arhei takzhe vzaimodejstvuyut i s bolee krupnymi organizmami Naprimer morskaya arheya zhivyot vnutri kak endosimbiont gubki Kommensalizm Arhei mogut byt kommensalami to est sushestvovat sovmestno s drugim organizmom ne prinosya emu ni polzy ni vreda no s vygodoj dlya sebya K primeru metanogen naibolee tipichnyj predstavitel arhej v mikroflore cheloveka Kazhdyj desyatyj prokariot v chelovecheskom pishevaritelnom trakte prinadlezhit k etomu vidu V pishevaritelnom trakte termitov i cheloveka eti metanogeny v dejstvitelnosti mogut byt mutualistami vzaimodejstvuyushimi s drugimi mikrobami pishevaritelnogo trakta i sposobstvuyushimi pishevareniyu Arhei takzhe vzaimodejstvuyut s drugimi organizmami k primeru zhivut na vneshnej poverhnosti korallov i v chasti pochvy prilegayushej k kornyam rastenij rizosfere KlassifikaciyaFilogeneticheskoe drevo postroennoe na osnovanii analiza rRNK pokazyvaet razdelenie bakterij arhej i eukariotOsnovnaya statya Sistematika arhej Klassifikaciya arhej kak i prokariot v celom bystro menyaetsya i vo mnogom ostayotsya spornoj Sovremennye sistemy klassifikacii stremyatsya obedinit arhei v gruppy organizmov so shozhimi strukturnymi svojstvami i obshimi predkami Eti klassifikacii osnovany na analize struktury genov rRNK dlya ustanovleniya rodstvennyh otnoshenij mezhdu organizmami molekulyarnaya filogenetika Bolshuyu chast arhej vyrashivaemyh v laboratoriyah i horosho izuchennyh otnosyat k dvum glavnym tipam krenarheoty Crenarchaeota i evriarheoty Euryarchaeota Drugie gruppy byli vydeleny v poryadke rabochej gipotezy Naprimer dovolno neobychnyj vid Nanoarchaeum equitans otkrytyj v 2003 godu byl vydelen v samostoyatelnyj tip Nanoarchaeota Byl takzhe predlozhen novyj tip Korarchaeota On obedinyaet nebolshuyu gruppu termofilnyh vidov obladayushih osobennostyami oboih osnovnyh tipov no bolee rodstvenno blizkih k krenarheotam Drugie nedavno otkrytye vidy imeyut lish dalnee rodstvo s vysheperechislennymi gruppami naprimer arhejnye acidofilnye nanoorganizmy Richmondskih rudnikov ARMAN otkrytye v 2006 godu i yavlyayushiesya odnimi iz samyh melkih izvestnyh na segodnyashnij moment organizmov ARMAN novaya gruppa arhej obitayushaya v Razdelenie arhej na vidy takzhe sporno V biologii vid opredelyaetsya kak gruppa blizkorodstvennyh organizmov Obychnyj kriterij kotorym polzuyutsya v podobnyh situaciyah organizmy odnogo vida mogut skreshivatsya drug s drugom no ne s osobyami drugih vidov v dannom sluchae ne rabotaet poskolku arhei razmnozhayutsya tolko bespolym putyom Arhei demonstriruyut vysokij uroven gorizontalnogo perenosa genov mezhdu liniyami Nekotorye issledovateli predpolagayut chto osobi mozhno obedinyat v populyacii pohozhie na vidy pri uslovii vysokoj stepeni shozhesti ih genomov i redko sluchayushegosya perenosa genov mezhdu organizmami s menee shozhimi genomami kak v sluchae roda ferroplazma Ferroplasma S drugoj storony izuchenie roda pokazalo sushestvovanie znachimoj peredachi genov mezhdu dalnerodstvennymi populyaciyami chto ogranichivaet primenimost etogo kriteriya Vtoraya problema sostoit v tom kakoe prakticheskoe znachenie mozhet imet podobnoe razdelenie na vidy Sovremennye dannye o geneticheskom raznoobrazii arhej fragmentarny i obshee chislo ih vidov ne mozhet byt oceneno s kakoj libo tochnostyu Sravnitelnyj analiz struktur 16S rRNK arhej pozvolil predpolozhit sushestvovanie 18 23 filogeneticheskih grupp urovnya tipov prichyom predstaviteli lish vosmi grupp vyrasheny neposredstvenno v laboratorii i izucheny s uchyotom veroyatnoj polifilii nekotoryh vydelyaemyh v dannyj moment tipov Mnogie iz etih gipoteticheskih grupp izvestny lish po odnoj posledovatelnosti rRNK chto govorit o tom chto predely raznoobraziya etih organizmov ostayutsya neyasnymi Mnogie bakterii takzhe nikogda ne kultivirovalis v laboratorii chto privodit k shozhim problemam pri ih harakteristike Po ustoyavshejsya klassifikacii na aprel 2021 goda vydelyayut ne menee 12 tipov arhej Crenarchaeota Garrityand Holt 2001 Krenarheoty termofily termoacidofily sernye anaerobnye bakterii Euryarchaeota Garrityand Holt 2001 Evriarheoty metanogennye i galofilnye arhei Thaumarchaeota Brochier Armanet et al 2008 v osnovnom okisliteli ammoniya kak naprimer morskoj ammonij okislitel i ammonij okislitel preimushestvenno pochvennogo proishozhdeniya Nedavnie 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